Variation in reproductive success in a fragmented Meadow Pipit population: a role for vegetation succession?

Abstract

Most farmland birds experience strong declines across Europe. These declines are typically associated with agricultural intensification but research on alternative local causes remains scarce. We investigated variation in reproductive success as a potential driver for the observed population declines in a fragmented population of the Meadow Pipit Anthus pratensis, a representative inhabitant of extensively managed mountain grasslands across Europe. Intense nest surveys in the entire Meadow Pipit metapopulation of the Northern Black Forest (SW Germany) between 2020 and 2022 provided information on reproductive success for 53 females distributed across nine habitat patches along an 18 km ridge of the Northern Black Forest. Hatching dates delayed by approx. 5.0 days per 100 m altitude and were almost 10 days later in a year with cold and rainy spring weather. Mean reproductive success per female and year (3.45 fledglings) was low compared to literature values (approx. 4.5) and may thus drive ongoing population declines. Mayfield nest survival estimates (approx. 51% across the nesting period) were comparably high, with most nest failures linked with predation or adverse weather. Low reproductive success further associated with comparably small clutch sizes and low fractions of second broods in habitat patches characterized by homogeneously dense swards. We suggest that restoration through extensive permanent cattle grazing coupled with succession control may be a key factor to increase population productivity.

Full text

Vol.:(0123456789)
1 3
Journal of Ornithology
https://doi.org/10.1007/s10336-023-02121-4
ORIGINAL ARTICLE
Variation inreproductive success inafragmented Meadow Pipit
population: arole forvegetation succession?
FabianAnger1 · MarcI.Förschler2 · NilsAnthes1
Received: 17 January 2023 / Revised: 29 September 2023 / Accepted: 10 October 2023
© The Author(s) 2023
Abstract
Most farmland birds experience strong declines across Europe. These declines are typically associated with agricultural
intensification but research on alternative local causes remains scarce. We investigated variation in reproductive success as
a potential driver for the observed population declines in a fragmented population of the Meadow Pipit Anthus pratensis,
a representative inhabitant of extensively managed mountain grasslands across Europe. Intense nest surveys in the entire
Meadow Pipit metapopulation of the Northern Black Forest (SW Germany) between 2020 and 2022 provided information
on reproductive success for 53 females distributed across nine habitat patches along an 18km ridge of the Northern Black
Forest. Hatching dates delayed by approx. 5.0days per 100m altitude and were almost 10days later in a year with cold and
rainy spring weather. Mean reproductive success per female and year (3.45 fledglings) was low compared to literature values
(approx. 4.5) and may thus drive ongoing population declines. Mayfield nest survival estimates (approx. 51% across the
nesting period) were comparably high, with most nest failures linked with predation or adverse weather. Low reproductive
success further associated with comparably small clutch sizes and low fractions of second broods in habitat patches char-
acterized by homogeneously dense swards. We suggest that restoration through extensive permanent cattle grazing coupled
with succession control may be a key factor to increase population productivity.
Keywords Meadow Pipit· Population declines· Nest survival· Nesting phenology· Conservation· Predation· Restoration
Zusammenfassung
Variation im Fortpflanzungserfolg einer fragmentierten Wiesenpieper-Population: Spielen Unterschiede in der
Vegetationsstruktur eine Rolle?
Die meisten Offenlandvögel in Europa zeigen in jüngster Zeit starke Bestandsrückgänge, die insbesondere auf die
Intensivierung der Landwirtschaft zurückgeführt werden. Untersuchungen zu alternativen Rückgangsursachen sind allerdings
selten. Wir untersuchten Unterschiede im Fortpflanzungserfolg als mögliche Ursache für den beobachteten Rückgang einer
fragmentierten Population des Wiesenpiepers Anthus pratensis, einer typischen Vogelart extensiv genutzter Grünlandflächen
in Europa. Eine intensive Nestersuche zwischen 2020 und 2022 lieferte Informationen über den Bruterfolg von 53 Weibchen in
Communicated by F. Bairlein.
* Fabian Anger
fabiananger@web.de
Marc I. Förschler
marc.foerschler@nlp.bwl.de
Nils Anthes
nils.anthes@uni-tuebingen.de
1 Institute ofEvolution andEcology, University ofTübingen,
Auf der Morgenstelle 28E, 72076Tübingen, Germany
2 Department ofEcosystem Monitoring, Research
andConservation, Black Forest National Park, Kniebisstr. 67,
72250Freudenstadt, Germany

Journal of Ornithology
1 3
neun inselartigen Offenland-Habitaten entlang eines 18km langen Höhenrückens im Nordschwarzwald. Der Schlupfzeitpunkt
verzögerte sich um etwa 5Tage pro 100 Höhenmeter und lag in einem Jahr mit kalter und regnerischer Witterung etwa
10Tage später. Der Fortpflanzungserfolg pro Weibchen und Jahr (3.45 flügge Jungvögel) war im Vergleich zu Literaturwerten
(ca. 4.5) relativ gering und könnte daher eine der Ursachen für den anhaltenden Rückgang der Population sein. Die nach der
Mayfield-Methode ermittelte Überlebensrate der Nester (ca. 51%) war vergleichsweise hoch, wobei die meisten Nestverluste
durch Prädation oder ungünstige Witterungsbedingungen verursacht wurden. Der geringe Fortpflanzungserfolg war zudem
durch relativ kleine Gelegegrößen und geringe Anteile an Zweitbruten in Teilflächen mit dichter und homogener Krautschicht
gekennzeichnet. Um den Fortpflanzungserfolg der Population zu erhöhen, halten wir auf Basis der Ergebnisse eine extensive
Beweidung mit Rindern sowie ein Zurückdrängen der Gehölze für besonders zielführend.
Introduction
Population declines are documented for many European bird
species (Keller etal. 2020; Burns etal. 2021) but are particu-
larly pronounced among the inhabitants of open agricultural
landscapes (Bauer etal. 2019; Kamp etal. 2020; PECBMS
2023). Farmland bird declines are typically attributed to the
pervasive agricultural intensification during recent decades
(Donald etal. 2006; Newton 2004). Yet, similar declines
also occur in mires, heathlands, coastal areas, and other
habitats that suffer far less, or more indirectly, from land
use intensification (Menke 2015; Förschler etal. 2016a).
Research on alternative local causes of population declines,
however, remains scarce.
In our study, we assessed associations between local land
use variables other than agricultural intensification and the
reproductive success of a representative inhabitant of exten-
sively managed moist grasslands, the ground nesting insec-
tivorous Meadow Pipit Anthus pratensis (Glutz von Blotz-
heim and Bauer 1985). Its European population declined by
approx. 63% between 1980 and 2021 (European Bird Census
Council 2022), the German population by approx. 60% in
just half that time interval between 1990 and 2009 (Gedeon
etal. 2014) with signs for stabilisation on a low level since
2010 (Kamp etal. 2020). In the south German federal state
of Baden-Württemberg, the Meadow Pipit rates as “criti-
cally endangered” (Kramer etal. 2022) given a population
decline from about 600 territories around 1995 (Hölzinger
and Ebenhöh 1999) to 120–160 territories in 2012–2016
(Kramer etal. 2022). The highly fragmented population
today concentrates in just three strongholds in the Southern
Black Forest, the Northern Black Forest, and at Lake Fed-
ersee (Gedeon etal. 2014), each isolated from their nearest
neighbouring population by at least 70km. Such small and
fragmented populations are meanwhile typical for several
farmland bird species within the intensively used agricul-
tural landscape of SW Germany (e.g., Anthes etal. 2017;
Seidt etal. 2017; Einstein etal. 2021).
In the Northern Black Forest, Meadow Pipits inhabit
raised bogs and extensively used grassland (heathland)
that is restricted by traditional land use to mountain tops
at 900 to 1200m a.s.l. (Förschler etal. 2016b). The local
population size declined from 85 territories in 1995–1997
to 28 territories in 2015 for still unknown causes (Förschler
etal. 2016a). Agricultural intensification, as discussed as the
primary factor for Meadow Pipit declines in general (e.g.,
Gedeon etal. 2014; Keller etal. 2020; BirdLife International
2021), does not qualify as a local factor in the absence of
agricultural intensification during the last decades. Yet, no
earlier study has investigated local breeding biology in detail
to better understand potential alternative drivers of popula-
tion declines.
We, therefore, focussed on reproductive success (= total
female productivity) as one key candidate cause for local
population declines, as suggested previously for other farm-
land bird species (Donald etal. 2002; Boatman etal. 2004;
Plard etal. 2020). From nest monitoring data of almost the
entire Meadow Pipit population between 2020 and 2022, we
first analysed variation in breeding phenology between years
and along the investigated altitudinal gradient as shown for
Meadow Pipits in Great Britain (Coulson 1956). Second, we
investigated the degree to which clutch size and reproduc-
tive success varied between brood types, grassland patches
and study years. Finally, we document reasons for nest fail-
ure and nestling mortality, quantify nest survival rates, and
derive suggestions for targeted conservation measures.
Methods
Study area
Our study took place in the “Grindenschwarzwald” in the
Northern Black Forest range (Fig.1, 48°32 N, 8°13 E,
Germany). “Grinden” are a regional type of semi-open
grassland (mountain heathland) that is embedded in
extensive coniferous forests. The “Grinden” heathlands
originated from deforestation or partial land clearance
by burning, followed by livestock grazing and litter use
between the sixteenth and nineteenth century (Förschler
etal. 2016b). They are part of the annex I habitat type
4030—European dry heaths and thus protected under the
NATURA 2000 Habitats Directive (Olmeda etal. 2020).
To preserve this habitat type with its highly diverse and

Journal of Ornithology
1 3
threatened flora and fauna, low-intensity grazing was
reintroduced on small grassland patches in 1995, but the
number of herds and area coverage continuously expanded
thereafter. Beyond the “Grinden”, grassland patches in
the study area are restricted to mowed or mulched ski
slopes. Surveys in the current study focused on all (seven)
grassland patches for which Meadow Pipit breeding was
confirmed or suggested during a recent large-scale sur-
vey (Förschler etal. 2016a) but also included another ten
grassland patches with historic breeding that had recently
been rated unoccupied (Fig.1). Grassland patches in the
nearby valleys were abandoned by Meadow Pipits sev-
eral decades ago (Förschler etal. 2016a) and thus not
investigated.
For statistical comparisons, we combined the individ-
ual grassland patches into three grassland patch groups:
Hornisgrinde North plus Hornisgrinde South into Hornis-
grinde, Schliffkopf plus Großer Geißkopf into Schliffkopf,
and all other grassland patches into “Others” given other-
wise small sample size (Fig.1).
Fig. 1 Locations and names
of the nine grassland patches
with Meadow Pipit breeding
occurrence (red) and grass-
land patches without breed-
ing occurrence (white) in the
Northern Black Forest close to
Baiersbronn during the study
period 2020–2022. Topographic
altitude: © LGL, www-lgl.de
(colour figure online)

Journal of Ornithology
1 3
Nesting survey
We visited each grassland patch about once per week during
the pre-breeding season, starting with the arrival of Meadow
Pipits in March, and the entire breeding season until early
August in three successive years 2020–2022. Pre-breeding
visits were necessary since Meadow Pipits exhibit intense
song displays during the first days after arrival but become
rather cryptic once pair bonds have been established (Süd-
beck etal. 2005) so that territories can easily be overlooked.
In the core breeding season between early May and August,
surveys focussed on nest building, incubating, and food pro-
visioning adult birds, spending at least half an hour per terri-
tory in that Meadow Pipit presence was confirmed during the
pre-breeding visits. From these weekly visits, we extracted
the number of breeding pairs (pairs with confirmed nest-
ing) and the number of solitary males (males with intense
and continuous singing activity without confirmed pairing
or nesting).
To localize nests, we pursued adult birds with nesting
material, when returning to nests during incubation, or
with nestling food items from typically > 60m distance to
minimize disturbance. Nest sites localized within approxi-
mately ± 4m were carefully approached and revisited about
every third day to document nest position, nest success
and the number of fledglings. After fledging or nest fail-
ure, we continued surveys to detect replacement or second
broods. Causes for nest failure were inferred from traces
in and around the nest, the presence and behaviour of the
adults, and weather conditions on the days preceding nest
failure. For each nest, we documented the presumed initial
clutch size (i.e., the largest number of eggs documented,
or the number of nestlings plus unhatched eggs), the num-
ber of fledglings (i.e., the number of nestlings during the
last visit before fledglings could be confirmed in close nest
vicinity), and assigned it—to our best knowledge—to first
brood, second brood or replacement brood based on date
and observation circumstances (first brood: nest of a pair
for that no earlier signs of breeding attempts were available;
second brood: nest of a pair for that a successful first brood
was confirmed; replacement brood: nest of a pair for that a
previous nest loss was documented or inferred from abrupt
termination of feeding or incubation activity).
Reproductive success
Our detailed surveys allowed us to closely approximate full
reproductive success as the total number of fledglings over
successive broods for each female and year. This was pos-
sible because most individuals of the breeding population
were individually colour-ringed with combinations of 3
colour rings (seven colours) and one metal ring. Our ring-
ing total of 32 adult birds caught from mist nets and 157
nestlings resulted in colour-ringed adult fractions of approx.
51% in 2020, 64% in 2021, and 62% in 2022. Since colour-
ringed females showed strong territory-fidelity with a given
male per season, un-ringed females were also assumed iden-
tical individuals for replacement or second broods in a given
male’s territory. Only in one case, a colour-ringed female
changed territory and thus its partner after losing its first
brood.
Unknown hatching dates were estimated based on nest-
ling size, nestling behaviour, and feather development by
comparison with local nestlings of known age and literature
reports assuming a total nestling period of 13days (Glutz
von Blotzheim and Bauer 1985; Hölzinger and Ebenhöh
1999).
We obtained reproductive success values (= fledgling
counts) per year for all 53 females that stayed in the breed-
ing area well into the breeding period. For 10 of these, val-
ues represent lower bounds to true fledgling count, either
because they rest on observations from a distance for one
brood (then as a minimum estimate of fledged young)
or because there is small chance, we missed a second or
replacement brood when a female could not be followed
into the late breeding period. We report findings based on
the full sample below, but provide the analysis reduced to
43 females with complete information in Online Resource
C.2, with near-identical qualitative (and even quantitative)
findings.
Statistical analyses
All statistical analyses were implemented in R version
4.2.2 (R Core Team 2022). For linear models, we used the
glmmTMB package (Brooks etal. 2017). The first model
describes variation in hatching dates of first broods (n = 40
nests, Gaussian error family) along the altitudinal gradient
and between study years, including their interaction. Further
models quantify variation in clutch size (n = 62 clutches) and
in reproductive success (i.e., fledgling counts per female,
n = 53 females) between patch groups, study years and
brood types (i.e., first broods versus replacement or second
broods), including the PatchGroup: BroodType and Year:
BroodType interactions. Both models used the generalized
poisson model family (‘genpois’ with a log-link) to reflect
underdispersion in their count responses. For model assess-
ment, we inspected residuals standardized for their distri-
bution family (independence of fitted values, homogeneity
across predictor variables) and conducted posterior predic-
tive checks on model-simulated data (dispersion, zero infla-
tion, and distribution relative to observed data) using the
routines provided by Santon etal. (2023). Mild zero inflation
in the fledgling model was captured by adding a zi-formula
that modelled extra zeroes by grassland patch groups.

Journal of Ornithology
1 3
We complement our descriptions of apparent nest
success and causes of nest failure with a formal analy-
sis of daily nest survival rates (Mayfield DSR) from a
binary logistic regression on nest outcome (0 = success,
1 = failure) as implemented in MARK (White and Burn-
ham 1999) and accessed through the R package RMark
(Laake and Rexstad 2008). As above, we described DSR
as a function of grassland patch group, study year and
brood type, and included season and nest age as covari-
ates because both often affect DSR (Rotella etal. 2004).
From overall mean DSR, we estimated nest survival prob-
ability as DSR raised to the power of the duration of incu-
bation and nestling stages (26days) (Johnson 1979).
We refrain from presenting P values and their associ-
ated evaluation of binary null hypotheses in accordance
with current recommendations for objective statistical
reporting (Halsey etal. 2015; Berner and Amrhein 2022).
Instead, we report effect size estimates with their compat-
ibility intervals, which are identical to classic confidence
intervals, but terminology shifts emphasis from trust in
hypothesis testing to a description of the central 95% den-
sity interval of effect values that are most compatible with
the observed data given the statistical model (Berner and
Amrhein 2022).
Results
Population size
Meadow Pipits were confirmed breeding in nine grassland
patches along an 18km section of the main ridge of the
Northern Black Forest (Fig.1). Seven of these grassland
patches are extensively grazed mountain heathlands, two
are mulched and mowed ski slopes (Table 1). Territory
numbers as well as the numbers of breeding females and
solitary males varied strikingly between study years despite
comparably intense survey effort. Grassland patch occu-
pancy dynamics included three recolonisation events and
one extinction event in marginal subpopulations (Table1).
Breeding phenology
Out of 62 documented nests (Table2), one was found dur-
ing nest building, 16 during incubation, and 45 during the
nestling stage. 44 nests were classified as first broods, 12 as
second broods and 6 as replacement broods.
Nestlings hatched between mid-May and end of July.
Lumped across study years, average hatching dates of first
broods increased with nest altitude by 4.98 (95% compat-
ibility interval, CI 2.33–7.63, Online Resource A.1) days
on average per 100m altitude, which implies hatching
Table 1 Number of Meadow
Pipit territories, breeding
females and solitary males in
the Northern Black Forest per
grassland patch (cf. Fig.1) in
2020–2022
Patch name Habitat Territories Breeding females Solitary males
2020 2021 2022 2020 2021 2022 2020 2021 2022
Mehliskopf Ski slope 0 1 0 0 1 0 0 0 0
Hochkopf Heathland 2 2 1 2 1 1 0 1 0
Unterstmatt Ski slope 0 0 1 0 0 1 0 0 0
Hornisgrinde N Heathland 2 2 2 1 1 2 1 1 0
Hornisgrinde S Heathland, raised bog 11 6 6 8 6 6 3 0 0
Schweinkopf Heathland 0 1 1 0 1 1 0 0 0
Großer Geißkopf Heathland 5 1 2 4 1 1 1 0 1
Schliffkopf Heathland 6 4 3 4 2 2 2 2 1
Zollstock-Heide Heathland 1 5 2 1 4 2 0 1 0
Total 27 22 18 20 17 16 7 5 2
Table 2 Numbers of females,
nests with complete information
on reproductive success,
percentage of females with
successful second broods, and
percentage of successful nests
per grassland patch group and
year
Females Nests % females with
successful 2nd
brood
% nests with apparent
success
2020 2021 2022 Total 20 21 22 Total 20 21 22 Total 20 21 22 Total
Hornisgrinde 9 7 8 24 13 6 9 28 57 0 60 37 92 100 89 93
Schliffkopf 8 3 3 14 9 2 4 15 14 0 0 9 78 100 50 73
Others 3 7 5 15 2 11 6 19 0 14 0 8 100 45 67 58
All areas 20 17 16 53 24 19 19 62 31 6 45 19 88 68 74 77

Journal of Ornithology
1 3
delays compared to the Zollstock-Heide site (950m a.s.l.)
by 3.5days at Schliffkopf (1020m a.s.l.) and 10.5days
at Hornisgrinde (1160m a.s.l.). Regression slopes var-
ied slightly between years but were consistently posi-
tive (Fig.2, Online Resource A.2). 2021 stands out with
particularly late first broods, with hatching dates approx.
10days later than in 2020 and 2022 at medium altitude
(Fig.2, Online Resource A.2).
Clutch size andhatching success
Most clutches contained four eggs, with an overall average
clutch size of 3.79 (CI 3.60–3.98). We could not detect any
relevant variation in mean clutch sizes between sites, years,
or brood types (Fig.3, Online Resource B). Overall hatching
rate was high, with 193 out of 222 non-predated eggs (87%)
hatching. 11 of these non-hatching eggs were contributed
by a single female from which all eggs of three successive
broods with the same male did not develop.
Reproductive success
Fledgling counts varied between zero and nine (Fig.4a,
b). Descriptive analysis revealed an overall average of
3.45 ± 2.36 (mean ± SD) fledglings per female and year, with
a particularly high value at Hornisgrinde (4.25 ± 2.47) com-
pared to Schliffkopf (2.64 ± 1.95) and Others (2.93 ± 2.25),
and a particularly low value in 2021 (2.71 ± 2.08) compared
to 2020 (4.10 ± 2.57) and 2022 (3.44 ± 2.25). We explored
possible reasons for these differences through a formal
analysis of the effects of year, brood type, and grassland
patch groups. Average reproductive success per female was
strikingly linked to brood type, where females with only a
single brood had clearly lower average reproductive success
than those with a replacement or second brood (Fig.4a).
Brood type effects did not vary among years (Fig.4a, Online
Resource C.1), so that the low average productivity in 2021
cannot be explained by low average nest success, but associ-
ated with a low fraction of females that initiated a second
brood in that year (Fig.4c). Average reproductive success of
Fig. 2 Relationship between altitude and hatching dates split by year,
including linear regression lines with their 95% compatibility inter-
vals (coloured shading, GLMM, N = 40 first broods). For statistical
details see Online Resource A.2 (colour figure online)
Fig. 3 Variation in clutch sizes of Meadow Pipits between years (left) and grassland patch groups (right) split by first vs. second or replacement
broods. Bold markers indicate predicted means and error bars their 95% compatibility intervals. For statistical details see Online Resource B

Journal of Ornithology
1 3
single-brooded females remained stable across patch groups
(Fig.4b, Online Resource C.1). In contrast, the benefit for
females with 2nd breeding attempts compared to single-
brooded females varied strikingly among patch groups
(Fig.4b, Online Resource C.1), with almost twice the repro-
ductive success at Hornisgrinde but far lower productivity
benefit in the other two patch groups (within-site contrasts
in Online Resource C.1). This difference goes along with
2nd breeding attempts at Hornisgrinde constituting largely
of true second broods (after completed first broods), while
those at the other patch groups largely relating to replace-
ment broods after first brood failure (Fig.4d).
Nestling mortality andnest survival
14 nests (23% of 62 nests) failed before fledging for vari-
able reasons between years (Table3). Six nest losses could
be associated with predation (nestlings or eggs depredated,
parents alive), three with adverse weather (nestlings dead
but without apparent damage in nest during cold and rainy
weather period, parents alive), and two with the loss of
a parent (nestlings dead in nest, only one parent present:
1 confirmed roadkill of female, 1 female disappeared for
unknown reason). In one case of depredation, the predator
was identified as a small carnivore (marten or weasel) from
bite marks on colour rings found in the nest. Beside carni-
vores, potential nest predators in the study area include birds
of prey, corvids, or European adders Vipera berus.
Fig. 4 Variation in the number of fledglings per female between years
(a) and grassland patch groups (b), and fractions of brood types for
years (c) and grassland patch groups (d). Bold markers indicate pre-
dicted means and error bars their 95% compatibility intervals. For
statistical details see Online Resource C
Table 3 Causes of nest losses per study year
2020 2021 2022 Sum
Predation 0 1 5 6
Adverse weather 1 2 0 3
Loss of parent 2 0 0 2
Unfertilized eggs 0 3 0 3
Total 3 6 5 14

Journal of Ornithology
1 3
One nest out of two found on a ski slope was rescued
from destruction by mowing through a targeted late mowing
arrangement. Another likely loss was prevented by fencing
the nest before intense sheep grazing. By contrast, none of
five nests recorded in low intensity grazing cattle pastures
was damaged by livestock trampling.
Overall daily nest survival was 0.9744 ± 0.0067 SE
(95% CI 0.9572–0.9848), resulting in a mean probability of
0.510 (95% CI 0.321–0.672) for nests to survive the entire
26-day nesting period (13days incubation, 13days nestling
period). Variation in daily survival rate was best explained
by a grassland patch group model, with highest mean DSR
at Hornisgrinde (Fig.5a, Online Resource D). Nest age was
the only other predictor that came close in predictor power
to the intercept-only model (Online Resource D), possibly
indicating a modest increase in DSR with nest age (Fig.5b).
Discussion
We studied nesting ecology and reproductive success in a
continuously declining, fragmented grassland population of
Meadow Pipits in the Northern Black Forest in 2020–2022.
Hatching dates of first broods were earlier in lower altitudes
and varied between study years. Clutch sizes showed low
variation and associated with neither year, patch group nor
brood type. Reproductive success per female varied strik-
ingly between grassland patch groups and was largely driven
by nest survival and the fraction of females that raised sec-
ond broods.
The observed increase in hatching dates with alti-
tude matches previous findings but was more pronounced
compared to Britain populations where average hatching
dates increased by only 2.5days per 100m altitude (Coul-
son 1956). Local habitat conditions may have intensified the
altitudinal effect in our study area: our highest altitude patch
group, Hornisgrinde, is characterized by a raised bog, where
cool and moist microclimate may favour particularly late
hatching. Hatching date differences between years correlate
well with weather conditions, where particularly late hatch-
ing in 2021 was associated with a cold spring and snowfall
until May. Mean air temperatures in May were 10.9°C and
13.3°C in 2020 and 2022, but only 7.9°C in 2021 at a
nearby weather station at 800m a.s.l. (Wetterdienst 2022).
The documented average clutch size of 3.79 eggs per nest
was low compared to literature values that vary between 3.89
and 5.4, depending on study region (Davies 1958; Constant
and Eybert 1980; Hötker and Sudfeldt 1982; Rose 1982;
Hölzinger and Ebenhöh 1999; Malm etal. 2020). However,
clutch size has also been described to increase with latitude
(Hötker and Sudfeldt 1982), and the Northern Black For-
est is close to the southern range margin of Meadow Pipit
(Keller etal. 2020). Average clutch sizes in more southern
populations were still slightly higher, with 4.15 in the South-
ern Black Forest during 1981–1996 (Hölzinger and Ebenhöh
1999) and 4.40 in the Swiss Jura during 1972–1974 (Pedroli
1978). Given a lack of previous data from the Northern
Black Forest we cannot assess, however, whether current
average clutch sizes are lower compared to the mid-twentieth
century when local Meadow Pipit populations were consid-
ered stable.
Our reproductive success data are difficult to compare to
literature values since reproductive success obviously dif-
fers between years, and demography as well as life history
Fig. 5 Estimated Mayfield daily survival rates for Meadow Pipit nests
and their variation among grassland patch groups (left, showing the
global mean DSR as dashed line) and with nest age spanning the
26-day period of incubation (13days) and nestling feeding (13days)
(right), including 95% compatibility intervals. See Online Resource D
for model comparisons

Journal of Ornithology
1 3
strategies may differ strongly between populations (Barras
etal. 2021). Yet, it is striking that average reproductive suc-
cess per female and year in our study area was about one
fledgling below that reported from Lower Saxony (Northern
Germany) with 4.45 raised juveniles per year and female
over a 5-year period (Hötker and Sudfeldt 1982). The authors
hint at a large fraction of second and even third broods and
estimate 2.3 broods or breeding attempts per female and year
(Hötker and Sudfeldt 1982), which compares to only 1.4
breeding attempts per female and year in our study. This
indicates that low reproductive success in the Black Forest,
which goes along a lack of (successful) second and third
broods, is insufficient to maintain population stability.
Differences in reproductive success between years might
be explained by the cold climatic conditions in 2021, where
delayed first broods and a shortage of second broods directly
contributed to the low average reproductive success com-
pared to 2020 and 2022. Adverse weather conditions thus not
only directly cause nest failures, but also indirectly reduce
reproductive success through a reduction in successive brood
numbers (e.g., Frey 1989; Förschler etal. 2005). Such effects
may intensify given that weather (and rainfall) extremes are
predicted to become more frequent (e.g., Seneviratne etal.
2012; Zeder and Fischer 2020).
Apparent nest success (77%) was higher than that
reported for four different years in Scotland (range:
18–65%, Malm etal. 2020) and for a population in Poland
(71%, Halupka 1998a). Also, our overall Mayfield esti-
mate for nest survival of 50.9% (49.6% when adjusted to
a 27-day period) was similar to the 48.2% (27-day period)
reported from a population in Poland (Halupka 1998a).
While these findings may suggest nest losses as a minor
reason for low productivity in the Northern Black Forest
population, we also found a striking link between nest sur-
vival and the high reproductive success per female at the
Hornisgrinde patch group, which grounded in a combina-
tion of generally higher nest survival and—to some extent
as a consequence of higher nest survival of first broods—a
larger fraction of females that raise second broods. This
finding is indeed opposite to the expectation of larger frac-
tions of second broods at lower altitudes where an earlier
onset of breeding prolongs the reproductive period (Bears
etal. 2009). We propose these differences in productivity
to be linked to differences in habitat structure and thus suit-
ability between grassland patch groups. First, a Meadow
Pipit population in Poland showed higher nest survival at
moist and hidden microrelief structures Halupka (1998b).
Consistent with this finding, the Hornisgrinde raised bog
provides highly structured microrelief coupled with short
and sparse vegetation and close-cropped grass areas that
provide well-protected nest sites and the required acces-
sible foraging habitat (Hölzinger and Ebenhöh 1999; Van-
denberghe etal. 2009) throughout the breeding season.
This contrasts to the other two patch groups, which are
characterised by comparably little microrelief and rather
dense and homogenous ground vegetation cover that is less
penetrable for foraging, in particular in the late breeding
season. Second, earlier work found that edge effects can
affect nest survival (Vetter etal. 2013). The Hornisgrinde
patch group contains the largest grassland patch in the
study area, thus showing the smallest possible edge effects
among all our study sites.
Conservation implications
Based on our findings above, we propose to expand low-
intensity permanent cattle grazing to break up dense ground
vegetation during the late breeding period (Bunzel-Drüke
etal. 2019) and thus help to create more suitable structures
like well-protected nest sites, accessible foraging habitat, and
thus ultimately for successful first and second broods. Such
predictable, permanent, and low intensity grazing regimes
are not expected to result in relevant nest losses from tram-
pling (Beintema and Muskens 1987, own data), contrary to
rotational or paddock grazing systems, intense sheep graz-
ing, or mowing that all go along with substantial nest losses
in Meadow Pipit (own data, Pavel 2004) and other ground
nesting birds (e.g., Handschuh and Klamm 2022). Since
nestlings fledge by mid-August and require another approx.
2weeks until showing full escape flights, intense grazing or
mowing should start no earlier than late August (Glutz von
Blotzheim and Bauer 1985). Management measures should
further aim at restoring open grassland habitats lost to shrub
succession. Besides space for more territories, larger grass-
land patches might also increase reproductive success due
to more possibilities for foraging and nesting and reduced
edge effects.
Supplementary Information The online version contains supplemen-
tary material available at https:// doi. org/ 10. 1007/ s10336- 023- 02121-4.
Acknowledgements We thank Johannes Kamp for constructive com-
ments on earlier versions of the manuscript. We acknowledge support
from the Open Access PublishingFund of the University of Tübingen.
Author contributions FA: conceptualization (lead); data curation
(lead); investigation (lead); methodology (lead); project administration
(lead); formal analysis (equal); software (support); writing—original
draft preparation (lead); funding acquisition (lead). MIF: conceptu-
alization (support); investigation (support), project administration
(support); resources (lead); writing—review & editing (support). NA:
conceptualization (support); formal analysis (equal); software (lead);
methodology (support); writing—review & editing (lead).
Funding Open Access funding enabled and organized by Projekt
DEAL. FA was temporarily funded by the Landesgraduiertenförderung
Baden-Württemberg, Germany.

Journal of Ornithology
1 3
Data availability All data generated or analyzed during this study are
included in the supplementary information files.
Declarations
Conflict of interest The authors declare no conflict of interest.
Ethical approval Field work in this study was conducted under permits
for the years 2020–2022 issued by the Black Forest National Park and
the regional conservation authorities (Regierungspräsidium Freiburg
and Karlsruhe, Az 84-8675.12 and Az 55-8841.03; 8853.17). Meadow
Pipits in the study area are accustomed to human presence and animal
welfare had priority. Nest visits were made as short as possible and
none of the parents deserted after nest visits.
Open Access This article is licensed under a Creative Commons Attri-
bution 4.0 International License, which permits use, sharing, adapta-
tion, distribution and reproduction in any medium or format, as long
as you give appropriate credit to the original author(s) and the source,
provide a link to the Creative Commons licence, and indicate if changes
were made. The images or other third party material in this article are
included in the article’s Creative Commons licence, unless indicated
otherwise in a credit line to the material. If material is not included in
the article’s Creative Commons licence and your intended use is not
permitted by statutory regulation or exceeds the permitted use, you will
need to obtain permission directly from the copyright holder. To view a
copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.
References
Anthes N, Boschert M, Daniels-Trautner J (2017) Verbreitung und
Bestandsentwicklung der Grauammer Emberiza calandra in
Baden-Württemberg. Ornithol Jh Bad-Württ 33:27–44
Barras AG, Blache S, Schaub M, Arlettaz R (2021) Variation in demog-
raphy and life-history strategies across the range of a declining
mountain bird species. Front Ecol Evol 9:780706. https:// doi. org/
10. 3389/ fevo. 2021. 780706
Bauer HG, Heine G, Schmitz D, Segelbacher G, Werner S (2019)
Starke Bestandsveränderungen der Brutvogelwelt des Boden-
seegebietes – Ergebnisse aus vier flächendeckenden Brutvo-
gelkartierungen in drei Jahrzehnten. Vogelwelt 139:3–29
Bears H, Martin K, White GC (2009) Breeding in high-elevation habi-
tat results in shift to slower life-history strategy within a single
species. J Anim Ecol 78:365–375. https:// doi. org/ 10. 1111/j. 1365-
2656. 2008. 01491.x
Beintema AJ, Muskens GJDM (1987) Nesting success of birds breed-
ing in Dutch agricultural grasslands. J Appl Ecol 24:743–758.
https:// doi. org/ 10. 2307/ 24039 78
Berner D, Amrhein V (2022) Why and how we should join the shift
from significance testing to estimation. J Evol Biol 35:777–787.
https:// doi. org/ 10. 1111/ jeb. 14009
Bird Life International (2021) Anthus pratensis. The IUCN red list of
threatened species 2021: e.T22718556A154480081. https:// doi.
org/ 10. 2305/ IUCN. UK. 2021-3. RLTS. T22718 556 A154480081.
en. Accessed 20 Dec 2022
Boatman ND, Brickle NW, Hart JD, Milsom TP, Morris AJ, Murray
AWA, Robertson PA (2004) Evidence for the indirect effects of
pesticides on farmland birds. Ibis 146:131–143. https:// doi. org/
10. 1111/j. 1474- 919X. 2004. 00347.x
Brooks ME, Kristensen K, van Benthem KJ, Magnusson A, Berg CW,
Nielsen A etal (2017) glmmTMB balances speed and flexibil-
ity among packages for zero-inflated generalized linear mixed
modeling. R Journal 9(2):378–400. https:// doi. org/ 10. 3929/ ethz-
b- 00024 0890
Bunzel-Drüke M, Reisinger E, Böhm C, Buse J, Dalbeck L, Ellwanger
G etal (2019) Naturnahe Beweidung und NATURA 2000, 2nd
edn. Arbeitsgemeinschaft Biologischer Umweltschutz, Bad
Sassendorf
Burns F, Eaton MA, Burfield IJ, Klvaňová A, Šilarová E, Staneva A,
Gregory RD (2021) Abundance decline in the avifauna of the
European Union reveals cross-continental similarities in bio-
diversity change. Ecol Evol 11(23):16647–16660. https:// doi.
org/ 10. 1002/ ece3. 8282
Constant P, Eybert MC (1980) Données sur la biologie de la repro-
duction du pipit farlouse, Anthus pratensis L., dans les landes
bretonnes. Oiseaux 35:349–360
Coulson JC (1956) Mortality and egg production of the Meadow Pipit
with special reference to altitude. Bird Study 3:119–132
Davies SJJF (1958) The breeding of the Meadow Pipit in Swedish
Lapland. Bird Study 5:184–191. https:// doi. org/ 10. 1080/ 00063
65580 94759 19
Donald PF, Evans AD, Muirhead LB, Buckingham DL, Kirby WB,
Schmitt SIA (2002) Survival rates, causes of failure and produc-
tivity of Skylark Alauda arvensis nests in lowland farmland. Ibis
144:652–664. https:// doi. org/ 10. 1046/j. 1474- 919X. 2002. 00101.x
Donald PF, Sanderson FJ, Burfield IJ, van Bommel FPJ (2006) Further
evidence of continentwide impacts of agricultural intensification
on European farmland birds, 1990–2000. Agric Ecosyst Environ
116:189–196. https:// doi. org/ 10. 1016/j. agee. 2006. 02. 007
Einstein J, Harry I, Kramer M (2021) Bestandsentwicklung und Ver-
breitung des Braunkehlchens (Saxicola rubetra) in Baden-Würt-
temberg seit 1950. Ornithol Jh Bad-Württ 37:7–19
European Bird Census Council (2022) Pan-European common bird
monitoring scheme. https:// pecbms. info/ trends- and- indic ators/
speci es- trends/ speci es/ anthus- prate nsis/. Accessed 3 Nov 2023
Förschler MI, Borras A, Cabrera J, Cabrera T, Senar JC (2005) Inter-
locality variation in reproductive success of the Citril finch Seri-
nus citrinella. J Ornithol 146:137–140. https:// doi. org/ 10. 1007/
s10336- 005- 0072-y
Förschler MI, Anger F, del Val E, Aichele D, Dreiser C (2016a) Zur
aktuellen und historischen Bestandssituation des Wiesenpiepers
Anthus pratensis im Nordschwarzwald. Ornithol Jh Bad-Württ
32:45–51
Förschler MI, Richter C, Gamio T (2016b) Grinden – waldfreie
Bergheiden im Nationalpark Schwarzwald. NaturschutzInfo
2(2016):28–31
Frey M (1989) Brutbiologie des Hänflings Carduelis cannabina unter
den Einflüssen des Gebirgsklimas. Ornithol Beob 86:265–289
Gedeon K, Grüneberg C, Mitschke A, Sudfeldt C, Eickhorst W, Fischer
S etal (2014) Atlas Deutscher Brutvogelarten. Stiftung Vogel-
monitoring und Dachverband Deutscher Avifaunisten, Münster
Glutz von Blotzheim UN, Bauer KM (1985) Handbuch der Vögel
Mitteleuropas. Passeriformes (1 Teil), 2nd edn. AULA-Verlag,
Wiesbaden
Halsey LG, Curran-Everett D, Vowler SL, Drummond GB (2015)
The fickle P value generates irreproducible results. Nat Methods
12:179–185. https:// doi. org/ 10. 1038/ nmeth. 3288
Halupka K (1998a) Nest predation in Meadow Pipits Anthus pratensis
nesting in natural conditions. Ornis Fenn 75:139–143
Halupka K (1998b) Nest-site selection and nest predation in meadow
pipits. Folia Zool 47:29–37
Handschuh M, Klamm A (2022) Nationally important population of the
Corn Bunting Emberiza calandra in the Hainich National Park.
Anzeiger Des Vereins Thüringer Ornithologen 10:43–78
Hölzinger J, Ebenhöh H (1999) Anthus pratensis (Linnaeus, 1758)
Wiesenpieper. Seite 146–155 in: Hölzinger, J. (Herausgeber):
Die Vögel Baden-Württembergs. Band 3.1, Singvögel 1. Ulmer,
Stuttgart. Jochen Hölzinger

Journal of Ornithology
1 3
Hötker H, Sudfeldt C (1982) Untersuchungen zur Brutbiologie des
Wiesenpiepers (Anthus pratensis). J Ornithol 123:183–201.
https:// doi. org/ 10. 1007/ BF016 45057
Johnson DH (1979) Estimating nest success: the Mayfield method and
an alternative. Auk 96:651–661. https:// doi. org/ 10. 1093/ auk/ 96.4.
651
Kamp J, Frank C, Trautmann S, Busch M, Dröschmeister R, Flade M
etal (2020) Population trends of common breeding birds in Ger-
many 1990–2018. J Ornithol 162:1–15. https:// doi. org/ 10. 1007/
s10336- 020- 01830-4
Keller V, Herrando S, Voříšek P, Franch M, Kipson M, Milanesi P etal
(2020) European breeding bird atlas 2: distribution, abundance
and change. European Bird Census Council & Lynx Edicions,
Barcelona
Kramer M, Bauer HG, Bindrich F, Einstein J, Mahler U (2022) Red
List of Breeding Birds of Baden-Württemberg. 7th edition, as of
31 December 2019. – Naturschutz-Praxis Artenschutz 11. LUBW
Landesanstalt für Umwelt Baden-Württemberg, Karlsruhe. www.
lubw. baden- wuert tembe rg. de
Laake J, Rexstad E (2008) RMark – an alternative approach to building
linear models in MARK. http:// www. phidot. org/ softw are/ mark/
docs/ book/ pdf/ app_3. pdf. Accessed 11 Aug 2023
Malm LE, Pearce-Higgins JW, Littlewood NA, Karley AJ, Karasze-
wska E, Jaques R etal (2020) Livestock grazing impacts compo-
nents of the breeding productivity of a common upland insectivo-
rous passerine: results from a long-term experiment. J Appl Ecol
57:1514–1523. https:// doi. org/ 10. 1111/ 1365- 2664. 13647
Menke W (2015) Die Entwicklung des Brutvogelbestands im Elis-
abeth-Außengroden (Gemeinde Wangerland, Kreis Friesland).
Nachrichten Des Marschenrates 52:57–72
Newton I (2004) The recent declines of farmland bird populations in
Britain: an appraisal of causal factors and conservation actions.
Ibis 146:579–600. https:// doi. org/ 10. 1111/j. 1474- 919X. 2004.
00375.x
Olmeda C, Šefferová V, Underwood E, Millan L, Gil T, Naumann
S (2020) EU Action plan to maintain and restore to favourable
conservation status the habitat type 4030 European dry heaths.
European Commission, pp 1–116
Pavel V (2004) The impact of grazing animals on nesting success of
grassland passerines in farmland and natural habitats: a field
experiment. Folia Zool 53:171–178
PECBMS (2023) Common farmland birds indicator. https:// pecbms.
info/ trends- and- indic ators/ indic ators/ indic ators/E_ C_ Fa/. Last
Accessed 10 Jan 2023
Pedroli JC (1978) Breeding success of the Meadow Pipit Anthus prat-
ensis in the Swiss Jura. Ornis Scand 9:168–171. https:// doi. org/
10. 2307/ 36758 78
Plard F, Bruns HA, Cimiotti DV, Helmecke A, Hötker H, Jeromin H
etal (2020) Low productivity and unsuitable management drive
the decline of central European lapwing populations. Anim Con-
serv 23:286–296. https:// doi. org/ 10. 1111/ acv. 12540
Rose LN (1982) Breeding ecology of British pipits and their cuckoo
parasite. Bird Study 29:27–40. https:// doi. org/ 10. 1080/ 00063
65820 94767 35
Rotella JJ, Dinsmore SJ, Shaffer TL (2004) Modelling nest-survival
data: a comparison of recently developed methods that can be
implemented in MARK and SAS. Anim Biodivers Conserv
27:187–205
Santon M, Korner-Nievergelt F, Michiels NK, Anthes N (2023) A
versatile workflow for linear modelling in R. Front Ecol Evol
11:1065273. https:// doi. org/ 10. 3389/ fevo. 2023. 10652 73
Seidt M, Geißler-Strobel S, Kramer M, Kratzer R, Straub F, Anthes
N (2017) Bestandsentwicklung und Grundlagen für den Schutz
des Rebhuhns Perdix perdix im Landkreis Tübingen. Ornithol Jh
Bad-Württ 33:3–12
Seneviratne SI, Nicholls N, Easterling D, Goodess CM, Kanae S,
Kossin J, Luo Y, Marengo J, McInnes K, Rahimi M, Reichstein
M, Sorteberg A, Vera C, Zhang X (2012) Changes in climate
extremes and their impacts on the natural physical environment.
In: Field CB, Barros V, Stocke TF etal (eds) Managing the risks
of extreme events and disasters to advance climate change adapta-
tion. Cambridge University Press, Cambridge and New York, pp
109–230. https:// doi. org/ 10. 7916/ d8- 6nbt- s431
Südbeck P, Andretzke H, Fischer S, Gedeon K, Schikore T, Schröder
K, Sudfeldt C (2005) Methodenstandards zur Erfassung der Brut-
vögel Deutschlands. Selbstverlag, Radolfzell
Vandenberghe C, Prior G, Littlewood NA, Brooker R, Pakeman R
(2009) Influence of livestock grazing on meadow pipit foraging
behaviour in upland grassland. Basic Appl Ecol 10:662–670.
https:// doi. org/ 10. 1016/j. baae. 2009. 03. 009
Vetter D, Rücker G, Storch I (2013) A meta-analysis of tropical forest
edge effects on bird nest predation risk: edge effects in avian nest
predation. Biol Conserv 159:382–395. https:// doi. org/ 10. 1016/j.
biocon. 2012. 12. 023
Wetterdienst D (2022) Klimadaten Deutschland, Station Freudenstadt.
https:// www. dwd. de/ DE/ leist ung en/ klima daten deuts chland/ klima
daten deuts chland. html. Accessed 20 Dec 2022
White GC, Burnham KP (1999) Program MARK: survival estimation
from populations of marked animals. Bird Study 46:120–139.
https:// doi. org/ 10. 1080/ 00063 65990 94772 39
Zeder J, Fischer EM (2020) Observed extreme precipitation trends and
scaling in Central Europe. Weather Clim Extremes 29:100266.
https:// doi. org/ 10. 1016/j. wace. 2020. 100266
Publisher's Note Springer Nature remains neutral with regard to
jurisdictional claims in published maps and institutional affiliations.