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Survival of Resident Neotropical Birds: Considerations for Sampling and Analysis

Based on 20 Years of Bird-Banding Efforts in Mexico

Article in The Auk · July 2012

DOI: 10.1525/auk.2012.11171

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 The Auk 129(3):500−509, 2012
 The American Ornithologists’ Union, 2012.
Printed in USA.

SURVIVAL OF RESIDENT NEOTROPICAL BIRDS: CONSIDERATIONS FOR

SAMPLING AND ANALYSIS BASED ON 20 YEARS OF BIRD-BANDING
EFFORTS IN MEXICO
VIVIANA RUIZ-GUTIÉRREZ,1,5,6 PAUL F. DOHERTY, JR.,1 EDUARDO SANTANA C.,2,3
SARAHY CONTRERAS MARTÍNEZ,2 JORGE SCHONDUBE,4 HERIBERTO VERDUGO MUNGUÍA,2
AND EDUARDO IÑIGO-ELIAS5
1
Department of Fish, Wildlife and Conservation Biology, Colorado State University, Fort Collins, Colorado, USA;
2
Instituto Manantlán de Ecología y Conservación de la Biodiversidad-DERN, Universidad de Guadalajara, Jalisco, Mexico;
3
Department of Forest and Wildlife Ecology, University of Wisconsin-Madison, Madison, Wisconsin, USA;
4
Centro de Investigaciones Ecosistemas, Universidad Nacional Autónoma de México, Morelia, Michoacán, Mexico; and
5
Cornell Laboratory of Ornithology, Cornell University, Ithaca, New York, USA

A.—Long-term bird-banding programs provide knowledge of the demographic rates of avian populations, but we currently
lack information on demographic rates of most bird populations in species-rich tropical ecosystems. Banding in tropical regions is typically
conducted with protracted or irregular sampling occasions that make the choice of the proper capture–mark–recapture (CMR) model
difficult. Here, we address common challenges related to collecting and analyzing data to estimate survival rates of resident Neotropical
birds using  years of banding efforts in Mexico as a case study. We applied Cormack-Jolly-Seber and Barker models to estimate apparent
survival and recapture probabilities of species with sufficient data for survival analyses. We were able to analyze  resident species of
 total species; apparent survival probabilities ranged from . to ., and recapture probabilities from . to .. For monitoring
programs with existing data collected at continuous, uneven, or irregular intervals, we recommend the application of the Barker model over
other models because it was more efficient in the use of available banding data and less often violated CMR assumptions. We recommend
that monitoring programs last > years and provide additional protocol suggestions for primary and secondary sampling occasions, as well
as the number of nets, potential net configurations, and the extent of the spatial scale. These baseline recommendations are likely to foster
an increase in our knowledge of avian survival rates in tropical ecosystems, which is imperative for managing tropical bird populations
under changing environmental conditions. Received  August , accepted  April .

Key words: Barker model, mark–recapture statistics, mist netting, population modeling, tropical birds.

Supervivencia de Aves Residentes Neotropicales: Consideraciones para el Muestreo y Análisis con base en un
Esfuerzo de  Años de Anillamiento de Aves en México

R.—Los programas de anillamiento de aves a largo plazo brindan conocimiento sobre las tasas demográficas de las poblaciones
de aves, pero todavía hace falta información sobre las tasas demográficas de la mayoría de poblaciones de aves en ecosistemas tropicales
áltamente diversos. El anillamiento en las regiones tropicales es típicamente conducido con muestreos prolongados o irregulares, que
hacen difícil la escogencia del modelo apropiado de captura-marcado-recaptura (CMR). En este trabajo abordamos las dificultades más
comunes relacionadas con la recolección y análisis de datos para estimar las tasas de supervivencia de aves neotropicales residentes, usando
 años de esfuerzos de anillamiento en México como un caso de estudio. Aplicamos los modelos de Cormack-Jolly-Seber y de Barker para
estimar la supervivencia aparente y las probabilidades de recaptura de especies con datos suficientes para los análisis de supervivencia.
Pudimos analizar seis especies residentes de un total de ; las probabilidades aparentes de supervivencia estuvieron entre . y ., y las
probabilidades de recaptura entre . y .. Para los programas de monitoreo con datos recolectados en intervalos continuos, desiguales
o irregulares, recomendamos la aplicación del modelo de Barker por encima de otros modelos, ya que éste fue más eficiente en el uso de los
datos de anillamiento disponibles, y viola menos frecuentemente los supuestos del CMR. Recomendamos que los programas de monitoreo
duren más de  años y presentamos sugerencias adicionales para el protocolo de ocasiones de anillamiento primario y secundario, así
como del número de redes, de la configuración potencial de las redes y de la extensión de la escala espacial. Estas recomendaciones base

6
E-mail: vruizg@colostate.edu

The Auk, Vol. , Number , pages −. ISSN -, electronic ISSN -.   by The American Ornithologists’ Union. All rights reserved. Please direct all
requests for permission to photocopy or reproduce article content through the University of California Press’s Rights and Permissions website, http://www.ucpressjournals.
com/reprintInfo.asp. DOI: ./auk..

— 500 —
JULY 2012 — PROTOCOL TO ESTIMATE SURVIVAL FOR RESIDENT TROPICAL BIRDS — 501

podrían promover el incremento de nuestro conocimiento de las tasas de supervivencia de las aves en ecosistemas tropicales, lo que es
imperativo para la administración de las poblaciones de aves tropicales bajo condiciones ambientales cambiantes.

E   management of avian populations
requires baseline information on the relationship between demo-
graphic rates and environmental factors. This knowledge is fun-
damental to our understanding of how changing environmental
conditions will influence population vital rates (e.g., survival and
reproductive success) and is critical for predicting the viability and
likelihood of persistence of threatened populations. Several long-
term banding programs have been established to monitor bird
populations in Europe (e.g., Baillie ) and North America (e.g.,
DeSante et al. ), generating information on demographic rates
of many resident and migratory bird populations. Despite increas-
ing efforts, the geographic extent of most information generated by
bird-banding monitoring programs remains limited to the Nearc-
tic and Palaearctic regions, which support only ~% of the world’s
avian biodiversity (Newton and Dale ). Roughly % of the
world’s bird species reside in tropical ecosystems whose forests (e.g.,
lowland, moist, dry, subtropical, and montane) contain almost half
of the world’s threatened bird species (BirdLife International ).
Thus, we are faced with the challenge of preserving tropical forest
birds without a clear understanding of the changes in demographic
rates that drive population declines of threatened species. More im-
portantly, we lack information to develop and prioritize bird con-
servation objectives (e.g., minimize adult mortality and maximize
dispersal) and the means to determine whether management ac-
tions have succeeded in achieving the objectives themselves.
Most published research on bird population dynamics in tropi-
cal ecosystems has been conducted in the Neotropics, where % of
the world’s bird species reside. These studies generally focus on es-
timating survival rates, similar to monitoring programs established
in Europe and North America. Monitoring trends in survival is crit-
ical, especially for forest understory, resident Neotropical birds (i.e.,
understory forest species that do not seasonally migrate over long
distances, hereafter “resident Neotropical birds”), because some of
these species are long lived (Robinson et al. ) and we need ad-
ditional information about the relative importance of survival and
fecundity across latitudes, life histories, and environments (Martin
, ; Ricklefs and Wikelski ) to target conservation ef-
forts (Pfister , Clark and Martin ).
In the past two decades, advancements in capture–mark–
recapture (CMR) sampling methods and statistical techniques
(Williams et al. , Sandercock ) have been used to esti-
mate annual survival probabilities for ~% of resident Neotropi-
cal bird species (Table ), but these are restricted to a few sites and
leave % of species unstudied. Survival information is lacking for
Neotropical bird populations because data do not exist for many
species and, when data exist, they are communicated through in-
ternal project reports and not in peer-reviewed literature. Second,
the main objective of many banding projects is to gather data on
Neotropical migrants (DeSante et al. ), with less focus on res-
ident species. Third, data collection methods might present some
analysis and interpretation challenges. For example, current pro-
tocols for resident and migratory bird species suggest continuous,
monthly sampling events (e.g., Ralph et al. , Latta et al. ).
This may result in pooling of data across space and time, which
may violate the common assumptions in many CMR models of
demographic and geographic closure during sampling.
Here, we address several of the challenges typically faced when
collecting and analyzing data to estimate survival rates of resident
Neotropical birds while adhering to CMR assumptions. We focus
our analyses on the assumption of closure within a sampling pe-
riod and forgo discussing the details of “why” and “what” to moni-
tor, given that these topics have been described at length elsewhere
(Yoccoz et al. ). Instead, we assume that “what to monitor” has
been identified as survival and deal specifically with “how to moni-
tor.” We use  years of banding efforts from the Sierra de Man-
antlán Bird Monitoring and Conservation Project, which is part of

TABLE 1. A compilation of banding studies that used a capture–mark–recapture framework to estimate survival probabilities of understory resident
bird species in tropical ecosystems in Latin America: publication, location and years of data collection, sampling intervals with number of days per
primary sampling event, months sampled, number of sites sampled, and number of species for which survival and recapture probabilities were re-
ported (NI = not included).

Sites Species
Reference Location Years Intervals Months (n) (n)

Karr et al. 1990 Panama a 1977–1986 3 days (primary)/month March, July 8 25

Brawn et al. 1995 Panama a 1977–1986 3 days (primary)/month March, July 8 25
Faaborg and Arendt 1995 Puerto Rico 1973–1990 3 days (primary)/year January or February 1 9
Johnston et al. 1997 Trinidad 1986–1995 NI June–August 3 17
Francis et al. 1999 Peru 1973–1982 3–4 days/year NI 6 14
Brawn et al. 1999 Panama a 1977–1997 3 days (primary)/month March, July 8–12 11
March–April,
Jullien and Clobert 2000 French Guiana 1991–1994 4 days (primary)/2 months September–October 7 17
Blake and Loiselle 2002 Costa Rica 1985–1994 2 days (primary)/5–6 weeks December–October 3 3
Parker et al. 2006 Ecuador 1998–2004 3 days (primary)/year December–February 2 5
Pearce-Higgins et al. 2007 Bolivia 1994–1999 2 weeks (primary) July–September 2 1
Blake and Loiselle 2008 Ecuador 2001–2006 1 day (primary) January, March 16 31
Ruiz-Gutierrez et al. 2008 Costa Rica 1993–2006 2–4 days (primary)/year January–March 8 1
Hau et al. 2010 Panama a 1979–2006 3 days (primary)/month March, July 8 NI
Present study Mexico 1993–2010 2–3 days (primary)/month January–December 1 9
a
The four published works for Panama are for the same study.
502 — RUIZ-GUTIÉRREZ
Mexico’s Long-term Ecological Research program, as a case study
(Santana , Guallar et al. ). We use subsets of available
banding data to represent common variations in CMR analysis and
evaluate the potential of the Barker model to estimate apparent an-
nual survival probabilities using data collected continuously or at
irregular intervals. Further, we use our results to provide sugges-
tions for protocols that () incorporate sampling approaches found
in the published literature, () meet CMR assumptions, () correct
for sampling bias, and () consider the ecology and life histories of
resident Neotropical birds. By providing a broader range of CMR
models to analyze existing bird-banding data, as well as suggestions
concerning sampling protocols that might lead to more powerful
data collection and analysis, we hope to improve the ability of inves-
tigators to estimate survival of Neotropical and other birds.
M ETHODS
Study site and sampling.—The study was conducted at the ,-ha
Las Joyas Research Station located within the Sierra de Manantlán
Biosphere Reserve and managed by the Manantlán Institute of Ecol-
ogy and Conservation of Biodiversity. Our analyses were restricted
to the San Campus (SC) banding site, one of seven at the station,
because SC has been actively monitored since  (Santana ).
The habitat surrounding SC is early-successional montane pine
(Pinus spp.) and oak (Quercus spp.) forest (Cuevas and Jardel ).
Sampling occurred in SC for  to  consecutive days. From  to
, sampling occurred once or twice a year; from  to ,
sampling occurred nearly monthly; from  to , sampling
occurred from November through March. Sampling consisted of
constant-effort mist netting, with nets placed at the same location
in each site throughout the duration of the study. Birds were sam-
pled using  to  mist nets (-mm mesh) that were opened for 
to  consecutive hours, starting at dawn. Nets were distributed over
an area of approximately  to . ha. Nets were checked every –
min, and all individuals were identified to species and banded with
a uniquely numbered aluminum band.
Methods for estimating survival.—To estimate survival, we
used three methods (Table  and Table ). Methods  and  are
Cormack-Jolly-Seber (CJS) models (Cormack , Jolly , Se-
ber ), and method  is a model developed by Barker ().
Method  came from the realization that assuming both a closed
population and constant site fidelity across all sampled months
(e.g., January–December), spanning both breeding and non-
breeding seasons, was not a likely biological scenario for resident
Neotropical birds, or for many bird species in general. To correctly
estimate survival, the length of the sampling occasion should also
only be a small fraction of the interval over which we want to make
inferences (Williams et al. ). To meet CMR assumptions, we
generated capture histories that collapsed data collected only in
January and February, from  to . Data were not collected
in either of those months from  to , and so we had to dis-
card data from those years.
Method  examined the potential applicability of an alternate
approach to CJS developed by Barker (). In a CJS context, uneven
intervals and continuous sampling violate the closure assumption
and can bias estimates of apparent survival and recapture probabili-
ties. When sampling occurs “instantaneously” (e.g., – consecutive
days) but at multiple times during a year, we can correct for potential
bias by defining each interval as a fraction of the year (e.g., Blake and
ET AL.— AUK, VOL. 129
Loiselle ) or by defining relatively smaller intervals (e.g., days and
months). But when different months are sampled inconsistently or
sampling is not instantaneous (e.g., sampling occurs once per month,
as recommended by Ralph et al. []), correcting for potential bias
can be difficult. The Barker () model can correct for potential
bias under these sampling scenarios by allowing the consideration
of a “primary sampling occasion” for which we can assume a closed
population, define intervals (e.g., yearly) between primary occasions,
but also include recaptures collected outside of this primary period
as ancillary information. Although the Barker model was intended
to mainly incorporate information from dead recoveries and resight-
ings outside of the primary sampling occasion, the model has been
successfully applied to live-capture data by setting the dead-recov-
ery probability to zero (Collins and Doherty , LeDee et al. ).
For method , we generated capture histories from banding data col-
lected from  to , defining January–February as the primary
sampling occasion, and used recaptures from March–December as
ancillary information.
For method , we generated capture histories from data col-
lected in January and February from  years of sampling (–
). After , sampling outside of the primary occasion
(January–February) was minimal or nonexistent, and there were
not enough recaptures outside of the primary occasion to yield
robust estimates using the Barker model from –. We in-
clude this method because it probably represents our best survival
estimates given the long period.
Statistical analysis for survival.—We fit a set of models that
varied in their assumptions of apparent survival (φ) and recapture
probabilities (p) in Program MARK (White and Burnham ).
Apparent survival is the probability that an individual alive at time
t is also alive and has not emigrated from the study area at time
t + ; the product of true survival and fidelity (F), where F is defined
as the probability that an individual that was alive and present in
the study area in time t is also alive and has not emigrated from
the study area at time t + . We considered time-dependent (t) and
constant (.) models for both survival and recapture probabilities.
We also used a time-since-marking (TSM) model to estimate ap-
parent survival (Pradel et al. ) to correct for the potential un-
derestimation of these parameters as a result of transient and/
or young individuals (Johnston et al. ). The TSM model is a
version of the CJS model and allows the partitioning of apparent
survival into survival during the first year after banding (φ) and
survival from subsequent years (φ+), commonly referred to as sur-
vival of resident or adult birds. Time-since-marking models have
been applied to estimate apparent survival and recapture prob-
abilities of tropical birds and are recommended for analyses of
tropical bird data (Johnston et al. , Brawn et al. , Parker
TABLE 2. The types of survival models, months, and years of data used in
methods 1–3 of the present study, and numbers of species (n) for which
we were able to estimate survival and recapture probabilities.
Method Survival model Months Years n
1 CJS a January–February 1994–1999 1
2 Barker b January–February (primary) 1994–1999 4
March–December (auxiliary) 1994–1999
3 CJS January–February 1994–2010 6
a
CJS = Cormack-Jolly-Seber model (Cormack 1964, Jolly 1965, Seber 1965).
b
Barker = the model developed by Barker (1997).
JULY 2012 — PROTOCOL TO ESTIMATE SURVIVAL FOR RESIDENT TROPICAL BIRDS — 503

TABLE 3. Notation and interpretation of the candidate model set tested using methods 1–3.

Model notation Interpretation

Methods 1 and 3 (Cormack-Jolly-Seber)

φ1(.)φ2+(.) p(.) Constant survival during the first year (φ1), survival during subsequent years (φ2+),
and recapture (p)
φ1(.)φ2+(t) p(t) Constant survival during the first year (φ1), time variation in recapture (p), and
time variation in survival for subsequent years (φ2+)
φ(.) p(.) Constant survival (φ) and recapture (p)
φ(t) p(.) Time variation in survival (φ) and constant recapture (p)
φ(t) p(t) Time variation in survival (φ) and recapture (p)
Method 2 (Barker)
All models have recovery rate (r) fixed to zero, with constant probabilities of be-
ing alive and having been recaptured (R), dying and having been recaptured alive
(R′), fidelity (F), and return probability of temporary emigrants (F′)
r = 0,S1(.)S2+(.)p(.)R(.)R’(.)F(.)F’(.) Constant survival after the first year (S1) and survival during subsequent years (S2+)
r = 0,S(.)p(.)R(.)R’(.)F(.)F’(.) Constant survival (S) and recapture probability (p)
r = 0,S(t)p(.)R(.)R’(.)F(.)F’(.) Time variation in survival (S) and constant recapture probability (p)
r = 0,S(t)p(t)R(.)R’(.)F(.)F’(.) Time variation in survival (S) and recapture probability (p)

et al. , Sandercock ). We had only enough data to model
the first-year survival as constant (φ[.]), but considered both con-
stant (φ+[.]) and time dependence [φ+[t]) on survival and recap-
ture probabilities of birds considered to be residents during the
sampling event, for methods  and  (Table ).
The Barker model (method ) in Program MARK has seven
estimable parameters: St = survival (probability that an individual
that was alive at time t is also alive at time t + ); pt = recapture
(probability that an individual that is alive in the study area at time
t is recaptured); rt = recovery rate (probability that an individual
dies between time t and t +  and was found dead and reported);
Rt = probability that an individual that survived from time t and
t +  was recaptured and alive between t and t + ; Rt ’ = probability
that an individual that died between t and t +  was recaptured and
alive between t and t + ; Ft = fidelity probability (probability of re-
turning to the study area given that the individual survives); and
Ft ’ = return of temporary emigrants (probability that an individual
that has emigrated temporarily from the study area and was not
recaptured during time t is alive and returns to the study area at
t + ). We set r = . for all models because the data did not contain
band recoveries from dead birds (e.g., Collins and Doherty ).
We cannot make any inferences on fidelity and other related pro-
cesses (F, F’, R, and R’) because all recaptures for the secondary oc-
casions were from the same area as the study site. Therefore, these
parameters were all set to be individually constant (.) in the candi-
date model set (Table ). Because of our sampling scheme and the
inherent inability to estimate fidelity, we interpret what is typi-
cally defined as true survival (S) in the Barker model as apparent
survival. Similarly, we interpret recapture probability as the prob-
ability that the individual (conditional on its being alive) remains
in the study area and is encountered. We modeled constancy and
time dependence on survival and recapture probabilities, survival
during the first year after banding (φ), and survival from subse-
quent years (φ+) of the TSM model (Table ).
We constructed our models for each species independently.
We evaluated overdispersion by generating a variance inflation
factor (ĉ) using the “median c-hat” estimation capabilities within
Program MARK for our most parameterized models. When ĉ > ,
we used the quasi-likelihood modified Akaike’s information crite-
rion corrected for small sample sizes (QAICc) for model selection
and model averaging (Burnham and Anderson ). We present
model-averaged estimates (Burnham and Anderson ).
R ESULTS
The -year data set from the SC study site constitutes one of the
largest known data sets in the Neotropics: , total captures,
of which , are recaptures of , individuals belonging to
 species. However, only  species had > total captures and
> individuals, with most species averaging <. recaptures in-
dividual– (Fig. ). Using the subset of data from  to  re-
sulted in model convergence on  of the  species. Of these, we
had only enough marked individuals and recaptures for survival
analyses for  migratory and  resident species using CMR meth-
ods (e.g., SE range: –) for method . Of the six resident species,
two belong to the family Trochilidae (Colibri thalassinus [COTH],
Hylocharis leucotis [HYLE]), two are in Turdidae (Myadestes
occidentalis [MYOC] and Catharus occidentalis [CAOC]), one
in Traupidae (Diglossa baritula [DIBA]), and one in Emberizidae
(Atlapetes pileatus [ATPI]). The models with the most support
across species had recapture probabilities as constant, p(.), and ap-
parent survival as constant, φ(.) or S(.), or had a constant effect of
transient or juvenile individuals on apparent survival, φ(.)φ+(.) or
S(.)S+(.) (Table ).
Using method , we had only enough data for DIBA to model
apparent survival and recapture probabilities. Apparent survival
was φ+ = . ± . (SE), and recapture probability was p = .
± . (Table ). Using the Barker model (method ), we were able
to generate estimates for four species: HYLE, COTH, DIBA, and
ATPI (φ+ = .–.; p = .–.; Table ). Using method ,
we obtained survival estimates for MYOC, CAOC, and the other
species analyzed with methods  and : COTH, HYLE, DIBA, and
ATPI (φ+ = .–.; p = .–.; Table ).
A subset of the data that included only captures from January
and February better met the biological and closure model assump-
tions (method ). However, this allowed us to obtain estimates for
504 — RUIZ-GUTIÉRREZ ET AL.— AUK, VOL. 129

from methods  and  (Table ). All estimates of apparent survival

and recapture probabilities using method  overlapped with es-
timates obtained through method , which represent  years of
monitoring efforts (Table ).

D ISCUSSION

Matching data collection with available analysis methods prior

FIG. 1. Summary of banding data collected at the San Campus site, 1991–
2010. Histograms represent total captures, total number of individuals
captured, total recaptures, and the ratio of average number of recaptures
per individual, estimated at the species level.
only  of  species (DIBA; Table ). The inclusion of captures
from March–December as supplementary information (method )
allowed us to estimate parameters for DIBA and three additional
species. The cost of adding the additional parameters of the Barker
model was evident from estimates of recapture probabilities,
which had very large standard errors compared with estimates
to sampling is the most efficient approach to avoid potential pit-
falls when making statistical inferences in CMR studies (Lindberg
). In Latin America, published banding protocols for moni-
toring survival and other demographic rates for Neotropical res-
ident birds are often based on continuous sampling throughout
the year (Ralph et al. , Latta et al. ). Challenges common
to many banding programs, like logistical constraints and a lack
of consistency in funding across years, could also result in data
being collected over protracted or uneven sampling periods. The
result is that a large proportion of existing tropical bird-banding
data likely represent banding events spread over multiple months,
uneven spacing and sampling effort across years, or both. We fo-
cused on  years of banding effort in the Las Joyas Research Sta-
tion in the Sierra de Manantlán Biosphere Reserve, Mexico, as a
case study to contrast different approaches for analyzing existing
bird-banding data to estimate survival probabilities of resident
Neotropical birds.
Our results suggest that the Barker model (method ) can
successfully be applied to such existing banding data for some
species, while still adhering to the closure and interval-length
assumptions required by CMR analyses. The Barker model also
enables the combination of primary banding data, with auxiliary
information from other sites that were not sampled as consis-
tently. The proportion of species for which we can estimate sur-
vival using the Barker model remains minimal (%), even after

TABLE 4. Species code, scientific name, total number of individuals (n), method used (1–3), best-supported (top) models, model
averaged estimates of apparent survival of resident adult birds (S2+ or φ2+) and recapture probabilities (p), with their respective
unconditional standard error (SE) and lower (LCI) and upper (UCI) bounds of the confidence interval.

Code Species n Method Top models S2+ or φ2+ SE LCI UCI p SE LCI UCI

DIBA Diglossa baritula 291 1 φ(.) p(.) 0.49 0.23 0.14 0.85 0.40 0.23 0.09 0.81
φ1(.)φ2+(.)p(.)
103 2 S1(.)S2+(.)p(.) 0.77 0.13 0.43 0.94 0.44 0.19 0.15 0.78
S(.)p(.)
50 3 φ1(.)φ2+(.) p(.) 0.68 0.09 0.48 0.83 0.30 0.10 0.14 0.53
φ(.) p(.)
HYLE Hylocharis leucotis 74 2 S1(.)S2+(.)p(.) 0.52 0.22 0.16 0.86 0.24 0.21 0.03 0.75
193 3 φ1(.)φ2+(.) p(.) 0.52 0.12 0.29 0.74 0.36 0.16 0.13 0.68
MYOC Myadestes occidentalis 152 3 φ(.) p(.) 0.65 0.09 0.46 0.80 0.17 0.07 0.07 0.36
φ1(.)φ2+(.) p(.)
ATPI Atlapetes pileatus 58 2 S(.)p(.) 0.62 0.15 0.32 0.85 0.52 0.31 0.09 0.92
φ1(.)φ2+(.) p(.)
48 3 φ1(.)φ2+(.) p(.) 0.74 0.08 0.56 0.87 0.44 0.10 0.27 0.62
φ(.) p(.)
CAOC Catharus occidentalis 260 3 φ1(.)φ2+(.) p(.) 0.68 0.04 0.59 0.75 0.36 0.05 0.26 0.46
COTH Colibri thalassinus 71 2 S(.)p(.) 0.30 0.17 0.08 0.67 0.21 0.24 0.01 0.83
S(t) p(.)
121 3 φ(.) p(.) 0.43 0.17 0.16 0.75 0.11 0.13 0.01 0.61
φ1(.)φ2+(.) p(.)
JULY 2012 — PROTOCOL TO ESTIMATE SURVIVAL
 years of continuous monitoring (– in our study). How-
ever, this rate was not unlike many published studies that gener-
ated data for < species, even after as much as  years of data
collection (Table ).
The Barker model remains preferable over truncating data to
create a sampling interval closer to  year (e.g., method ), which
greatly reduced the data and the number of species for which we
could estimate annual survival probabilities. Collapsing band-
ing data across several months to increase the number of species
for which we can estimate survival (e.g., Blake and Loiselle )
yields estimates that are difficult to interpret as annual survival
rates, even if effort is constant across years (e.g., McGregor et al.
) because of the violation of the closure assumption within
a sampling occasion. Small sample sizes also create the tempta-
tion to combine capture histories across species to provide a value
of “average” survival for a group of interest (e.g., across guilds or
habitat types). The biological justification for combining capture
histories of diverse tropical bird species is tenuous because of the
implicit assumption that all individuals across species have the
same survival and recapture probabilities, including rates of fidel-
ity, and temporary and permanent immigration–emigration. The
resulting survival estimates are a weighted average, and the high-
est weight will be associated with species with the most individu-
als and recaptures.
Below, we build on the work of Ralph et al. () and Latta
et al. () and present protocols that are specifically designed
to model survival as a discrete process using a CMR framework
(Table ), explicitly linking data collection to analysis (Lindberg et
al. ) using existing user-friendly software. Our suggested pro-
tocols also incorporate published methods used to estimate sur-
vival of Neotropical birds (Table ). Ralph et al. () developed
protocols to accomplish multiple objectives of monitoring of bird
populations, but Latta et al. () focused specifically on mon-
itoring Neotropical resident landbirds and wintering migrants.
The continuous sampling methods of Ralph et al. () and Latta
et al. () have successfully yielded critical information on molt
patterns, food habits, and temporal changes in body condition
(e.g., Santana , Schondube et al. , Guallar et al. ). Al-
though analytical methods to model survival and recapture prob-
abilities as a continuous process exist, continuous sampling (e.g.,
monthly) is likely to result in net avoidance behaviors, leading to
low recapture rates (see below). Discrete-time survival models are
also most commonly used and have received the most attention
TABLE 5. Summary of suggestions for sampling Neotropical resident birds to estimate apparent an-
nual survival. Each column represents a sampling topic, sampling considerations under each topic,
and our suggestions for sampling to increase sample size and recapture probabilities in relation to
our study.
Sampling topic Considerations
Time span Average survival
Trends in survival
Sampling approach Frequency and timing
Spatial extent Core netting area
Fidelity and movements
Spatial configuration Continuous sites
Discrete sites
a
Robust design stands for the model described by W. L. Kendall (1997).
FOR RESIDENT TROPICAL BIRDS — 505
from methods developers, and user-friendly software (e.g., Pro-
gram MARK) is available to conduct such discrete-time analyses.
Time span of monitoring.—The appropriate time frame over
which to estimate survival is likely to differ between temperate
and tropical forest species, given that the latter are considered to
be longer lived, on average (see review in Robinson et al. ).
We recommend against monitoring adult survival over short time
frames (e.g.,  month; Githiru and Lens ) in relation to the
longevity of the species (> years), because true survival rates
over short intervals would be ~. and would not be expected to
differ from one interval to the next. Therefore, observed differ-
ences in apparent monthly survival estimates are likely a result
of movement patterns and do not reflect differences in true sur-
vival (e.g., Cruz-Angón et al. ). For long-term monitoring
programs, DeSante et al. () recommended using ≥ years of
data to model temporal and spatial variation in annual survival
for bird species in North America. We suggest that a longer time
frame (more than – years) might be required to study popu-
lation dynamics of most resident bird species in Neotropical sys-
tems, given the longevity and low recapture probabilities (p < .;
Sandercock and Beissinger ) that are likely. We also note that
models that estimate population growth explicitly may be useful
with such long-term (approximately – years) data sets (De-
Sante et al. ).
Frequency and timing of primary sampling events.—Ralph et
al. () suggest operating each site for  day every  days from
June to August for temperate species and monitoring longer than
 continuous months in the Neotropics. Latta et al. () pro-
posed similar recommendations specifically for monitoring de-
mographic rates of tropical bird populations and suggested that
sampling continuously throughout the entire year at regular in-
tervals (e.g., daily, weekly, monthly, quarterly) was more produc-
tive than sampling once a year. Sampling month to month, over
multiple years, is known to cause long-term net-avoidance behav-
ior in tropical lowland birds (J. D. Brawn pers. comm.) and prob-
ably resulted in lower recapture probabilities for the entire length
of our study. To reduce this potential effect, we recommend moni-
toring during discrete primary sampling events. The timing and
number of the primary sampling events should be determined by
the specific objectives of the monitoring project and would need to
be optimized with simulations, but the beginning, middle, or end
of the peak of the breeding season for most species (e.g., March,
May, and July in Central America) may be a logical starting point.
Suggestions
~ 10 years
~ 10–30 years
Robust design a
Primary and secondary occasions
>25 ha
>1 km surrounding core
Clusters of nets
Scaling of clusters by habitat types or sizes
One focal site, surrounding ancilliary data
506 — RUIZ-GUTIÉRREZ
Examples include Faaborg and Arendt (), whose sampling
took place every January from  to the present; Brawn et al.
(), with sampling in March and July from  to the pres-
ent; and Blake and Loiselle (), with sampling in January and
March from  to the present (Table ). Reducing continuous
monitoring to one or a few primary sampling periods each year
could reduce the cost and effort associated with current protocols
and is likely to reduce the effects of net avoidance by individuals.
Secondary sampling occasions.—We recommend using a
robust-design (RD) sampling approach (Kendall et al. ), in
which sampling events are separated into primary and secondary
occasions. We have had success with primary sampling occasions
that are made up of  consecutive days of mist netting or secondary
occasions (e.g., Fabborg and Arendt , Brawn et al. , Ruiz-
Gutiérrez et al. ). The main objective of the RD analysis ap-
proach is that it allows for the additional estimation of temporary
emigration parameters. The probability of emigrating temporar-
ily is likely to be high for frugivorous birds (e.g., tanagers), insec-
tivores (e.g., flycatchers), and nectarivores (e.g., hummingbirds),
as well as differing between sexes and between breeding and non-
breeding seasons. High or variable rates of temporary emigration
will result in recapture probabilities that are biased low and possi-
bly underestimate survival rates (Sandercock and Beissinger ,
Parker et al. ). Although recapture rates will likely not be high
enough to estimate temporary emigration parameters for most
species, a benefit of the RD sampling approach is that banding in-
formation can be collapsed across secondary sampling occasions
(e.g., to generate one value for the entire primary sampling event),
without violating CMR closure assumptions (e.g., most work in
Table ). Although we used an RD sampling approach in our study,
we had consecutive primary sampling occasions (monthly) and
used very few mist nets (–) within a small area (< ha), which
resulted in too few recaptures and marked individuals to apply RD
models. However, the RD sampling approach allowed us to col-
lapse our capture histories across secondary occasions and apply
CJS and Barker models to our data. The RD can also be extended
to use ancillary recaptures between primary sessions (see Lind-
berg ).
Spatial extent of sampling.—The small spatial extent of our
sampling area (< ha) also could have contributed to our low sam-
ple sizes and low recapture rates. We recommend that the spatial
extent of study areas be large enough to maximize the number of
potential territories per site (e.g., > ha). Otherwise, permanent
emigration rates can underestimate dispersal distances and esti-
mates of apparent survival for bird populations (Cilimburg et al.
, Zimmermann et al. , Cooper et al. ). Banded birds
are a combination of hatch-year birds, nonbreeding and nonterri-
torial floaters, delayed dispersers, and resident breeders. Through
the application of TSM models, we can account for lower survival
of hatch-year birds and for some of the nonbreeding floaters that
never returned to the study area after the initial capture. How-
ever, Francis et al. () found that  of  species of tropical
birds in Peru were recaptured > m away after the year of initial
capture, with movement distances ranging from  m to > km.
This is likely the result of protracted breeding seasons (Schon-
dube et al. ), delayed dispersal, and individuals that disperse,
float, and fail to breed (Tarwater and Brawn ). Time-since-
marking models do not account for recaptured individuals that
ET AL.— AUK, VOL. 129
permanently emigrate or switch territories > year after initial
capture (e.g., widowed individuals). Although differences in fi-
delity between northern temperate and tropical bird species
have never been formally tested, the evidence suggests that the
common assumption that fidelity to a study site is ~ for tropi-
cal birds is also dependent on the spatial extent of sampling. Our
application of the Barker model did not allow us to estimate fi-
delity and, hence, true survival because we did not have any re-
captures beyond our small study area that would have enabled us
to model permanent, large-scale movements. To estimate differ-
ences in fidelity among tropical bird species and test for effects of
permanent emigration on apparent survival rates, we recommend
increasing the spatial extent of the effective study area. One way
in which this can be accomplished is by collecting ancillary move-
ment data, such as telemetry data (Powell et al. ) or resight-
ings of marked individuals (Cilimburg et al. ) at a reasonable
distance outside the study area (> km).
Spatial configuration of study sites.—Species-rich tropical
ecosystems generally contain low densities of individual terri-
tories for most bird species (Karr et al. ). Although the area
used by individuals can expand and contract between breeding
and non-breeding seasons, the number of individuals per species
that were consistently recaptured in our relatively small netting
sites was low. To increase sample sizes and recaptures, we recom-
mend that several clusters or net lanes (“N” in Fig. ) be used to
monitor continuous sites using the RD sampling approach. Moni-
toring can be consecutive, sampling one cluster per each second-
ary sampling occasion (e.g., Jullien and Clobert ; Fig. A), or
a subset of available nets per cluster can be opened at each occa-
sion per site (e.g., Johnston et al. ; Fig. B). Another possibil-
ity is to have a core set of net clusters or lanes that are monitored
on a regular basis, and other clusters or lanes outside of the core
FIG. 2. Examples of potential spatial configurations for monitoring sur-
vival of resident Neotropical birds. For continuous sites, (A) net lanes
or clusters (“N”) are sampled consecutively (N1–N3) at each sampling
occasion or (B) a subset of each net cluster–lane (all N1) is sampled si-
multaneously. For discrete sites or habitat types, (C) individuals can be
captured and marked in focal areas (“N”) and recaptures or resightings
(“R”) can take place outside the focal site. Sampling effort must also be
scaled across (D) habitat types and sizes.
JULY 2012 — PROTOCOL TO ESTIMATE SURVIVAL
(“R” in Fig. ) that are run less frequently but might generate valu-
able recaptures (e.g., Francis et al. ; Fig. C). To monitor mul-
tiple discrete sites (e.g., forest patches) or sites located in different
habitat types (e.g., shade vs. sun coffee), sampling designs should
take into consideration that, as with detection probabilities (Ruiz-
Gutiérrez et al. ), recapture probabilities are likely to differ
between species in a group and could contrast sharply between
habitat types (e.g., primary forest vs. pasture) and patch sizes. In
this case, sampling effort (e.g., number of clusters) could be scaled
across habitat types and patch sizes (Fig. D), and the focus should
be on increasing recapture probabilities across all site categories.
Individuals could also be captured and marked in focal habitats
or patches, and recaptures or resightings can take place outside of
the focal sites (e.g., surrounding matrix habitats; Fig. C) and po-
tentially incorporated in the analysis using Lindberg et al.’s ()
and Barker’s models (Barker and White ).
Number of nets.—Our relatively small number of nets (–
mist nets) could also have contributed to our low sample sizes and
recapture probabilities. The number of nets should be based on
the number and configuration of net clusters or lanes, limited by
the total number of birds that can be handled and processed safely.
Ralph et al. () recommend using  to  nets site–, but we ob-
served a range of – nets cluster– in the published literature
(cited in Table ) and have also had success with clusters of ~
nets. The density of the nets will vary depending on the latitude,
altitude, topography, and vegetation of the forest sites, but we rec-
ommend as even spacing as possible to reduce heterogeneity.
Overall, the estimation of annual survival through CMR
sampling and analysis methods for most Neotropical bird species
will require a substantial amount of long-term effort. We believe
that our low sample sizes, even after  years of monitoring, likely
resulted from a combination of net avoidance behavior related to
our continuous sampling, the small number of nets we used, and
the limited spatial extent of our sampling area. The sampling and
analysis protocols presented here are meant as suggestions for
long-term sampling and monitoring and are designed to increase
the potential number of species for which we can obtain survival
rates. To evaluate the performance of our suggestions, existing
banding data should be frequently analyzed and, if needed, the
protocols should be adjusted on the basis of preliminary findings
or results from simulations. To this end, we need trained quan-
titative ecologists working for, and collaborating with, agencies
and institutions in charge of monitoring resident Neotropical bird
populations. Much of our current knowledge on survival stems
from scientists who are trained in CMR analysis techniques, have
access to such training, or collaborate with individuals who are
well versed in the subject (Table ). Our knowledge of survival
rates of resident Neotropical birds will not likely increase un-
less existing and future international bird-monitoring programs
shift from a blanket approach of teaching the mechanics of how
to capture and band birds, toward need-based, local capacity-
building efforts focused on teaching the data analysis and inter-
pretation methodology that we have applied and discussed here.
All the above recommendations, along with other complementary
ones aimed at establishing long-term monitoring projects in Latin
America (Villaseñor and Santana ), will increase the amount
of knowledge of survival rates, providing insight into the popula-
tion ecology of resident Neotropical birds, as well as help guide
future conservation and management actions.
FOR RESIDENT TROPICAL BIRDS — 507
ACKNOWLEDGMENTS
Data analysis was funded by the Cornell Laboratory of Ornithol-
ogy (CLO); the MacArthur Foundation (E.I.E. and V.R.G.); the
Department of Fish, Wildlife and Conservation Biology (V.R.G.
and P.F.D.); and the Colorado AGEP Postdoctoral Fellowship Pro-
gram (V.R.G.). The field work was funded by the Universidad de
Guadalajara, CONACYT, CONANP, Point Reyes Bird Observa-
tory, Rocky Mountain Bird Observatory, CLO, Catalán Institute
of Ornithology, University of Wisconsin-Madison, University of
Arizona-Tucson, San Francisco State University, World Wildlife
Fund, National Fish and Wildlife Foundation, Audubon Society
of Greater Denver, U.S. Fish and Wildlife Service, U.S.-A.I.D.,
U.S. Forest Service, National Institutes of Health MIRT program,
Grace J. Calder Trust, General Services Foundation, Colorado
Wildlife Heritage Foundation, Vilas Scholarship, WWF-Interna-
tional Prince Bernhard Fellowship, and the Paul and Bay Founda-
tions’ Biodiversity Leadership Award. Comments by J. M. Lepak,
M. Lindberg, J. D. Brawn, and C. M. Francis on the manuscript
were very much appreciated. The Sierra de Manantlán long-term
banding project was possible thanks to > individuals who
helped in multiple technical and logistical chores, too many to list;
we acknowledge and are grateful for their support.
LITERATURE CITED
B, S. R. . The contribution of ringing to the conservation
and management of bird populations: A review. Ardea :–
.
B, R. J. . Joint modeling of live-recapture, tag-resight, and
tag-recovery data. Biometrics :–.
B, R. J.,  G. C. W. . Joint analysis of live and dead
encounters of marked animals. Pages – in Proceedings
of the Second International Wildlife Management Congress (R.
Field, R. J. Warren, H. Okarma, and P. R. Sievert, Eds.). Wildlife
Society, Bethesda, Maryland.
BL I. . Threatened birds occur in all hab-
itats, but the majority are found in forest. [Online.] Available from
www.birdlife.org/datazone/sowb/casestudy/.
B, J. G.,  B. A. L. . Manakins (Pipridae) in
second-growth and old-growth forests: Patterns of habitat use,
movement, and survival. Auk :–.
B, J. G.,  B. A. L. . Estimates of apparent sur-
vival rates for forest birds in eastern Ecuador. Biotropica :–
.
B, J. D., J. R. K,  J. D. N. . Demography of
birds in a Neotropical forest: Effects of allometry, taxonomy, and
ecology. Ecology :–.
B, J. D., J. R. K, J. D. N,  W. D. R. .
Demography of forest birds in Panama: How do transients affect
estimates of survival? Pages – in Proceedings nd Inter-
national Ornithological Congress (N. J. Adams and R. H. Slotow,
Eds.). Bird Life South Africa, Johannesburg.
B, K. P.,  D. R. A. . Model Selection
and Multimodel Inference: A Practical Information-Theoretic
Approach, nd ed. Springer-Verlag, New York.
C, A. B., M. S. L, J. J. T,  S. J.
H. . Effects of dispersal on survival probability of adult
Yellow Warblers (Dendroica petechia). Auk :–.
508 — RUIZ-GUTIÉRREZ
C, M. E.,  T. E. M. . Modeling tradeoffs in avian
life history traits and consequences for population growth. Eco-
logical Modelling :–.
C, C. T.,  P. F. D, J. . Survival estimates for
Royal Terns in southern California. Journal of Field Ornithology
:–.
C, C. B., S. J. D,  J. R. W. . Can we
improve estimates of juvenile dispersal distance and survival?
Ecology :–.
C, R. M. . Estimates of survival from the sighting of
marked animals. Biometrika :–.
C-A, A., T. S. S,  R. G. . An
experimental study of habitat selection by birds in a coffee planta-
tion. Ecology :–.
C, G. R.,  E. J. J. . Flora y Vegetación de la
Estación Científica Las Joyas. Departamento de Ecología y
Recursos Naturales-IMECBIO, Centro Universitario de la
Costa Sur, Universidad de Guadalajara, Jalisco, Mexico.
DS, D. F., M. P. N,  D. R. O’G. . Identify-
ing the proximate demographic cause(s) of population change by
modeling spatial variation in productivity, survivorship, and pop-
ulation trends. Ardea :–.
DS, D. F., T. S. S, R. B. S, J. F. S, C. A.
R  V A, S. M, A. C, D. R.
K, M. G,  B. M. . MoSI (Moni-
toreo de Sobrevivencia Invernal): Assessing habitat-specific over-
wintering survival of Neotropical migratory landbirds. Pages
– in Bird Conservation Implementation and Integration
in the Americas (C. J. Ralph and T. D. Rich, Eds.). U.S. Depart-
ment of Agriculture, Forest Service General Technical Report
PSW-GTR-.
F, J.,  W. J. A. . Survival rates of Puerto Rican
birds: Are islands really that different? Auk :–.
F, C. M., J. S. T,  J. W. F. .
Survival rates of understory forest birds in Peru. Pages –
in Proceedings nd International Ornithological Congress
(N. J. Adams and R. H. Slotow, Eds.). BirdLife South Africa,
Johannesburg.
G, M.,  L. L. . Demography of an Afrotropical
passerine in a highly fragmented landscape. Animal Conserva-
tion :–.
G, S., E. S C., S. C, H. V, 
A. G. . Paseriformes del occidente de México: Mor-
fometría, datación y sexado. Monografía no. , Museu de Ciències
Naturals, Barcelona.
H, M., R. E. R, M. W, K. A. L,  J. D.
B. . Corticosterone, testosterone and life-history strat-
egies of birds. Proceedings of the Royal Society of London, Series
B :–.
J, J. P., W. J. P, R. D. G,  S. A. W.
. Survival rates of tropical and temperate passerines: A Trini-
dadian perspective. American Naturalist :–.
J, G. M. . Explicit estimates from capture–recapture data
with both death and immigration-stochastic model. Biometrika
:–.
J, M.,  J. C. . The survival value of flocking in
Neotropical birds: Reality or fiction? Ecology :–.
ET AL.— AUK, VOL. 129
K, J. R., J. D. N, M. K. K,  J. D. B.
. Survival rates of birds of tropical and temperate forests:
Will the dogma survive? American Naturalist :–.
K, W. L., J. D. N,  J. E. H. . Estimating
temporary emigration using capture–recapture data with Pol-
lock’s robust design. Ecology :–.
L, S. C., C. J. R,  G. R. G. . Strategies for
the conservation monitoring of resident landbirds and wintering
Neotropical migrants in the Americas. Ornitologia Neotropical
:–.
LD, O. E., T. W. A, E. A. R,  F. J. C.
. Use of breeding and nonbreeding encounters to estimate
survival and breeding-site fidelity of the Piping Plover at the Great
Lakes. Condor :–.
L, M. S. . Designs for capture–mark–recapture studies:
Review, assessment, and emerging issues. Journal of Ornithology
 (Supplement ):S–S.
L, M. S., W. L. K, J. E. H,  M. G. A-
. . Combining band recovery data and Pollock’s robust
design to model temporary and permanent emigration. Biomet-
rics :–.
M, T. E. . Life history evolution in tropical and south tem-
perate birds: What do we really know? Journal of Avian Biology
:–.
M, T. E. . Avian life-history evolution has an eminent
past: Does it have a bright future? Auk :–.
MG, R., M. J. W,  W. C. .
Survival rates of tropical birds in Nigeria, West Africa. Ibis
:–.
N, I.,  L. D. . A comparative analysis of the avi-
faunas of different zoogeographical regions. Journal of Zoology
(London) :–.
P, T. H., C. D. B, B. K. S,  A. E.
A. . Apparent survival estimates for five species of
tropical birds in an endangered forest habitat in western Ecuador.
Biotropica :–.
P-H, J. W., R. C. B,  J. H. .
Survival of Band-tailed Manakins. Condor :–.
P, C. A. . Patterns of variance in stage-structured popula-
tions: Evolutionary predictions and ecological implications. Pro-
ceedings of the National Academy of Sciences USA :–.
P, L. A., M. J. C, J. E. H, J. D. N, 
D. G. K. . Simultaneous use of mark–recapture
and radio telemetry to estimate survival, movement, and capture
rates. Journal of Wildlife Management :–.
P, R., J. E. H, J.-D. L,  J. D. N. .
Capture–recapture survival models taking account of transients.
Biometrics :–.
R, C. J., G. R. G, P. P, T. E. M,  D. F.
DS. . Handbook of field methods for monitoring land-
birds. U.S. Department of Agriculture, Forest Service General
Technical Report PSW-GTR-.
R, R. E.,  M. W. . The physiology/life-
history nexus. Trends in Ecology & Evolution :–.
R, W. D., M. H, K. C. K, M. W, J. D. B,
S. H. A, C. E. T,  R. E. R. . Diversi-
fication of life histories in New World birds. Auk :–.
 JULY 2012 — PROTOCOL TO ESTIMATE SURVIVAL
R-G, V., T. A. G,  A. A. D. . Habi-
tat fragmentation lowers survival of a tropical forest bird. Ecolog-
ical Applications :–.
R-G, V., E. F. Z,  A. A. D. .
Occupancy dynamics in a tropical bird community: Unexpect-
edly high forest use by birds classified as non-forest species. Jour-
nal of Applied Ecology :–.
S, B. K. . Estimation of demographic parameters
from live-encounter data: A summary review. Journal of Wildlife
Management :–.
S, B. K.,  S. R. B. . Estimating rates
of population change for a Neotropical parrot with ratio, mark–
recapture and matrix methods. Journal of Applied Statistics
:–.
S [C.], E. . Dynamics of understory birds along a cloud
forest successional gradient. Ph.D. dissertation, University of
Wisconsin, Madison.
S, J., E. S C.,  I. R-T. . Bian-
nual cycles of the Cinnamon-bellied Flowerpiercer. Biotropica
:–.
S, G. A. F. . A note on the multiple-recapture census.
Biometrika :–.
View publication stats
FOR RESIDENT TROPICAL BIRDS — 509
T, C. E.,  J. D. B. . Family living in a Neo-
tropical bird: Variation in timing of dispersal and higher survival
for delayed dispersers. Animal Behaviour :–.
V, F.,  E. S C. . El monitoreo de poblacio-
nes: Herramienta necesaria para la conservación de aves en México.
Pages – in Conservación de Aves: Experiencias en México
(H. Gómez de Silva and A. Oliveras de Ita, Eds.). CIPAMEX, Mexico.
W, G. C.,  K. P. B. . Program MARK: Sur-
vival estimation from populations of marked animals. Bird Study
 (Supplement):S–S.
W, B. K., J. D. N,  M. J. C. . Analy-
sis and Management of Animal Populations. Academic Press, San
Diego, California.
Y, N. G., J. D. N,  T. B. . Monitor-
ing of biological diversity in space and time. Trends in Ecology &
Evolution :–.
Z, G. S., R. J. G,  W. S. L. .
Finite study areas and vital rates: Sampling effects on estimates of
Spotted Owl survival and population trends. Journal of Applied
Ecology :–.
Associate Editor: M. Lindberg