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One step forward: contrasting the effects of Toe clipping
and PIT tagging on frog survival and recapture probability
Murilo Guimar~
aes
1
,D
ecio T. Corr^
ea
1,2
,S
ergio S. Filho
3
, Thiago A. L. Oliveira
3
, Paul F. Doherty Jr
4
&
Ricardo J. Sawaya
5
1
Programa de P
os-Graduac
ß
~
ao em Ecologia, Departamento de Biologia Animal, Instituto de Biologia, Universidade Estadual de Campinas, CP 6109,
Campinas, SP cep 13083-970, Brazil
2
Ecology, Evolution, and Behavior Graduate Program, Department of Integrative Biology, The University of Texas at Austin, 1 University Station
#C0930, Austin, Texas 78712
3
Programa de P
os-Graduac
ß
~
ao em Biologia Animal, Departamento de Zoologia e Bot^
anica, Universidade Estadual Paulista, S~
ao Jos
e do Rio Preto,
SP Cep 15054-000, Brazil
4
Fish, Wildlife and Conservation Biology Department, Colorado State University, PO 80523, Fort Collins, Colorado
5
Departamento de Ci^
encias Biol
ogicas, Universidade Federal de S~
ao Paulo, Diadema, SP Cep 09972-270, Brazil
Keywords
Amphibians, detection probability, Hylidae,
mark–recapture, multimodel inference, return
rate.
Correspondence
Murilo Guimar~
aes, Instituto Nacional de
Pesquisas da Amaz^
onia, Avenida Andr
e
Ara
ujo, 2.936, CEP 69067-375 Manaus,
Brazil. Tel: (+55) 92 3643-1909;
Fax: (+55) 92 3643-3124;
E-mail: mu.guima@gmail.com
Funding Information
The authors thank CAPES (229611-0), CNPq
(140684/2009-3, 309229/2009-0), FADA-
UNIFESP, FAPESP (2008/54472-2), INCTTOX,
and the Ecology Graduate Program of
UNICAMP for financial support.
Received: 21 November 2013; Revised: 19
February 2014; Accepted: 25 February 2014
Ecology and Evolution 2014; 4(8): 1480–
1490
doi: 10.1002/ece3.1047
Abstract
Amphibians have been declining worldwide and the comprehension of the
threats that they face could be improved by using mark–recapture models to
estimate vital rates of natural populations. Recently, the consequences of mark-
ing amphibians have been under discussion and the effects of toe clipping on
survival are debatable, although it is still the most common technique for indi-
vidually identifying amphibians. The passive integrated transponder (PIT tag) is
an alternative technique, but comparisons among marking techniques in free-
ranging populations are still lacking. We compared these two marking tech-
niques using mark–recapture models to estimate apparent survival and recap-
ture probability of a neotropical population of the blacksmith tree frog,
Hypsiboas faber. We tested the effects of marking technique and number of toe
pads removed while controlling for sex. Survival was similar among groups,
although slightly decreased from individuals with one toe pad removed, to indi-
viduals with two and three toe pads removed, and finally to PIT-tagged individ-
uals. No sex differences were detected. Recapture probability slightly increased
with the number of toe pads removed and was the lowest for PIT-tagged indi-
viduals. Sex was an important predictor for recapture probability, with males
being nearly five times more likely to be recaptured. Potential negative effects
of both techniques may include reduced locomotion and high stress levels. We
recommend the use of covariates in models to better understand the effects of
marking techniques on frogs. Accounting for the effect of the technique on the
results should be considered, because most techniques may reduce survival.
Based on our results, but also on logistical and cost issues associated with PIT
tagging, we suggest the use of toe clipping with anurans like the blacksmith tree
frog.
Introduction
With the current state of amphibian declines (Stuart et al.
2004), quantitative links between vital rates and explana-
tory covariates are fundamental to understand the
dynamics of and threats to populations (Biek et al. 2002).
The results obtained by marking individuals provide
accurate information on population trends and demo-
graphic estimates (Manly et al. 2005), especially when
population dynamics are poorly understood, as in the
Neotropics (Hiert et al. 2012).
Although field biologists strive to apply the least harm-
ful marking technique to their study species, most tech-
niques remain at least somewhat invasive and may affect
1480 ª2014 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
This is an open access article under the terms of the Creative Commons Attribution License, which permits use,
distribution and reproduction in any medium, provided the original work is properly cited.
individual behavior and survival (Lemckert 1996; Bloch
and Irschick 2004; Ferner 2007; Schmidt and Schwarzkopf
2010). Among the different techniques used to mark anu-
rans (Donnelly et al. 1994), the most common is toe clip-
ping (Bogert 1947), which consists of removing different
combinations of digits to give individuals unique marks.
Nevertheless, the scientific community has divergent
opinions regarding the impacts of marking individuals,
especially via toe clipping (May 2004; Funk et al. 2005).
Besides that, environmental agencies and common sense
from different countries have also expressed concerns
about the efficacy of such a potential unappealing tech-
nique for studying frogs and other vertebrates (Ferner
2007). Their opinion derives from several recent papers
that have related the number of toes clipped to individual
response of amphibians, including low return or survival
rates (Parris and McCarthy 2001; McCarthy and Parris
2004; Waddle et al. 2008). In fact, philosophical and legal
views deserve attention as methodological efficacy is not
the only concern when considering toe clipping (for a
review, see Perry et al. 2011). In Brazil, for instance, envi-
ronmental agencies and nongovernmental organizations
claim that toe clipping is a form of mutilation and its use
should be prohibited (Corr^
ea et al. 2013). In general, eth-
ical standard policies on animal welfare state that marking
techniques should not cause distress or inflict pain,
reducing individual survival. And because of the contro-
versial results to date, Brazil suggested the use of alterna-
tive marking techniques, including visible implanted
elastomers or photographs of natural marks (Brown 1997;
Hoffmann et al. 2008; Campbell et al. 2009; Kenyon et al.
2009), but these methods remain to be contrasted.
The passive integrated transponder (PIT tag) is used
worldwide and recommended as an alternative to toe clip-
ping (Donnelly et al. 1994; Gibbons and Andrews 2004;
Phillott et al. 2008). It consists of a glass-encapsulated
electromagnetic coil with a unique alphanumeric code.
The tag is lodged under the skin or in the body cavity of
an animal and read by a handheld scanner (Gibbons and
Andrews 2004). Because of the possibility to mark a great
number of individuals, PIT tags have been used in anurans
as an alternative marking technique (Christy 1996; Jehle
and H€
odl 1998). Nevertheless, negative effects on frog sur-
vival have been reported (Scherer et al. 2005), and little is
known on the impacts of PIT tags on anurans (e.g.,
Christy 1996; Brown 1997; Phillott et al. 2008), which
may include behavioral and physiological deleterious
effects from the injection of the tag. Overall, direct com-
parisons between different marking techniques in frogs are
lacking, which does not allow discussion and the clarifica-
tion of the effects of marking to advance.
Another issue for studies that have attempted to quan-
tify the effects of marking techniques on frogs is that
studies should explicitly consider individual detectability.
Past studies looking at toe-clipping effects have used the
return rate (e.g., McCarthy and Parris 2004). The return
rate assumes that detection probability does not change,
which is unrealistic in natural systems due to behavioral
heterogeneity (e.g., between sexes) and climatic condi-
tions, such as rainfall, which influences amphibian activity
(Duellman and Trueb 1986). Despite the number of stud-
ies reporting decreased return rates with increasing num-
ber of toes removed, only a few studies have incorporated
detection probability (e.g., Waddle et al. 2008; Grafe et al.
2011), which is likely less than one, into survival esti-
mates. If recapture probability differs among groups, but
not survival probability, one could conclude through the
use of return rates, that toe clipping reduces survival
when in fact only recapture probability is reduced.
To further advance the discussion on the topic, we
contrasted the effects of toe clipping and PIT tagging on
a free-ranging neotropical tree frog population using
mark–recapture models, which allowed us to disentangle
survival and recapture probabilities (Schmidt 2003; Wad-
dle et al. 2008). We specifically compared survival and
recapture probabilities between marking techniques while
controlling for sex differences. Our intention is to estab-
lish a direct comparison between two of the commonest
marking techniques and to provide scientific basis on
amphibian conservation biology for field biologists and
policy makers.
Materials and Methods
Study site and study species
We conducted this study in a 970-m
2
permanent pond in
Estac
ß
~
ao Ecol
ogica de Jata
ı (21°33059.75″S, 47°43033.19″
W), a protected area in the state of S~
ao Paulo, southeast-
ern Brazil. The reserve is located in a transitional area
between the Atlantic Forest and Cerrado biomes, com-
posed of open grassy areas and semi-deciduous forests.
Average temperature in the coldest months (June to
August) is about 11°C, and about 30°C in the hottest
months (December to February). Annual rainfall is about
1500 mm. Precipitation during the rainy season (October
to March) typically exceeds 270 mm per month, but does
not exceed 27 mm per month during the dry season
(April to September).
We sampled an adult population of the blacksmith tree
frog Hypsiboas faber (Anura, Hylidae, Appendix A), a
large tree frog (snout-vent length =92.3 4.8 mm,
N=305; this population) distributed from northern
Argentina to eastern Brazil (Martins 1993). As in most
amphibian species behavior is sexually divergent, and
males H. faber occupy a pond and build nests at the
ª2014 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd. 1481
M. Guimar~
aes et al. Effects of Marking Techniques on Frogs
beginning of the breeding season, generally from October
to March (pers. obs. D.T.C.), then begin to vocalize until
a female approaches to inspect the nest (Martins and
Haddad 1988).
Data Collection
We collected data during two reproductive seasons, from
November to March, in 2010–2011 and 2011–2012. We
captured individuals during three nights per month and
pooled nights within the same month together, resulting
in 10 sampling occasions, five for each breeding season.
On each capture occasion, three observers systematically
walked around the pond covering from the margins until
approximately 150 cm of water depth. Adult individuals
were captured by hand based on visual and acoustic cues
in all accessible microhabitats.
We determined sex and randomly assigned one type of
marking technique to each individual, toe clipping or PIT
tagging (IBAMA permit number: 10423-1, COTEC permit
number: 010.157/2010). We placed individuals in four
different groups: one toe clipped, two toes clipped, three
toes clipped, and PIT tags.
For toe-clipping groups, we adapted the marking tech-
nique of Waichman (1992), removing only the toe pad
(“toe tipping”, sensu Phillott et al. 2007), which is enough
for individual recognition as tissue regeneration is rarely
observed (L€
uddecke and Am
ezquita 1999; Phillott et al.
2007; Grafe et al. 2011). We marked individuals in the
toe pad-clipping groups starting with the removal of one
toe pad. When all combinations for removing one toe
pad were used, we started removing two toe pads in
unique combinations, and finally, three toe pad removal
combinations, clipping up to two toe pads per limb. This
sampling design generates uneven toe pad removal groups
over the study period (see Grafe et al. 2011), which we
accounted for by including a linear trend model in the
parameter estimates (see below).
The PIT-tagging group received a 2.2 mm 912.2 mm,
0.5 g internal transponder (Animall Tag Company)
implanted in a posterior laterally dorsum position, using
sterilized needles and followed by the use of glue to help
healing. The PIT tag/tree frog mass ratio was 1%. To con-
trol for possible effects of the glue, all individuals from
the toe pad-clipping groups also received the same
amount of glue on the same body region of the PIT tag.
The effects of marking technique and number of toes
tipped were assessed in three different ways: (1) by com-
paring the effects of toe pad clipping against PIT tagging,
where we combined groups one, two, and three and com-
pared with group four (hereafter “marking technique
effect”); (2) by comparing the survival on individuals
with one, two, three toes pad clipped and PIT tagged
(hereafter “group effect”); and finally (3) by forcing a lin-
ear trend effect only on toe pad-clipping groups (hereaf-
ter “linear trend effect”), to test the hypothesis that
removal of more toes decreased survival. Also, the inclu-
sion of a linear trend in the recapture probability was
used to account for the uneven toe pad removal groups
over the study. We used sex as an individual covariate as
we expect differences between males and females.
Statistical analysis
We obtained maximum likelihood parameter estimates
using a Cormack–Jolly–Seber (CJS) model (Cormack
1964; Jolly 1965; Seber 1965) in Program MARK version
6.1 (White and Burnham 1999). The CJS model estimates
the apparent survival probability (Φ), which is a combi-
nation of true survival and site fidelity, and recapture
probability (P). We used the strategy proposed by Doher-
ty et al. (2010) to run all possible additive combinations
of factors (marking techniques and sex), except for com-
binations that did not make logical sense (e.g., different
representations of marking effects not considered a pri-
ori). Such a strategy is recommended over stepwise proce-
dures (Doherty et al. 2010), but it may generate a large
number of models.
Goodness-of-fit and a variance inflation factor (i.e.,
median ^
c) were assessed using the general model with no
individual or temporal covariates (Φ
group
P
group
) to test
the mark–recapture assumptions. We selected and ranked
models using Akaike Information Criterion (Akaike 1973)
adjusted for small sample sizes (AICc, Burnham and
Anderson 2002). Survival and recapture probabilities were
then model averaged, a weighted average of the model-
specific parameter estimates based on Akaike weights, to
include uncertainty in model selection (Burnham and
Anderson 2002). We then calculated the relative impor-
tance of each covariate through the cumulative AICc
weights to determine the important covariates for each
parameter. Following Barbieri and Berger (2004), we con-
sidered covariates with cumulative AICcweight above 0.5
to be important.
Results
Eighteen individuals (14 males and four females) had one
toe pad clipped, 150 (110 males and 40 females) had two
toe pads clipped, another 150 individuals (120 males and
30 females) had three toe pads clipped, and 227 individu-
als (177 males and 50 females) were PIT tagged. We
recaptured 117 of 545 individuals at least once. The good-
ness-of-fit test showed no problem with transient individ-
uals or trap dependence effects, and no extra binomial
variation was detected (^
c=0.96).
1482 ª2014 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
Effects of Marking Techniques on Frogs M. Guimar~
aes et al.
The top model (AICcweight =0.07) included constant
apparent survival and detection probability varying as an
additive effect of sex and linear trend on toe pad-clipped
groups. However, models had similar AICcweights, with
considerable model selection uncertainty (Appendix B).
Considering all models averaged, apparent monthly sur-
vival probability was similar among the four groups, with
slightly higher survival probability for individuals with
one toe pad clipped (0.77), then two (0.75) and three
(0.74) toe pads clipped, similar to McCarthy and Parris
(2004). Survival was the lowest for PIT-tagged individuals
(0.72), but there was also considerable uncertainty around
the estimates (Fig. 1, Appendix C). Assuming group one’s
survival estimate as the closest of a control group (as we
have no estimates for individuals with no toe pads
clipped), removing the second toe pad reduced survival
probability in 2.6% in relation to group one, and the
third toe pad in 3.6%. The use of PIT tag reduced sur-
vival in 5.8% in relation to group one’s point estimate.
No covariate was important for describing the apparent
survival probability, because all of them presented AICc
cumulative model weights below 0.5 (Appendix C).
PIT tagged males and females presented lower recapture
probability than toe pad-clipped groups, but confidence
intervals greatly overlapped (Fig. 2). The same trend was
also showed by the bestimate (b
technique
=0.35, CI 0.07 to
0.8, second top model). The AICcmodel weight of the linear
trend effect on toe pad clipping was close to the 0.5 cutoff
(Appendix B), with a slightly higher probability of recapture
with the increase of toe pads removed, but confidence inter-
vals overlapped among groups and included zero
(b
toe_lin
=0.15, CI 0.01 to 0.3, top model). Assuming indi-
viduals with one toe pad clipped as the closest of a control
group, we observed an increase on recapture probability of
6% and 8% for individuals with two toe pads clipped, 17%
and 26% for individuals with three toe pads clipped, and a
decrease of 9% and 21% for PIT-tagged individuals in males
and females, respectively. Sex was the most important covar-
iate predicting recapture probability (0.98 of model weight;
Appendix C), where males (from 0.20 to 0.26) and females
(from 0.03 to 0.05) greatly differed (Fig. 2).
Discussion
We found subtle differences in survival probability among
individuals marked with different techniques. Although
there was uncertainty around the estimates, we should
look at the potential biological differences among groups.
Looking only at the point estimates, we see that survival
probability of individuals with only one toe pad clipped
was slightly higher than other groups, declining 2.6% with
two toe pads clipped, 3.6% with three toe pads clipped,
and finally declining 5.8% in PIT-tagged individuals. Our
estimated declines were lower than those predicted by
McCarthy and Parris (2004) that reported declines from
4% to 11% when clipping two and three toes respectively.
The difference between both studies may relate to differ-
ent species and procedures used, but also because they
used return rates that confound survival and recapture
probabilities. Overall, clipping multiple toes can reduce
survival substantially.
Differences were observed among groups concerning
recapture probability, but confidence intervals overlapped.
Among toe pad-clipped groups, recapture probability
increased with more toe pads clipped –from 6% to 26%
for males and females in individuals with two and three
Figure 1. Model-averaged monthly apparent survival probability (and
95% CI) among groups.
Figure 2. Model-averaged monthly recapture probability (and 95%
CI) among groups and sexes.
ª2014 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd. 1483
M. Guimar~
aes et al. Effects of Marking Techniques on Frogs
toe pads removed. This may be related to reduction of
individual mobility due to the removal of toe pads. Addi-
tionally, recapture probability of individuals with two and
three toe pads removed could be higher because more
individuals were allocated to those groups, or even because
individuals in group one were marked at the onset of the
first reproductive season, and they could leave the site first.
All these hypotheses would be better addressed randomly
assigning individuals to the different groups since the
beginning of the study. PIT-tagged individuals presented
the lowest recapture probability, decreasing 9% and 21%
in males and females, respectively. Sex was the strongest
covariate influencing recapture probability, carrying the
most cumulative AICcweight.
Toe clipping may be a stressor for amphibians if com-
pared to handling only (Narayan et al. 2011; but see Fisher
et al. 2013), and negative effects of toe clipping on frog sur-
vival and capturing have been observed previously (Lemck-
ert 1996; van Gelder and Strijbosch 1996; Hartel and
Nemes 2006). In our study, the removal of toe pads was
quicker than marking with the PIT tag, and bleeding usu-
ally did not occur. The application of the PIT tag took from
two to four times longer (pers. obs. M.G.), and possibly
increased handling stress. Many males, especially from toe
pad-clipping groups, were seen in normal reproductive
activities right after being manipulated and no injures pos-
sibly caused by toe clipping were observed during the study.
Despite its low cost and ease of use, the number of individ-
uals to be marked using toe clipping/tipping is limited and
should always be the smallest as possible. In our study, we
did not find evidence of toe regeneration, but if it occurs,
as described by Hoffmann et al. (2008), the mark–recap-
ture assumption of mark retention will be violated, under-
estimating survival and other vital rates (Lebreton et al.
1992; Williams et al. 2002).
Males and females H.faber present behavioral differ-
ences (Duellman and Trueb 1986; Martins and Haddad
1988; Martins 1993). Females spend less time in the
ponds, and this may explain the lower recapture probabil-
ity observed for them. The difference in the recapture
probability of males and females indicates that pooling
the sexes in the analysis would mask results. In this way,
considering the return rate a survival estimate would have
provided underestimated estimates (Martin et al. 1995).
The importance of sex to our analysis highlights the
importance of considering individual (and also temporal,
though not used here) covariates when studying potential
effects of marking techniques. Most of the articles so far
(Waddle et al. 2008; Grafe et al. 2011) do not present
comparisons including such covariates.
The use of PIT tags in anurans might be less common
than toe clipping, but may be a reliable technique for cer-
tain species and has not generally been demonstrated to
cause serious problems, such as detrimental effects on body
condition or mortality (Christy 1996; Brown 1997; Jehle
and H€
odl 1998; McAllister et al. 2004; but see Scherer
et al. 2005). However, PIT tagging is more costly, increases
handling time, requires more skill from the field biologist,
and may be unfeasible in small frogs. We are unable to
clearly demonstrate the difference between both marking
techniques based only on our data and the effects and dif-
ferences among techniques should be emphasized in future
studies. Assuming the difference is real, the stress of higher
handling time and PIT tag implant procedure could make
individuals to leave the reproductive site, reducing their
recapture probability. In this case, individuals would need
more time to recover, after being tagged. In addition, while
being considered a permanent marking technique (Gibbons
and Andrews 2004), PIT tags could be expelled from
(Roark and Dorcas 2000) or migrate to another location in
the body (Tracy et al. 2011) causing apparent tag loss and
affecting population estimates.
In general, studies looking at the impacts of different
marking techniques on vital rates of wild populations are
scarce. Studies comparing toe clipping and PIT tagging
have shown similar effects on survival and growth rates of
salamanders (Ott and Scott 1999) and free-living naked
mole rats (Braude and Ciszek 1998). It is also noteworthy
that the effects of marking techniques vary by species,
reproductive strategies, habitats (e.g., arboreal vs. fosso-
rial), and behaviors (Liner and Smith 2007). Frog species
will respond in different ways to marking and investiga-
tors must consider the characteristics of each species, as
well as the use of the most practical and least harmful
technique, evaluating all methods together, as suggested
by Phillott et al. (2008). Stress response should be
included as an important trait to be measured in individ-
uals (Perry et al. 2011), but few studies considered this
trait when testing the impacts of different invasive mark-
ing techniques on amphibians.
Estimating vital rates of a control group of nonmarked
individuals in the field would be ideal, as we were unable
to compare survival of individuals that were not marked,
and because we believe that both techniques may decrease
survival. Perry et al. (2011) suggest the use of Visual
Implanted Elastomers (VIE) as a true control group, but
handling and inserting the elastomers under a frog’s skin
might also cause stress. Photography may be a good can-
didate for a control group in free-ranging populations.
Photography also presents problems, like identifiable
characteristics on the target species and obtaining good
quality pictures without disturbing individuals, as han-
dling only may be an important stressor itself (Fisher
et al. 2013). Controlled laboratory experiments may be
useful for the inclusion of a nonmarked group, for
comparing survival (e.g., looking for inflammation or
1484 ª2014 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
Effects of Marking Techniques on Frogs M. Guimar~
aes et al.
stress responses) and for allowing estimation of tag loss
(Brown 1997). However, laboratory experiments are not
the best solution to observe the effects of marking on spe-
cies interactions (e.g., predation, competition) as well as
the effects of weather variability on marked individuals.
In summary, we showed slight differences between both
marking techniques. Considering only the statistical results
would make us conclude that both techniques performed
similarly. Although not discussed here, the decision to
adopt a particular marking method should be multidisci-
plinary, also involving law, ethics and philosophy. How-
ever, given the urge of studying and preserving
populations, as well as the pros and cons of each tech-
nique, a decision has to be put into practice. No perfect
technique is available, but being aware of the problems
and accounting for the effect of the chosen technique in
the analysis is better than ignoring such problems. Based
on our estimates of survival and recapture, but also given
the lack of comparisons among alternative marking tech-
niques in the literature, as well as logistical issues, such as
budget and processing time, we agree with others authors
and recommend the use of toe clipping instead of PIT tag-
ging with the blacksmith tree frog. However, there should
be a threshold, where toe clipping is not worthwhile (when
removing multiple toes, for instance) and another tech-
nique to individualize frogs should be considered.
Acknowledgments
We thank the Wagar 113 group for discussions on earlier
versions of this manuscript. Eduardo Martins, Gonc
ßalo
Ferraz, Larissa Bailey, and four anonymous reviewers pro-
vided insightful comments that improved the manuscript.
Andr
e Scatigna, Jos
e Ferrarini, Karina Silva, Juliana Sou-
za, Lucas Crivelari and Thiago Pires provided field assis-
tance. Edson Montilha, Instituto Florestal and Fundac
ß
~
ao
Florestal provided support at Estac
ß
~
ao Ecol
ogica de Jata
ı.
The authors thank CAPES (229611-0), CNPq (140684/
2009-3, 309229/2009-0), FADA-UNIFESP, FAPESP (2008/
54472-2), INCTTOX, and the Ecology Graduate Program
of UNICAMP for financial support.
Conflict of Interest
None declared.
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Appendix A
An adult male Hypsiboas faber. Photo credit: D. T. Corr^
ea.
Appendix B
Model results.
Model AICcDAICcw k Deviance
Phi(.) p(sex+toe_lin) 849.53 0.00 0.07 4 841.47
Phi(.) p(sex+tech) 850.27 0.74 0.05 4 842.20
Phi(tech) p(sex) 850.91 1.38 0.04 4 842.85
Phi(sex) p(sex+toe_lin) 850.93 1.41 0.04 5 840.84
Phi(.) p(sex) 850.95 1.42 0.04 3 844.91
Phi(tech) p(sex+toe_lin) 851.29 1.76 0.03 5 841.19
Phi(.) p(sex+tech+toe_lin) 851.54 2.01 0.03 5 841.45
Phi(toe_lin) p(sex+toe_lin) 851.55 2.03 0.03 5 841.46
Phi(toe_lin) p(sex) 851.66 2.13 0.03 4 843.60
Phi(tech+toe_lin) p(sex) 851.68 2.15 0.03 5 841.58
Phi(sex) p(sex+tech) 851.70 2.17 0.03 5 841.60
Phi(tech) p(sex+tech) 851.98 2.46 0.02 5 841.89
Phi(tech+toe_lin) p
(sex+toe_lin)
852.13 2.60 0.02 6 840.00
Phi(toe_lin) p(sex+tech) 852.27 2.74 0.02 5 842.18
Phi(sex) p(sex) 852.33 2.80 0.02 4 844.27
Phi(tech+toe_lin) p
(sex+toe_lin)
852.44 2.91 0.02 6 840.30
Phi(sex+toe_lin) p(sex) 852.62 3.09 0.02 5 842.53
Phi(tech+toe_lin) p(sex) 852.76 3.23 0.01 5 842.67
Phi(sex) p(sex+tech+toe_lin) 852.95 3.42 0.01 6 840.81
Appendix B. Continued.
Model AICcDAICcw k Deviance
Phi(sex+toe_lin) p
(sex+toe_lin)
852.96 3.43 0.01 6 840.83
Phi(sex+tech) p(sex+tech) 853.12 3.59 0.01 6 840.98
Phi(tech) p
(sex+tech+toe_lin)
853.22 3.69 0.01 6 841.09
Phi(tech+toe_lin) p
(sex+tech+toe_lin)
853.31 3.78 0.01 7 839.13
Phi(.) p(sex) 853.36 3.83 0.01 6 841.22
Phi(.) p(g+sex+tech) 853.36 3.83 0.01 6 841.22
Phi(.) p(g+sex+toe_lin) 853.36 3.83 0.01 6 841.22
Phi(.) p(g+sex+tech+toe_lin) 853.36 3.83 0.01 6 841.22
Phi(sex+tech+toe_lin) p
(sex+toe_lin)
853.44 3.91 0.01 7 839.26
Phi(sex+tech+toe_lin) p(sex) 853.53 4.00 0.01 6 841.40
Phi(toe_lin) p
(sex+tech+toe_lin)
853.57 4.05 0.01 6 841.44
Phi(sex+toe_lin) p(sex+tech) 853.61 4.08 0.01 6 841.47
Phi(tech+toe_lin) p
(sex+tech)
853.65 4.12 0.01 6 841.51
Phi(g) p(sex+toe_lin) 853.75 4.22 0.01 7 839.57
Phi(g+tech) p(sex+toe_lin) 853.75 4.22 0.01 7 839.57
Phi(g+toe_lin) p(sex+toe_lin) 853.75 4.22 0.01 7 839.57
p(g+tech+toe_lin) p
(sex+toe_lin)
853.75 4.22 0.01 7 839.57
Phi(g) p(sex) 853.75 4.22 0.01 6 841.62
Phi(g+tech) p(sex) 853.75 4.22 0.01 6 841.62
Phi(g+toe_lin) p(sex) 853.75 4.22 0.01 6 841.62
Phi(g+tech+toe_lin) p(sex) 853.75 4.22 0.01 6 841.62
Phi(sex+tech) p
(sex+tech+toe_lin)
854.29 4.76 0.01 7 840.11
Phi(sex+tech+toe_lin) p
(sex+tech+toe_lin)
854.46 4.93 0.01 8 838.23
Phi(g+sex) p(sex) 854.53 5.00 0.01 7 840.35
Phi(g+sex+tech) p(sex) 854.53 5.00 0.01 7 840.35
Phi(g+sex+toe_lin) p(sex) 854.53 5.00 0.01 7 840.35
Phi(g+sex+tech+toe_lin) p
(sex)
854.53 5.00 0.01 7 840.35
Phi(sex) p(g+sex) 854.75 5.22 0.01 7 840.58
Phi(sex) p(g) p(sex+tech) 854.75 5.22 0.01 7 840.58
Phi(sex) p(g+sex+toe_lin) 854.75 5.22 0.01 7 840.58
Phi(sex) p
(g+sex+tech+toe_lin)
854.75 5.22 0.01 7 840.58
Phi(sex+tech+toe_lin) p
(sex+tech)
854.83 5.30 0.01 7 840.65
Phi(g) p(sex+tech) 854.92 5.39 0.01 7 840.75
Phi(g+tech) p(sex+tech) 854.92 5.39 0.01 7 840.75
Phi(g+toe_lin) p(sex+tech) 854.92 5.39 0.01 7 840.75
Phi(g+tech+toe_lin) p
(sex+tech)
854.92 5.39 0.01 7 840.75
Phi(sex+toe_lin) p
(sex+tech+toe_lin)
854.97 5.44 0.00 7 840.80
Phi(g+sex) p(sex+toe_lin) 855.00 5.47 0.00 8 838.77
Phi(g+sex+tech) p
(sex+toe_lin)
855.00 5.47 0.00 8 838.77
Phi(g+sex+toe_lin) p
(sex+toe_lin)
855.00 5.47 0.00 8 838.77
ª2014 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd. 1487
M. Guimar~
aes et al. Effects of Marking Techniques on Frogs
Appendix B. Continued.
Model AICcDAICcw k Deviance
Phi(tech) p(g+sex) 855.02 5.49 0.00 7 840.84
Phi(tech) p(g+sex+tech) 855.02 5.49 0.00 7 840.84
Phi(tech) p(g+sex+toe_lin) 855.02 5.49 0.00 7 840.84
Phi(tech) p
(g+sex+tech+toe_lin)
855.02 5.49 0.00 7 840.84
Phi(g) p(sex+tech+toe_lin) 855.08 5.56 0.00 8 838.86
Phi(g+tech) p
(sex+tech+toe_lin)
855.08 5.56 0.00 8 838.86
Phi(g+toe_lin) p
(sex+tech+toe_lin)
855.08 5.56 0.00 8 838.86
Phi(tech+toe_lin) p(g+sex) 855.10 5.57 0.00 8 838.87
Phi(tech+toe_lin) p
(g+sex+tech)
855.10 5.57 0.00 8 838.87
Phi(tech+toe_lin) p
(g+sex+toe_lin)
855.10 5.57 0.00 8 838.87
Phi(toe_lin) p(g+sex) 855.40 5.87 0.00 7 841.22
Phi(toe_lin) p(g+sex+tech) 855.40 5.87 0.00 7 841.22
Phi(toe_lin) p(g+sex+toe_lin) 855.40 5.87 0.00 7 841.22
Phi(toe_lin) p
(g+sex+tech+toe_lin)
855.40 5.87 0.00 7 841.22
Phi(g+sex) p(sex+tech) 856.05 6.52 0.00 8 839.82
Phi(g+sex+tech) p(sex+tech) 856.05 6.52 0.00 8 839.82
Phi(g+sex+toe_lin) p
(sex+tech)
856.05 6.52 0.00 8 839.82
Phi(sex+tech) p(g+sex) 856.05 6.53 0.00 8 839.83
Phi(sex+tech) p(g+sex+tech) 856.05 6.53 0.00 8 839.83
Phi(sex+tech) p
(g+sex+toe_lin)
856.05 6.53 0.00 8 839.83
Phi(g+sex) p
(sex+tech+toe_lin)
856.20 6.67 0.00 9 837.92
Phi(g+sex+tech) p
(sex+tech+toe_lin)
856.20 6.67 0.00 9 837.92
Phi(g+sex+toe_lin) p
(sex+tech+toe_lin)
856.20 6.67 0.00 9 837.92
Phi(g+sex+tech+toe_lin) p
(sex+tech+toe_lin)
856.20 6.67 0.00 9 837.92
Phi(sex+tech+toe_lin) p
(g+sex)
856.23 6.70 0.00 9 837.94
Phi(sex+tech+toe_lin) p
(g+sex+tech)
856.23 6.70 0.00 9 837.94
Phi(sex+tech+toe_lin) p
(g+sex+toe_lin)
856.23 6.70 0.00 9 837.94
Phi(sex+tech+toe_lin) p
(g+sex+tech+toe_lin)
856.23 6.70 0.00 9 837.94
Phi(sex+toe_lin) p(g+sex) 856.78 7.25 0.00 8 840.55
Phi(sex+toe_lin) p
(g+sex+tech)
856.78 7.25 0.00 8 840.55
Phi(sex+toe_lin) p
(g+sex+toe_lin)
856.78 7.25 0.00 8 840.55
Phi(sex+toe_lin) p
(g+sex+tech+toe_lin)
856.78 7.25 0.00 8 840.55
Phi(g+sex+tech+toe_lin p
(sex+toe_lin)
857.06 7.53 0.00 9 838.77
Phi(.) phi g p(g+sex) 857.07 7.54 0.00 9 838.79
Phi(g+tech) p(g+sex) 857.07 7.54 0.00 9 838.79
Phi(g+toe_lin) p(g+sex) 857.07 7.54 0.00 9 838.79
Appendix B. Continued.
Model AICcDAICcw k Deviance
Phi(g) p(g+sex+tech) 857.07 7.54 0.00 9 838.79
Phi(g) p(g+sex+toe_lin) 857.07 7.54 0.00 9 838.79
Phi(g+tech) p
(g+sex+toe_lin)
857.07 7.54 0.00 9 838.79
Phi(g+toe_lin) p
(g+sex+toe_lin)
857.07 7.54 0.00 9 838.79
Phi(g+tech+toe_lin) p
(g+sex+toe_lin)
857.07 7.54 0.00 9 838.79
Phi(g) p
(g+sex+tech+toe_lin)
857.07 7.54 0.00 9 838.79
Phi(g+tech) p
(g+sex+tech+toe_lin)
857.07 7.54 0.00 9 838.79
Phi(g+tech+toe_lin) p
(sex+tech+toe_lin)
857.14 7.61 0.00 9 838.86
Phi(tech+toe_lin) p
(g+sex+tech+toe_lin)
857.16 7.63 0.00 9 838.87
Phi(g+sex+tech+toe_lin) p
(sex+tech)
858.11 8.58 0.00 9 839.82
Phi(sex+tech) p(g) p
(sex+tech+toe_lin)
858.11 8.58 0.00 9 839.83
Phi(g+sex) p(g+sex) 858.19 8.66 0.00 10 837.84
Phi(g+sex+tech) p(g+sex) 858.19 8.66 0.00 10 837.84
Phi(g+sex+toe_lin) p(g+sex) 858.19 8.66 0.00 10 837.84
Phi(g+sex) p(g+sex+tech) 858.19 8.66 0.00 10 837.84
Phi(g+sex+tech) p
(g+sex+tech)
858.19 8.66 0.00 10 837.84
Phi(g+sex+toe_lin) p
(g+sex+tech)
858.19 8.66 0.00 10 837.84
Phi(g+sex) p(g+sex+toe_lin) 858.19 8.66 0.00 10 837.84
Phi(g+sex+tech) p
(g+sex+toe_lin)
858.19 8.66 0.00 10 837.84
Phi(g+tech+toe_lin) p
(g+sex)
859.14 9.61 0.00 10 838.79
Phi(g+tech) p(g+sex+tech) 859.14 9.61 0.00 10 838.79
Phi(g+toe_lin) p
(g+sex+tech)
859.14 9.61 0.00 10 838.79
Phi(g+tech+toe_lin) p
(g+sex+tech)
859.14 9.61 0.00 10 838.79
Phi(g+toe_lin) p
(g+sex+tech+toe_lin)
859.14 9.61 0.00 10 838.79
Phi(g+sex+tech+toe_lin) p
(g+sex)
860.26 10.73 0.00 11 837.84
Phi(g+sex+toe_lin) p
(g+sex+toe_lin)
860.26 10.73 0.00 11 837.84
Phi(g+sex) p
(g+sex+tech+toe_lin)
860.26 10.73 0.00 11 837.84
Phi(g+sex+toe_lin) p
(g+sex+tech+toe_lin)
860.26 10.73 0.00 11 837.84
Phi(g+tech+toe_lin) p
(g+sex+tech+toe_lin)
861.21 11.68 0.00 11 838.79
Phi(g+sex+tech+toe_lin) p
(g+sex+tech)
862.34 12.81 0.00 12 837.84
Phi(g+sex+tech+toe_lin) p
(g+sex+toe_lin)
862.34 12.81 0.00 12 837.84
Phi(g+sex+tech) p
(g+sex+tech+toe_lin)
862.34 12.81 0.00 12 837.84
1488 ª2014 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
Effects of Marking Techniques on Frogs M. Guimar~
aes et al.
Appendix B. Continued.
Model AICcDAICcw k Deviance
Phi(g+sex+tech+toe_lin) p
(g+sex+tech+toe_lin)
866.51 16.98 0.00 14 837.84
Phi(sex) p(toe_lin) 870.13 20.60 0.00 4 862.07
Phi(sex) p(tech) 870.94 21.41 0.00 4 862.88
Phi(sex+toe_lin) p(toe_lin) 871.07 21.54 0.00 5 860.98
Phi(sex+tech+toe_lin) p
(toe_lin)
871.98 22.45 0.00 6 859.85
Phi(sex) p(.) 871.99 22.46 0.00 3 865.95
Phi(sex+tech) p(toe_lin) 872.05 22.52 0.00 5 861.96
Phi(sex) p(tech+toe_lin) 872.14 22.62 0.00 5 862.05
Phi(sex+toe_lin) p(tech) 872.43 22.90 0.00 5 862.33
Phi(sex+tech) p(tech) 872.85 23.32 0.00 5 862.75
Phi(sex+toe_lin) p
(tech+toe_lin)
873.10 23.57 0.00 6 860.97
Phi(sex+tech) p(.) 873.35 23.82 0.00 4 865.29
Phi(sex+tech+toe_lin) p
(tech+toe_lin)
873.35 23.82 0.00 7 859.17
Phi(g+sex) p(toe_lin) 873.60 24.07 0.00 7 859.42
Phi(g+sex+tech) p(toe_lin) 873.60 24.07 0.00 7 859.42
Phi(g+sex+toe_lin) p(toe_lin) 873.60 24.07 0.00 7 859.42
Phi(sex+toe_lin) p(.) 873.75 24.22 0.00 4 865.68
Phi(sex) p(g) 873.83 24.30 0.00 6 861.69
Phi(sex) p(g+tech) 873.83 24.30 0.00 6 861.69
Phi(sex) p(g+toe_lin) 873.83 24.30 0.00 6 861.69
Phi(sex+tech) p
(tech+toe_lin)
874.09 24.56 0.00 6 861.95
Phi(sex+tech+toe_lin) p
(tech)
874.22 24.69 0.00 6 862.08
Phi(sex+toe_lin) p(g) 874.85 25.32 0.00 7 860.67
Phi(sex+toe_lin) p(g+tech) 874.85 25.32 0.00 7 860.67
Phi(sex+toe_lin) p(g+toe_lin) 874.85 25.32 0.00 7 860.67
Phi(sex+tech+toe_lin) p(g) 875.03 25.50 0.00 8 858.80
Phi(sex+tech+toe_lin) p
(g+tech)
875.03 25.50 0.00 8 858.80
Phi(sex+tech+toe_lin) p
(g+toe_lin)
875.03 25.50 0.00 8 858.80
Phi(g+sex) p(tech+toe_lin) 875.09 25.56 0.00 8 858.86
Phi(g+sex+tech) p
(tech+toe_lin)
875.09 25.56 0.00 8 858.86
Phi(g+sex+toe_lin) p
(tech+toe_lin)
875.09 25.56 0.00 8 858.86
Phi(sex+tech+toe_lin p(.) 875.09 25.57 0.00 5 865.00
Phi(g+sex) p(tech) 875.49 25.96 0.00 7 861.32
Phi(g+sex+tech) p(tech) 875.49 25.96 0.00 7 861.32
Phi(g+sex+toe_lin) p(tech) 875.49 25.96 0.00 7 861.32
Phi(g+sex+tech+toe_lin) p
(tech)
875.49 25.96 0.00 7 861.32
Phi(g+sex+tech+toe_lin) p
(toe_lin)
875.65 26.12 0.00 8 859.42
Phi(sex+tech) p(g) 875.80 26.27 0.00 7 861.62
Phi(sex+tech) p(g+tech) 875.80 26.27 0.00 7 861.62
Phi(sex+tech) p(g+toe_lin) 875.80 26.27 0.00 7 861.62
Phi(sex+tech) p
(g+tech+toe_lin)
875.80 26.27 0.00 7 861.62
Phi(sex) p(g+tech+toe_lin) 875.87 26.34 0.00 7 861.69
Phi(g+sex) p(.) 876.02 26.49 0.00 6 863.89
Appendix B. Continued.
Model AICcDAICcw k Deviance
Phi(g+sex+tech) p(.) 876.02 26.49 0.00 6 863.89
Phi(g+sex+toe_lin) p(.) 876.02 26.49 0.00 6 863.89
Phi(g+sex+tech+toe_lin) p(.) 876.02 26.49 0.00 6 863.89
Phi(sex+toe_lin) p
(g+tech+toe_lin)
876.90 27.37 0.00 8 860.67
Phi(g+sex) p(g) 877.02 27.49 0.00 9 858.73
Phi(g+sex+tech) p(g) 877.02 27.49 0.00 9 858.73
Phi(g+sex+toe_lin) p(g) 877.02 27.49 0.00 9 858.73
Phi(g+sex) p(g+tech) 877.02 27.49 0.00 9 858.73
Phi(g+sex+tech) p(g+tech) 877.02 27.49 0.00 9 858.73
Phi(g+sex+toe_lin) p
(g+tech)
877.02 27.49 0.00 9 858.73
Phi(g+sex) p(g+toe_lin) 877.02 27.49 0.00 9 858.73
Phi(g+sex+toe_lin) p
(g+toe_lin)
877.02 27.49 0.00 9 858.73
Phi(g+sex+tech+toe_lin) p
(g+toe_lin)
877.02 27.49 0.00 9 858.73
Phi(sex+tech+toe_lin) p
(g+tech+toe_lin)
877.08 27.55 0.00 9 858.80
Phi(g+sex+tech+toe_lin) p
(tech+toe_lin)
877.14 27.62 0.00 9 858.86
Phi(g+sex+tech+toe_lin) p
(g)
879.08 29.55 0.00 10 858.73
Phi(g+sex+tech+toe_lin) p
(g+tech)
879.08 29.55 0.00 10 858.73
Phi(g+sex+tech) p
(g+toe_lin)
879.08 29.55 0.00 10 858.73
Phi(g+sex) p
(g+tech+toe_lin)
879.08 29.55 0.00 10 858.73
Phi(g+sex+toe_lin) p
(g+tech+toe_lin)
879.08 29.55 0.00 10 858.73
Phi(g+sex+tech+toe_lin) p
(g+tech+toe_lin)
879.08 29.55 0.00 10 858.73
Phi(.) p(toe_lin) 880.41 30.88 0.00 3 874.37
Phi(g+sex+tech) p
(g+tech+toe_lin)
881.15 31.62 0.00 11 858.73
Phi(.) p(tech) 881.33 31.80 0.00 3 875.29
Phi(.) p(.) 881.96 32.43 0.00 2 877.94
Phi(tech) p(toe_lin) 882.33 32.80 0.00 4 874.27
Phi(tech) p(.) 882.36 32.83 0.00 3 876.32
Phi(.) p(tech+toe_lin) 882.36 32.83 0.00 4 874.30
Phi(toe_lin) p(toe_lin) 882.36 32.83 0.00 4 874.30
Phi(toe_lin) p(.) 882.95 33.42 0.00 3 876.91
Phi(tech+toe_lin) p(toe_lin) 883.13 33.60 0.00 5 873.04
Phi(tech) p(tech) 883.21 33.69 0.00 4 875.15
Phi(toe_lin) p(tech) 883.35 33.82 0.00 4 875.29
Phi(.) p(g) 883.77 34.24 0.00 5 873.68
Phi(.) p(g+tech) 883.77 34.24 0.00 5 873.68
Phi(.) p(g+toe_lin) 883.77 34.24 0.00 5 873.68
Phi(.) p(g+tech+toe_lin) 883.77 34.24 0.00 5 873.68
Phi(tech+toe_lin) p(tech) p
(toe_lin)
884.09 34.56 0.00 6 871.96
Phi(tech) p(tech+toe_lin) 884.21 34.69 0.00 5 874.12
Phi(tech+toe_lin) p(.) 884.24 34.71 0.00 4 876.18
Phi(toe_lin) p(tech+toe_lin) 884.33 34.80 0.00 5 874.24
Phi(g) p(toe_lin) 884.47 34.94 0.00 6 872.34
ª2014 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd. 1489
M. Guimar~
aes et al. Effects of Marking Techniques on Frogs
Appendix B. Continued.
Model AICcDAICcw k Deviance
Phi(g+tech) p(toe_lin) 884.47 34.94 0.00 6 872.34
Phi(g+toe_lin) p(toe_lin) 884.47 34.94 0.00 6 872.34
Phi(g+tech+toe_lin) p
(toe_lin)
884.47 34.94 0.00 6 872.34
Phi(g) p(.) 884.72 35.19 0.00 5 874.63
Phi(g+tech) p(.) 884.72 35.19 0.00 5 874.63
Phi(g+toe_lin) p(.) 884.72 35.19 0.00 5 874.63
Phi(tech+toe_lin) p(tech) 884.90 35.38 0.00 5 874.81
Phi(tech+toe_lin) p(g) 885.48 35.95 0.00 7 871.30
Phi(tech+toe_lin) p(g+tech) 885.48 35.95 0.00 7 871.30
Phi(tech+toe_lin) p
(g+toe_lin)
885.48 35.95 0.00 7 871.30
Phi(tech) p(g) 885.60 36.07 0.00 6 873.46
Phi(tech) p(g+tech) 885.60 36.07 0.00 6 873.46
Phi(tech) p(g+toe_lin) 885.60 36.07 0.00 6 873.46
Phi(g) p(tech+toe_lin) 885.68 36.15 0.00 7 871.50
Phi(g+tech) p(tech) p
(toe_lin)
885.68 36.15 0.00 7 871.50
Phi(g+toe_lin) p
(tech+toe_lin)
885.68 36.15 0.00 7 871.50
Phi(toe_lin) p(g) 885.77 36.24 0.00 6 873.64
Phi(toe_lin) p(g+tech) 885.77 36.24 0.00 6 873.64
Phi(toe_lin) p(g+toe_lin) 885.77 36.24 0.00 6 873.64
Phi(toe_lin) p
(g+tech+toe_lin)
885.77 36.24 0.00 6 873.64
Phi(g) p(tech) 885.77 36.24 0.00 6 873.64
Phi(g+tech) p(tech) 885.77 36.24 0.00 6 873.64
Phi(g+toe_lin) p(tech) 885.77 36.24 0.00 6 873.64
Phi(g+tech+toe_lin) p(tech) 885.77 36.24 0.00 6 873.64
p(g+tech+toe_lin) p(.) 886.76 37.23 0.00 6 874.63
Phi(g) p(g) 887.46 37.93 0.00 8 871.23
Phi(g+tech) p(g) 887.46 37.93 0.00 8 871.23
Phi(g+toe_lin) p(g) 887.46 37.93 0.00 8 871.23
Phi(g) p(g+tech) 887.46 37.93 0.00 8 871.23
Phi(g+toe_lin) p(g+tech) 887.46 37.93 0.00 8 871.23
Phi(g+tech+toe_lin) p
(g+tech)
887.46 37.93 0.00 8 871.23
Phi(g) p(g+toe_lin) 887.46 37.93 0.00 8 871.23
Phi(g+tech) p(g+toe_lin) 887.46 37.93 0.00 8 871.23
Appendix B. Continued.
Model AICcDAICcw k Deviance
Phi(tech+toe_lin) p
(g+tech+toe_lin)
887.53 38.00 0.00 8 871.30
Phi(tech) p(g+tech+toe_lin) 887.64 38.11 0.00 7 873.46
Phi(g+tech+toe_lin) p(g) 889.52 39.99 0.00 9 871.23
Phi(g+tech) p(g+tech) 889.52 39.99 0.00 9 871.23
Phi(g+toe_lin) p(g+toe_lin) 889.52 39.99 0.00 9 871.23
Phi(g) p(g+tech+toe_lin) 889.52 39.99 0.00 9 871.23
Phi(g+toe_lin) p
(g+tech+toe_lin)
889.52 39.99 0.00 9 871.23
p(g+tech+toe_lin) p
(g+tech+toe_lin)
889.52 39.99 0.00 9 871.23
p(g+tech+toe_lin) p
(tech+toe_lin)
889.79 40.26 0.00 9 871.50
p(g+tech+toe_lin) p
(g+toe_lin)
891.58 42.05 0.00 10 871.23
Phi(g+tech) p
(g+tech+toe_lin)
891.58 42.05 0.00 10 871.23
AICc, Akaike’s information criteria with small sample size correction;
DAICc, difference between top model and the current model; w
i
,
AICcweights; K, number of parameters; Deviance, difference of the
current model and the saturated model. Parameter abbreviations: (.),
constant; (sex), varies by sex; (g), varies by group; (toe_lin), varies with
linear trend effect; (tech), varies by marking technique.
Appendix C
Cumulative AICcweights for the covariates used for apparent survival
(Φ) and recapture probability (P).
Variables ΦP
sex 0.35 0.98
group 0.20 0.18
tech 0.40 0.38
toe_lin 0.35 0.48
1490 ª2014 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.
Effects of Marking Techniques on Frogs M. Guimar~
aes et al.