Zoologica Scripta - 2024 - Cerqueira - A New Antshrike Aves Thamnophilidae Endemic To The Caatinga and The Role of

Received: 18 December 2023
| Revised: 19 March 2024

| Accepted: 23 May 2024
DOI: 10.1111/zsc.12672
ORIGINAL ARTICLE
A new antshrike (Aves: Thamnophilidae) endemic to the

Caatinga and the role of climate oscillations and drainage
shift in shaping cryptic diversity of Neotropical seasonal
dry forests
Pablo Cerqueira1,2,3 | Gabriela R. Gonçalves1,2 | Tânia F. Quaresma3,4 |

Marcelo Silva4 | Mauro Pichorim5 | Alexandre Aleixo3,4
1
Programa de Pós-graduação Em
Zoologia, Museu Paraense Emílio Abstract
Goeldi/Universidade Federal do Pará, The Caatinga is the largest patch of Seasonal Dry Tropical Forest in the Neotropics,
Belém, Pará, Brazil
2
located in northeastern Brazil and characterized mainly by deciduous vegetation
Laboratório de Biogeografia da
Conservação e Macroecologia-
and extreme rainfall seasonality. It has historically been treated as a biologically
BIOMACRO, Instituto de Ciências impoverished domain, but recent studies uncovered new diversification patterns
Biológicas, Universidade Federal do and several new taxa of frogs, mammals, insects, and fishes. Here we employed a
Pará, Belém, Pará, Brazil
3
dense sampling regime to evaluate whether the São Francisco River (SFR) would
Instituto Tecnológico Vale, Belém,
Pará, Brazil have promoted genetic diversification and fixed phenotypic differences and how
4
Programa de Pós-graduação Em Quaternary climatic oscillations shaped distribution and population sizes in a
Biodiversidade e Evolução, Museu Caatinga endemic species, the Silvery-cheeked Antshrike (Sakesphoroides cris-
Paraense Emílio Goeldi, Belém, Pará,
Brazil
tatus). We adopted an integrative approach using multilocus genetic, plumage,
5
Departamento de Botânica e Zoologia, vocal data, and ecological niche modelling (ENM) to characterize evolutionary
Universidade Federal Do Rio Grande units and niche suitability in past scenarios. We recovered strong genetic struc-
Do Norte, Natal, Brazil
ture across the SFR that was congruent with plumage and vocal variation, reveal-
Correspondence ing a yet undescribed species named herein as Sakesphoroides niedeguidonae, sp.
A. Aleixo, Instituto Tecnológico nov. (urn:lsid:zoobank.org:act:2B9FC637-008A-4E9A-B92B-69ED7FE23823s).
Vale – ITV, Rua Boaventura da Silva,
The splitting time estimated between the newly described species and S. cristatus
955-66055-090, Belém, PA, Brazil.
Email: [email protected] is consistent with the establishment of the modern course of SFR, with a more
recent course shift apparently promoting the secondary contact between the two
Funding information
Coordenação de Aperfeiçoamento
species in the Raso da Catarina region. After their split, both species experienced
de Pessoal de Nível Superior, Grant/ increases in population sizes and range sizes at similar times during the Last
Award Number: 1537057 and 1537056; Glacial Maximum. We expect other Caatinga avian endemic lineages to show
Financiadora de Estudos e Projetos,
Grant/Award Number: 0118003100 similar patterns of genetic differentiation across the SFR that were enhanced by
Quaternary climatic oscillations.
Cover plate: Illustration of Sakesphoroides niedeguidonae (female MPEG 84680, male MPEG 84480). Illustration by Eduardo Brettas.
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.
© 2024 The Author(s). Zoologica Scripta published by John Wiley & Sons Ltd on behalf of Royal Swedish Academy of Sciences.
Zoologica Scripta. 2024;00:1–22. wileyonlinelibrary.com/journal/zsc | 1

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2 CERQUEIRA et al.
KEYWORDS
bioacoustics, biogeography, integrative taxonomy, niche modelling, São Francisco River
1 | I N T RO DU CT ION and trees (Caetano et al., 2008). Despite the recent in-
crease in the number of studies focusing on Caatinga di-
Seasonally Dry Tropical Forests (hereafter SDTFs) are versification patterns, additional studies are necessary to
tree-dominated ecosystems with a more or less continuous address the existence of congruent spatio-temporal histo-
canopy in which grasses are a minor element and thorny ries across taxa. In this sense, birds are one of the least
species are prominent. The vegetation is mostly decidu- sampled taxa by phylogenetic studies directed at Caatinga
ous during the dry season, with deciduousness increasing endemic lineages, in strong contrast to other domains
as rainfall declines in volume, although in the driest for- such as Amazonian and the Atlantic Forest (do Amaral
ests there is a marked increase in evergreen and succu- et al., 2013; Silva et al., 2019).
lent species (Mooney et al., 1995; Pennington et al., 2000). Herein, we present the first multi-character system-
The SDTFs has currently a discontinuous distribution atic study integrating genetic, morphological, vocal, and
with very heterogeneous patches, including formations as ecological data on a Caatinga endemic bird species, the
diverse as tall forest on moister sites to cactus scrub on Silvery-cheeked Antshrike Sakesphoroides cristatus (de
the driest. In South America this heterogeneity prompted Wied-Neuwied, 1831). Until recently, this species was
the recognition of distinct core areas throughout SDTFs placed in the genus Sakesphorus, but novel genetic and
such as the ‘Caatinga nucleus' in northeastern Brazil, the morphological data recovered it as non- monophyletic,
‘Misiones nucleus' in the Paraná-Paraguay basin, and the prompting the transfer of S. cristatus to the separate genus
‘Subandean Piedmont nucleus' in southwestern Bolivia Sakesphoroides (Bravo et al., 2021; Grantsau, 2010). The
and northwestern Argentina (Pennington et al., 2000, endemic Sakesphoroides cristatus is widely distributed in
2006; Prado, 2000; Prado & Gibbs, 1993; Ratter et al., 1978; the Caatinga domain, being currently regarded as mono-
Silva & Bates, 2002). typic. Based on our field observations and the examina-
The Caatinga is the largest area of SDTFs and en- tion of female specimens, which led to the first insights
compasses heterogeneous arid and semi-arid formations on plumage and song variation in Sakesphoroides, cou-
surrounded by more mesic formations, extending for pled with published evidence on vocal variation detected
~800 km2 and representing 11% of the Brazilian territory across the São Francisco River (Capelli et al., 2020), herein
(IBGE, 1985). The geomorphology of this region plays an we evaluated whether genetic differentiation and fixed
important role in its biogeographic history, with altitudes phenotypic differences would also be found on opposite
ranging from sea level in the north, an average 400–700 m banks of the São Francisco River (hereafter SFR). We also
through Bahia and Minas Gerais states in the central tested with the molecular data whether any differentia-
part, to over 1000 m on the slopes of the Espinhaço range tion across the SRF would correlate temporally with the
(Taylor & Zappi, 2004). In addition to geological processes establishment of its modern course. Finally, we carried
in the Paleogene−Neogene, Quaternary climatic oscilla- out ecological niche modelling and historical demography
tions also contributed to the high levels of observed ende- analyses to evaluate any congruences in distribution and
mism (Werneck, 2011). population sizes for S. cristatus across the Caatinga do-
The biogeographic history and diversification patterns main mediated by Quaternary climatic oscillations.
of SDTFs have received relatively less attention than those
of Neotropical rain forests (Amazonia and Atlantic Forest).
This is in part because Caatinga was historically treated 2 | METHODS
as a species impoverished domain with low endemism, a
concept falsified by recent studies (Mooney et al., 1995; 2.1 | Sampling
Pennington et al., 2000; Werneck, 2011). Many studies
have analysed Caatinga taxa under phylogenetic, phylo- We studied and analysed plumage and morphometric
geographic and genetic population dynamics approaches, variation in a total of 1079 Sakesphoroides cristatus speci-
including insects (Coutinho-Abreu et al., 2008; Franco mens (818 males, 261 females) including study skins
& Manfrin, 2013), lizards (Oliveira et al., 2015; Passoni (n = 92) and digital photographs available in online data-
et al., 2008; Siedchlag et al., 2010; Werneck et al., 2015), bases (n = 987; Table S1). To investigate song variation, we
frogs (Thomé et al., 2016), mammals (Faria et al., 2013; analysed a total of 115 different sound recordings (n = 95
Nascimento et al., 2011, 2013), fishes (Costa et al., 2018), loudsongs, n = 29 calls; more than one vocalization type
|
CERQUEIRA et al. 3
could be present in the same recording) available in on- location of females. We tested all measurements for ho-
line databases (Table S2). mogeneity of variance and normality before ANOVA, and
We sequenced 58 tissue samples from throughout S. performed statistical analyses using the package ‘stats' in
cristatus range (Table S3), 39 of them belonging to the left R software version 4.3.1 (R Core Team, 2023).
bank and 19 to the right bank of the SFR. As outgroups, we To support our interpretations of the degree of ge-
used sequences of 29 Thamnophilidae species closest re- netic and phenotypic differentiation in Sakesphoroides,
lated to the S. cristatus from GenBank based on topologies we inferred its dispersal ability in comparison to other
of phylogenies inferred by Brumfield and Edwards (2007) Thamnophilidae species occurring in dry open and humid
and Bravo et al. (2021; see Table S4 for species names and forest vegetation types. To do so, we used morphometric
GenBank accession numbers). data from the AVONET dataset (Tobias et al., 2022) to ex-
We gathered all available occurrence records of S. crista- tract measurements and indices related to wing size and
tus (n = 568) for Environmental Niche Modelling (ENM) shape variation (see Table S5 for the species and num-
from the Species Link database (http://splink.cria.org. ber of individuals analysed), which are regarded as good
br), Sound archive Xeno-canto (http://www.xeno-canto. proxies of flight efficiency and dispersal ability in birds
org), Wikiaves (http://www.wikiaves.com.br), Vertnet (Claramunt, 2021; Claramunt & Wright, 2017). We con-
(http://www.vertnet.org/index.html), Ebird (http://www. sidered the following wing attributes: (1) wing Length
ebird.org/), and museum specimens deposited at Museu (length from the carpal joint to the tip of longest primary
Paraense Emílio Goeldi (MPEG), Museu de Zoologia in millimetres); (2) Kipp's Distance (length from the tip
da Universidade de São Paulo (MZUSP), Universidade of the first secondary feather to the tip of the longest pri-
Federal de Pernambuco (UFPE), Universidade Federal mary in millimetres); (3) secondary1 (length from the
do Rio Grande do Norte (UFRN), Universidade Estadual carpal joint to the tip of the first secondary in millime-
de Feira de Santana (UEFS), and personal observations tres); and (4) Hand-Wing Index (100*DK/Lw, where DK is
gathered during fieldwork. We inspected all occurrences Kipp's distance and Lw is wing length). We used the Hand-
for errors in locality names and geographic coordinates Wing Index (HWI) and raw measurements reflecting
by comparison with the Brazilian ornithological gazetteer wing shape and size to perform a Principal Component
(Paynter Jr & Traylor Jr., 1991). Analysis (PCA) to determine whether these variables can
explain dispersal ability variation between Sakesphoroides
and other antbird species with inferred low dispersal abil-
2.2 | Plumage and ity. We further tested for significant differences in mean
morphometric analyses HWI between Sakesphoroides and other antbird species as
a direct comparison of dispersal capacity, also considering
To document geographic variation in plumage patterns, different taxonomic and ecological groupings (i.e. genera
we analysed study skins and digital photographs. We used and environment type, see Table S5). We checked for nor-
characters with discrete variation such as colours of the mality and homogeneity of variance prior to a Kruskal–
crown, back, throat, breast, tail, and presence of barring Wallis test and a pairwise Wilcox test (Mann–Whitney) if
and streaks. All colour standardizations were based on differences in HWI existed. We performed all statistical
Smithe (1975). We studied the type series of S. cristatus analyses using the package ‘stats' in R software version
through high quality digital photos, which include an 4.3.1 (R Core Team, 2023).
adult male (AMNH 6819), an adult female (AMNH 6820)
and a juvenile female (AMNH 6821) held at the American
Museum of Natural History, New York, USA. 2.3 | Vocal analyses
We used a digital calliper (nearest 0.1 mm) to measure
the following characters in study skins of 24 females and We used vocalizations to explore putative differentiation
54 males (Table S1): (1) length of flattened wing (LW), (2) across the SFR. Initially, we searched for qualitative char-
tarsus length (TarL), (3) tail length (TaiL), (4) bill length acters by visualizing spectrograms of loudsongs, which
from the skull insertion (BL1), (5) bill length from anterior are characterized as short series of ca. 10 notes, with the
edge of nostrils (BL2), (6) bill width at anterior edge of initial notes being sharp and strident, followed by a se-
nostrils (BW), and (7) bill depth at anterior edge of nos- ries of raspy notes (Zimmer et al., 2003). We could not
trils (BD). We tested for significant differences in these assign homologies among calls due to the small number
measurements using ANOVA with a Tukey test for a pos- of recordings across different locations in our sample and
teriori comparison using a dataset composed of males and the high degree of variation of this vocal type in differ-
females. Given their overall plumage conservatism, males ent stress contexts (i.e. playback, mate contact, presence
were assigned to different groups following the geographic of predators).
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4 CERQUEIRA et al.
All sound recordings were standardized before visu- included an initial cycle of 5 min at 94°C, followed by 35 cy-
alization. We converted all files to ‘.WAV’ format, with cles of 1 min at 94°C, 1 min at 57°C for Cytb and 55°C for
a sampling rate of 44,100 Hz and bit depth of 16 bits in G3PDH (annealing temperature), and 1 min at 72°C, with
the mono pattern. Spectrograms were generated and an- a final extension of 5 min at 72°C. For the ND2 fragment,
alysed on software Raven Pro 1.5 (Cornell Laboratory of the PCR reaction consisted of an initial cycle of 5 min at
Ornithology) with a Hamming window type, window size 94°C, followed by 35 cycles of 30 s at 94°C, 1 min at 55°C
of 512 samples, a time grid 90% overlap, and DFT size of (annealing temperature), and 30 s at 72°C, with a final ex-
2048 samples. We analysed the syntax and note structure tension of 5 min at 72°C. An aliquot of each amplification
qualitatively through a blind inspection and grouping was loaded in 1% agarose gel to check for the amplifica-
printed sonograms, followed by an assessment of whether tion products. Then, the amplified products were purified
the groupings matched the patterns under study, follow- using PEG-8000 (Polietilenoglicol-8000) to remove PCR
ing the procedure in Carneiro et al. (2012). residues. We sequenced the amplicons with a Big Dye
We also measured some parameters based on spectro- Terminator Cycle Sequencing Standard Version 3.1 kits
grams and oscillograms: number of sharp notes (NN1), in an automated sequencer model ABI 3130 (Applied
duration of all sharp notes including the intervals be- Biosystems). Sequences generated in this study were de-
tween notes in seconds (s; DS1), pace of sharp notes sec- posited in GenBank (www.ncbi.nlm.nih.gov; Accession
tion (dividing NN1 by DS1; Pace1), highest and lowest Numbers PP842021–PP842121).
frequency of all sharp notes in hertz (Hz; HighF1 and
LowF1), delta frequency of all sharp notes in hertz (differ-
ence between HighF1 and LowF1; DF1), peak frequency 2.5 | Phylogenetic analyses and
of all sharp notes in hertz (the frequency at which maxi- genetic distances
mum power occurs within the selection; PeakF1), number
of raspy notes (NN2), duration of all raspy notes includ- We edited and inspected sequences for any insertions,
ing the intervals between notes in seconds (DS2), pace of deletions and stop codons, using the software BioEdit
raspy notes section (dividing NN2 by DS2; Pace2), peak 7.1.3 (Hall, 1999). The alignment was performed using
frequency of all raspy notes in hertz (PeakF2), number of ClustalW algorithm implemented in the same software
notes along the entire song (NN), duration of total song BioEdit 7.1.3. Heterozygous sites of nuclear markers were
including the intervals between notes in seconds (DTS), translated following the IUPAC code.
pace of total song (dividing NN by DTS; Pace), and peak We estimated the evolutionary model that best ex-
frequency along the entire song in hertz (PeakF). We in- plained the obtained sequences in jModelTest 2.1.4
vestigated significant differences in these measurements (Darriba et al., 2012), and subsequently used in a Bayesian
using an ANOVA with a Tukey test for a posteriori com- Inference (BI) to estimate phylogenetic relationships
parison. We tested all measurements for homogeneity of among the specimens sequenced with MrBayes 3.1.2
variance and normality before performing ANOVA, we (Ronquist & Huelsenbeck, 2003). Analyses of the con-
performed statistical analysis using the package ‘stats' in catenated dataset were partitioned by gene. Four par-
R software version 4.3.1 (R Core Team, 2023). allel simultaneous runs of 3 × 106 generations each
were carried out, sampling one tree every 1000 genera-
tions. We used the Software TRACER 1.5 (Rambaut &
2.4 | DNA extraction, Drummond, 2009) to verify consistency of results and to
amplification, and sequencing check whether Effective Sample Size (ESS) values were
greater than 200 (Drummond et al., 2007). Trees obtained
We extracted DNA samples from muscle tissues to se- before the Markov chain reached stable and convergent
quence two mitochondrial (NADH dehydrogenase subu- likelihood values were discarded as burn-in. To estimate
nit 2– ND2 and Cytochrome b – Cytb) and one nuclear genetic distances between and within lineages recov-
markers (Glyceraldehyde 3-Phosphate Dehydrogenase – ered in phylogenies we obtained uncorrected pairwise
G3PDH). Total DNA was extracted using standard proce- p-distances based on the mtDNA dataset using the soft-
dures with the phenol-chloroform technique (Sambrook ware MEGA 5.2 (Tamura et al., 2011). We also compared
et al., 1989). We performed amplifications using a the genetic divergence between Sakesphoroides lineages
Polymerase Chain Reaction (PCR) under the following with those observed between closely related/sister species
amplification profile: 12.5 μL of Master Mix 1x, 8.5 μL pairs of other Neotropical Suboscine passerines (mainly
of H2O, 1.0 μL (200 ng/μL) of each primer (forward and Thamnophilidae; see Table S7). This set of species include
reverse) for all genes (see Table S6) and 2.0 μL (20 ng) of both those whose ranges are limited by major Amazonian
genomic DNA. Procedures for genes Cytb and G3PDH rivers as well as some species living in sympatry or with
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CERQUEIRA et al. 5
allopatric or parapatric ranges not separated by a major parameters of the Markov chain every 10,000 generations,
river (Table S7). We used sequences of the ND2 marker until the ESS of all parameters reached scores of 200. We
obtained from this study and GenBank, and analysed set for the mtDNA dataset (Cytb and ND2) the mutation
them with the software MEGA 5.2 (Tamura et al., 2011) rate of 2.1% of nucleotide substitutions per million years
based on uncorrected pairwise p-distances. (Weir & Schluter, 2008), whereas that of the nuclear locus
(G3PDH) was estimated in comparison to that of the
mtDNA rate under relaxed clock and uncorrelated log-
2.6 | Coalescence-based species normal options and a Yule process as a speciation prior.
delimitation (BPP) Based on the obtained results, the first 1000 samples of
each run were discarded as burn-in. We estimated values
To test for interspecific limits and genetic differentia- of posterior probability and divergence times through a
tion within S. cristatus we used the software Bayesian majority rule consensus of the remaining samples.
Phylogenetics and Phylogeography (BPP; version 2.2).
BPP uses a Bayesian approach to generate probabilities of
speciation between closely related taxa, using multilocus 2.8 | Historical demography
sequence data and a phylogenetic hypothesis as a guide
tree (Rannala & Yang, 2003, 2013; Yang & Rannala, 2010). To investigate the historical demography of S. cristatus
We performed BPP analysis using both algorithms 0 lineages we used the Extended Bayesian Skyline Plot
and 1 with different fine-tuning parameters (ε for 0, and α (EBSP; Heled & Drummond, 2008) under a linear model
and m for 1) following the software's recommendations to in BEAST 2.4.7 (Bouckaert et al., 2014). EBSP estimates
verify consistency of results. To assess variation of popula- changes in effective population sizes over time under
tion size parameters, we simulated three scenarios of prior a multilocus approach by using the times of coalescent
combinations representing different ancestral population events among gene trees. We used this method to test
sizes and different ages of separation between lineages: (1) demographic predictions expected under the Refugia
large effective population size (Ne) and deep divergence Hypothesis of Pleistocene climatic oscillations and to
with gamma distribution priors for θ (population size) and evaluate for congruence between historical demography
τ (divergence time) – G(1, 10) and G(1, 10); (2) small Ne and range size changes scenarios predicted by ecological
and shallow divergences with gamma distribution priors niche modelling for each lineage during the driest peri-
for θ and τ – G(2, 2000) and G(2, 2000); and (3) large Ne ods. We ran the analysis for 1 × 107 generations under an
and shallow divergences, with gamma distribution priors uncorrelated lognormal molecular clock with a substitu-
for θ and τ – G(1, 10) and G(2, 2000) (Smith et al., 2013). tion rate of 2.1% per million years applied to the mtDNA
As a guide tree, we used the phylogenetic hypothesis gen- dataset (Weir & Schluter, 2008), using this rate to estimate
erated by BI described above, running Markov chains for comparatively that of the nuclear locus.
100,000 generations, and sampling every five generations.
BPP tests different topologies, collapsing nodes of the
guide tree and counting support values. Speciation proba- 2.9 | Ecological niche modelling (ENM)
bilities are estimated from the sum of the probabilities of
all models for speciation events at each node of the tree To estimate the climatic niche dynamics of the different
guide. Daughter lineages (terminal taxa) from nodes that S. cristatus lineages in the last 120 ka we used 19 biocli-
had speciation probabilities >0.95 under all three prior matic layers at 30 arc-seconds (~1 km2) for the following
scenarios were classified as species (Smith et al., 2013). temporal window scenarios (see Table S8 for details of bi-
oclimatic layers): (1) current climatic scenario available
from the WordClim database (http://www.worldclim.
2.7 | Molecular dating and species tree org/current, Hijmans et al., 2005); (2) Middle Holocene
(~6 ka) and Last Glacial Maximum (LGM, ~21 ka) ob-
To estimate the divergence time among S. cristatus line- tained from Ecoclimate (http://www.ecoclimate.org/ ,
ages revealed by the BI phylogeny estimates, we carried Lima-Ribeiro et al., 2015; Varela et al., 2015); and (3)
out a species tree approach using the multilocus dataset Last Interglacial (LIG, ~120 ka) data obtained following
(mt and nucDNA) in *BEAST (Heled & Drummond, 2010). procedures described in Otto-Bliesner et al. (2006). The
We also estimated a separate time tree using mtDNA in climatic data used was based on the same general cir-
the BEAST 2.4.7 package (Bouckaert et al., 2014). For each culation model (GCM) available for all paleoscenarios:
multilocus and mitochondrial time tree, we performed the Community Climate System Model (CCSM). We
two independent runs of 1 × 107 generations, sampling the performed a Pearson's correlation test for all bioclimatic
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6 CERQUEIRA et al.
variables and removed the most highly correlated et al., 2012). We also performed an identity test to check
(r > 0.8) to avoid collinearity and model overfitting. the statistical significance of the obtained niche over-
Thereafter, we selected six bioclimatic variables to use lap metrics through 1000 replicates to generate a null
in models: Mean Diurnal Range (BIO2), Temperature distribution through similarity and equivalency tests. If
Seasonality (BIO4), Annual Precipitation (BIO12), the values of niche overlap metrics ranged within 95%
Precipitation Seasonality (BIO15), Precipitation of of the simulated null distribution, the null hypothesis
Warmest Quarter (BIO18) and Precipitation of Coldest (niche overlap) cannot be rejected. We carried out the
Quarter (BIO19). Our occurrence data was partitioned analysis using a set of packages (‘ecospat’, ‘rgeos', ‘ras-
following the ‘checkerboard’ method (Muscarella ter’, ‘tidyverse’ and ‘geodata’) in R software version 4.3.1
et al., 2014; Valavi et al., 2018), whereby the occurrence (R Core Team, 2023).
data is divided in geographic grids to maximize the envi-
ronmental independence and similarity among subsets.
We used a ROC's threshold to cut the suitability matri- 3 | RESULTS
ces modelled, which equalizes the omission and com-
mission errors (Barbet-Massin et al., 2012). 3.1 | Plumage and morphometric
We evaluated the ENMs using the True Skill Statistic analyses
(TSS; Allouche et al., 2006), a threshold dependent
method ranging from −1 to 1. Negative and posi- Of the 1079 individuals (skins and photographs) analysed,
tive values close to zero are not better than expected we could not identify any diagnostic plumage differences
by chance; values between 0.6 and 0.8 denote rea- among males (n = 818), but two main plumage patterns in
sonable fit models, whereas good fit models are indi- females (n = 261) were diagnosed mostly across the SFR
cated by values greater than 0.8, with 1 denoting best (Figure 1).
model fit (Allouche et al., 2006; Coetzee et al., 2009). The first pattern (hereafter plumage pattern 1) is char-
We used a combined result of the following four dif- acterized by females with tails showing large black and
ferent algorithms to model the species distribution and white bands and some ‘rufous-chestnut’ only on the edges
obtain a more reliable estimate of the environmen- of rectrices and near the white bands, in addition to an
tal niche (Rocchini et al., 2011): Generalized Linear Amber crest and olive brown back. In contrast, females
Model (GLM), Maximum Entropy (Maxent), Support sharing the second pattern (hereafter plumage pattern
Vector Machines (SVM) and Random Forest (RF; Guo 2) have entirely ‘rufous-chestnut’ tails, with weak dark
et al., 2005, Phillips et al., 2006, Prasad et al., 2006, brown marks, plus Chestnut crests and Cinnamon-Brown
Schölkopf et al., 2001, Tax & Duin, 2004). backs (Table 1; Figure S1).
The final model for current time and those projected We could not detect sexual dimorphism in S. crista-
for each paleoclimate scenario were obtained using a con- tus for the morphometric characters measured accord-
sensus model for each S. cristatus lineage. We considered ing to ANOVA tests. However, when both female and
only results with TSS values higher than 0.6 across the male specimens belonging to the two distinct groups
four algorithms used, presenting maps with the suitabil- defined based on the plumage of females are contrasted,
ity mean of the estimated environmental niche, named tail length (TaiL) and bill length from cranium insertion
‘ensemble’. We performed all procedures and model fit- (BL1) were the only morphometric characters that var-
ting described above using the package ‘ENMTML’ (de ied significantly (p < .05; Table 2). Despite these statisti-
Andrade et al., 2020) in R software version 4.3.1 (R Core cal differences neither measurement can be regarded as
Team, 2023), available at GitHub (https://github.com/ diagnostic between the two groups due to broad overlap
andrefaa/ENMTML). (Table 2).
To test for niche divergence, we performed a Based on wing size and shape attributes, we found
niche overlap analysis between the two recovered no differences in inferred dispersal ability between
Sakesphoroides lineages to verify whether climate has Sakesphoroides and other antbird species from both dry
influenced their Grinnellian niches (Soberón, 2007). We and wet environments (Figure S2), including species with
used the same 19 bioclimatic variables and occurrence recognized low dispersal features and genetic differentia-
data from the ENM analysis and performed a calibrated tion across Amazonian rivers (Figure S2). The PCA anal-
PCA (PCA-env) on the environmental space of both ysis explained 99.8% of total variance in the first two axes
lineages, including all occurrences. We calculated the (PC1: 65.28%, PC2: 34.51%), with Sakesphoroides having a
niche overlap using Schoener's D similarity metric rang- great overlap in space with other species from both dry and
ing from 0 (no overlap) to 1 (total overlap; Broennimann wet environments (Figure S2a–c). While variables related
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CERQUEIRA et al. 7
F I G U R E 1 Geographic distribution of diagnostic plumage characters in Sakesphoroides cristatus females. Green and blue circles
represent diagnostic patterns numbered 1 and 2, respectively, recovered from plumage analyses using study skins and digital photographs
(see text for details). The dark-blue line represents the modern course of the São Francisco River (SFR), with the red dashed lines
representing the estimated position of abandoned meanders of a Late Pleistocene paleocourse of the SFR. The colour gradient represents
altitudinal variation. Credit photos: Rocílio Ribeiro Rocha (pattern 1) and Oberdan Nunes (pattern 2).
T A B L E 1 Diagnostic body-part colour

Pattern 1 Pattern 2
comparisons of female skins belonging to
two distinct plumage patterns recovered Crown Amber (Colour 36) Chestnut (Colour 32)
in Sakesphoroides cristatus. Colour Back Olive brown (Colour 28) Cinnamon-Brown (Colour 33)
nomenclature follows Smithe (1975). Throat Pale horn Colour (Colour 92) Pale horn Colour (Colour 92)
Breast Buff (Colour 124) Buff (Colour 124)
Belly Buff (Colour 124) Buff (Colour 124)
Tail Jet Black (Colour 89), White and Amber (Colour 36) and dark
Amber (Colour 36) bands Amber
to wing size were positively correlated with PC1, the Hand- 3.2 | Vocal analyses
wing index was highly and negatively correlated with PC2
(Table S9). Specimens of Sakesphoroides clustered on Based on spectrograms of 95 sound recordings classified
the top left corner of the PCA graph and, therefore, were as loudsongs, we also detected two diagnosable vocal pat-
characterized by smaller wing size and Hand-wing Index terns roughly separated by the SFR in both males and
values, typically associated with species of lower dispersal females, consistent with differences detected in female
capacity. To reinforce this pattern, we found a significant plumage patterns (Figure 2).
difference in Hand-wing Index between taxa from distinct No sexual dimorphism was identified in spectrograms
dry and wet habitats (p < .05; Figure S2f), with dry habitat of recordings relative to sound structure and note shape.
species (which includes Sakesphoroides) showing signifi- The first diagnostic loudsong type (hereafter vocal pattern
cantly smaller Hand-wing Index values than wet habitat 1), is characterized by an initial series of sharp notes with
species (Figure S2d,e). frequency modulation (rapid increase and decrease, like an
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8 CERQUEIRA et al.
T A B L E 2 Measurements of
Pattern 1 Pattern 2
morphometric characters and statistical
(n = 59) (n = 19) comparisons among male and female
specimens of Sakesphoroides cristatus
Charactersa Mean SD Min-Max Mean SD Min–Max
belonging to two distinct plumage groups
LW 66.95 (a) 2.06 63–72 67.79 (a) 1.65 64–71 diagnosed only in females (see text for
TarL 25.28 (a) 0.93 23.46–26.85 25.32 (a) 1.04 23.69–27.33 details).
TaiL 57.03 (b) 4.04 48–68 61.74 (a) 2.86 56–67

BL1 17.75 (a) 0.73 15.14–19.23 16.87 (b) 0.9 14.63–18.59
BL2 10.09 (a) 0.53 9.08–11.81 10.02 (a) 0.39 9.09–10.67
BW 4.08 (a) 0.38 3.4–4.95 4.26 (a) 0.32 3.61–4.75
BD 5.18 (a) 0.31 4.71–6.08 5.29 (a) 0.28 4.77–5.80
Note: Males were assigned to each group based on the geographic location of females (Figure 1). All
measurements are in millimetres (mm), with reported sample sizes (n), mean plus standard deviation
(±SD), followed by minimum-maximum values. Mean values followed by different letters in parenthesis
on the same line differ significantly by p < .05 (in bold).
a
Morphometric characters: BD, bill depth at anterior edge of nostrils; BL1, bill length from cranium
insertion; BL2, bill length from anterior edge of nostrils; BW, bill width at anterior edge of nostrils; LW,
length of flattened wing; TaiL, tail length; TarL, tarsus length.
F I G U R E 2 Geographic distribution of diagnostic loudsong types of Sakesphoroides cristatus recovered during vocal analyses. Green and
blue triangles represent the distribution of patterns 1 and 2, respectively. (a) Spectrogram of type 1 loudsong showing the sharp notes with
ascending–descending frequency modulation (inverted ‘U' shape), accession number: XC320806; see detailed views of both sharp and raspy
notes under the spectrogram; (b) Spectrogram of type 2 loudsong showing the sharp notes with ascending–descending-stable-descending
frequency modulation (turned ‘S' shape), accession number: XC229164; see under the spectrogram detailed views of both sharp and raspy
notes. The dark-blue line represents the current course of the São Francisco River (SFR), with red dashed lines representing the estimated
position of abandoned meanders of a late Pleistocene paleocourse of the SFR. The colour gradient represents altitudinal variation.
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CERQUEIRA et al. 9
inverted ‘U’ in shape), followed by a short series of final raspy statistical support that Sakesphoroides cristatus is not
notes (Figure 2). The second loudsong type recovered (here- related to other Sakesphorus species, consistent with
after vocal pattern 2), has the initial series of sharp notes the topologies in Brumfield and Edwards (2007) and
with a different type of frequency modulation (i.e. rapid in- Bravo et al. (2021). Despite the overall uncertainty with
crease, decrease, short duration stable and decrease again, in respect to the true S. cristatus outgroup, our phylog-
a sigmoid shape), and short final raspy notes (Figure 2). eny estimates recovered this species as monophyletic
Comparisons using ANOVA test revealed seven vocal (Figure 3).
characters with statistically significant differences be- Within the monophyletic S. cristatus, two statis-
tween loudsong patterns 1 and 2: number of notes in total tically well supported independent genetic groups
song (NN), duration of total song (DTS), pace of total song were recovered roughly replacing each other across
(PTS), number of sharp notes (NN1), duration of all sharp the SFR (Figure 3; Figure S3), with clade 1 found for
notes (DS1), highest frequency of all sharp notes (HF1), the most part west and north of the river and clade 2
and delta frequency of all sharp notes (DF1) (Table 3). mostly east and south of it. Both clades are in close
Despite these statistical differences, none of the vocal and direct contact on the west bank of the mid-
characters above can be used as diagnostic between the dle SFR in Western Bahia. The uncorrected genetic
two vocal types due to overlap (Table 3). In summary, distance between clades 1 and 2 is 1.8% (standard
loudsong pattern 1 is significantly longer, yet slower and error ± 0.3%), whereas that within each clade is 0.2%
lower pitched than loudsong pattern 2. (standard error ± 0.1%). This average pairwise genetic
distance is similar to those found between some pairs
of closely related ‘biological’ species of antbirds such
3.3 | Phylogenetic analyses as Rhegmatorhina berlepschi/Rhegmatorhina hoff-
mannsi (1%) and Herpsilochmus atricapillus/Herp-
Our results based on BI phylogenies of the combined silochmus pileatus (1.7%; see Table S7 for complete
mtDNA and nuclear datasets recovered with high comparison).
TABLE 3 Variation in loudsong characters and statistical comparisons between two song types in Sakesphoroides cristatus.
Pattern 1 Pattern 2
(n = 19) (n = 30)
Measurementa Mean SD Min–Max Mean SD Min–Max

NN 12.74 (a) 1.33 10–15 11.1 (b) 1.42 08–14
DTS 3.04 (a) 0.39 2.35–3.56 2.47 (b) 0.29 1.76–3.06
Pace 4.21 (b) 0.37 3.65–5.09 4.51 (a) 0.39 3.61–5.42
PeakF 1707.94 (a) 97.38 1485.8–1938.0 1669.55 (a) 117.35 1442.7–1894.9
NN1 10 (a) 1.10 08–12 8.8 (b) 1.54 06–13
DS1 2.37 (a) 0.34 1.69–2.87 1.95 (b) 0.28 1.37–2.52
Pace1 4.25 (a) 0.45 3.58–5.20 4.51 (a) 0.47 3.64–5.62
HighF1 2186.44 (b) 210.78 1817.2–2626.0 2380.9 (a) 188.74 1964.4–2778.0
LowF1 576.12 (a) 141.29 368.5–934.2 553.11 (a) 125.85 341.9–788.9
DF1 1610.32 (b) 216.54 1110.0–1975.0 1834.84 (a) 232.75 1175.6–2243.5
PeakF1 1707.94 (a) 97.38 1485.8–1938.0 1662.37 (a) 119.20 1442.7–1894.9
NN2 2.74 (a) 0.93 1–4 2.3 (a) 0.84 1–5
DS2 0.6 (a) 0.22 0.22–1.06 0.6 (a) 0.76 0.14–4.51
Pace2 4.69 (a) 0.92 2.66–6.04 5.07 (a) 1.23 0.22–7.41
PeakF2 1712.46 (a) 198.96 1162.8–1981.1 1689.65 (a) 107.85 1507.3–1938.0
Note: Frequency is presented in hertz (Hz) and duration variables in seconds (s). Pace of notes per second is the total number of notes counted within a
particular time interval. Sample sizes (n), mean ± SD, followed by minimum-maximum values are presented for each character. For each character, any means
in bold followed by different letters in parenthesis depict significant differences at p < .05.
a
Measurement keys: DF1, delta frequency of sharp note section (HighF1 − LowF1); DS1, duration of all sharp notes including interval notes; DS2, duration of
all raspy notes including interval notes; DTS, duration of total song; HighF1, highest frequency of sharp note section; LowF1, lowest frequency of sharp note
section; NN, number of notes along the entire song; NN1, number of sharp notes; NN2, number of raspy notes; Pace, pace of total song; Pace1, pace of sharp
notes section; Pace2, pace of raspy notes section; PeakF, peak frequency along the whole song; PeakF1, peak frequency of sharp notes section; PeakF2, peak
frequency of raspy notes section.
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10 CERQUEIRA et al.
3.4 | Topology tests and species cristatus (clade 1 and 2) occurred during the Pleistocene be-
delimitation (BPP) tween 1.5 and 0.85 Mya (mean 1.2 Mya; Figure 3). In con-
trast, the splitting date estimated based on the multilocus
We recovered similar results using both algorithms 0 and dataset (species tree) for the two clades of S. cristatus was
1, with similar speciation probabilities in two independ- younger, but also centred in the Pleistocene, between 1.3
ent runs, indicating convergence of MCMC chains. Here and 0.35 Mya (mean 0.85; Figure 3). Because the confidence
we report results from algorithm 1 for three simulated intervals of splitting times estimated based on both datasets
evolutionary scenarios. overlapped broadly, and the multilocus approach provides
The BPP analysis strongly supported splitting S. cri- more robust estimates, we based our discussions mainly on
status in two deeply coalesced lineages for the pool of the dates obtained with the multilocus species tree.
molecular markers analysed, as indicated by high spe-
ciation probabilities (PP = 1.0; Figure 3) recovered across
all three evolutionary scenarios, a pattern consistent with 3.6 | Historical demography,
their recognition as separate species or highly structured current ecological niche model and
populations. Paleodistribution models
Extended Bayesian Skyline Plots (EBSPs) indicated a post-

3.5 | Molecular dating LGM increase in the effective population sizes of both lin-
eages of S. cristatus separated by the SFR (Figure 4). The
The molecular dating obtained with the mtDNA dataset only estimated expansion in population size of clade 1 (occur-
indicated that the split between S. cristatus and its closest ring west of the SFR) started about 22,000 years ago (22
relative among the sampled outgroups (Dysithamnus punc- kya) and was greater than that recovered for clade 2 (oc-
ticeps) took place during Late Miocene between 7 and 5.5 curring mostly east of the SFR), which began about 26 kya.
Mya (mean 6.295 Mya; Figure 3). According to the mtDNA The Ecological Niche Models (ENMs) estimated for the
chronogram, the divergence between two main clades of S. current and past times were obtained using six bioclimatic
F I G U R E 3 Phylogenetic hypotheses estimated from multilocus (species tree) and mtDNA (Bayesian chronogram) datasets. The larger
tree contains the topology recovered from the mtDNA dataset, with values above branches indicating mean estimated divergence dates for
each node and posterior probabilities shown as circles with different colours (black circles, PP≥0.99 and white circles, PP = 0.5). The smaller
species tree was inferred from the multilocus dataset, with values above branches representing mean estimated splitting times and values
below branches denoting speciation probabilities for each evolutionary scenario tested with BPP. Orange bars indicate 95% highest posterior
densities of divergence dates. The map on the left side shows the distribution of samples used in all analyses, with green representing clade
1 and blue representing clade 2 specimens. The black dashed square indicates the Western Bahia Region (WBR) and red dashed square Raso
da Catarina region. Clade 2 samples from Raso da Catarina region are in red in the main phylogenetic tree.
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CERQUEIRA et al. 11
F I G U R E 4 Demographic and range variation estimates through time for the two main clades of Sakesphoroides cristatus. Left: Extended
Bayesian Skyline Plots (EBSPs) showing demographic variation over time in thousands of years ago (kya) for each S. cristatus clade
recovered based on the multilocus dataset. Dashed black lines represent the median population size and the orange shaded area between
black lines represents the 95% confidence interval. Vertical red lines point to the time period when estimated population expansion began
in each clade. Map columns represent estimated Ecological Niche Models (ENMs) for the present, Mid-Holocene, Last Glacial Maximum
(LGM) and Last Interglacial (LIG) periods, respectively from the left to right. On frames showing the current distribution, grey circles
represent occurrence records used in analyses. Colours on maps ranging from cold (blue) to warm (red) indicate low and high suitability of
the ENMs, respectively. The São Francisco River is depicted on its current conformation as a blue line for reference in all maps, other rivers
in green and Caatinga boundaries under a black line.
variables (BIO2, BIO4, BIO12, BIO15, BIO18, BIO19; see both lineages have more similar niches (observed) than ex-
Table S8). All algorithms employed obtained TSS values pected by chance (simulated), at p > .05 (Figure S4).
equal to or higher than 0.6; therefore, we assumed that
they performed well to construct a consensus model. Of
the total occurrence data for S. cristatus as a whole, 153 and 3.7 | Taxonomy and species description
176 unique occurrences were selected to model the niches
of clade 1 and clade 2, respectively. For the LIG scenario, The BI and coalescence-based species tree analyses sup-
both S. cristatus lineages presented a very small suitable port the existence of two evolutionary independent line-
area (suitability>0.8; Figure 4). During the LGM, a strong ages in the otherwise monotypic Sakesphoroides cristatus
expansion of suitable areas for both lineages relative to the as currently treated. This is a novel finding for Caatinga
previous LIG period was detected (Figure 4). During the endemic birds, which historically were not thought to
Mid-Holocene there was a slight expansion of suitable areas have consistent phenotypic differentiation throughout the
relative to the previous scenario, whereas in the present (cur- Caatinga domain (Tomotani & Silveira, 2016).
rent scenario) a slightly smaller suitable area than during the The high statistical support for the split of
Mid-Holocene was estimated for both lineages of S. crista- Sakesphoroides cristatus in two separate lineages recov-
tus, congruent with the known limits of their modern distri- ered by the species delimitation analysis, combined with
butions (Figure 4). In the present scenario, ENMs estimated consistent phenotypic diagnoses of plumage and vocal
for clade 1 also predicted suitable niche conditions in most traits, together provide strong evidence of an evolutionary
of the clade 2 range, whereas clade 2 ENMs predicted fewer independent status for these lineages (Gill, 2014).
suitable niche conditions throughout the clade 1 distribu- The lone adult female of the type series of
tion (Figure 4). We recovered a small niche overlap between Sakesphoroides cristatus (AMNH 6820) collected by Wied
the two Sakesphoroides lineages (Schoener's D = 0.29). The exhibit the diagnostic plumage patterns characterizing fe-
identity test using similarity and equivalency indicated that males belonging to clade 2 (digital photographs of the type
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12 CERQUEIRA et al.
series held at the American Museum of Natural History: characters as described by Grantsau (2010), and its sister
AMNH 6819, 6820, 6821; see Figure S5). Wied's known relationship to Sakesphoroides cristatus.
itinerary bisected areas in the geographic range of clade 2, Males of Sakesphoroides niedeguidonae are similar in
never getting close to areas where clade 1 occurs (de Wied- plumage to Sakesphoroides cristatus, but females differ
Neuwied, 1940), reinforcing the notion that S. cristatus is from the former by distinct crown, back, and tail colours,
the taxon name associated with clade 2. The original notes (Table 1). Sakesphoroides niedeguidonae has an overall
on the type series of S. cristatus collected by Wied describe lighter (Amber) colour rather than Chestnut as S. crista-
the location as Campo Geral in Sertão of Bahia province tus. The back is olive brown in S. niedeguidonae, whereas
(Allen, 1889; de Wied-Neuwied, 1831). Throughout his that of S. cristatus is Cinnamon-Brown. The overall tail
itinerary, Wied provided detailed descriptions of the vege- colour in S. niedeguidonae is darker than in S. cristatus,
tation and general environments, passing through humid with the most conspicuous difference being the black and
Atlantic Forest, Caatinga, and ecotonal areas such as white barring, replaced in S. cristatus only by discreet dull
‘mata de cipó’, but never mentioned the exact place where brown and rufous bars and complete absence of white
the birds were collected. barring (Figure S1).
We suggest the area of Lajedo dos Beija- Flores, Voice: The loudsong of S. niedeguidonae is slightly similar
Municipality of Boa Nova, Bahia (14°19′ 51.2”S, in general pattern to that of S. cristatus, but differs conspic-
40°12′54.8”W) as a precise designated type locality for uously by the shape of the first notes. While the loudsong
Sakesphoroides cristatus. Wied visited the Boa Nova re- of S. niedeguidonae has initial notes with an ascending–de-
gion when moving from ‘General Quarter of Valo’ in the scending pattern of frequency modulation, looking visually
Bahia-Minas Gerais border to Salvador. Boa Nova is a like an inverted ‘U’ letter on sound spectrograms, that of
well-known ornithological location (Paynter Jr & Traylor S. cristatus has initial ascending notes reaching quickly a
Jr., 1991), maintaining Caatinga forests, with S. cristatus high frequency (~2300 Hz) and then descending to the
being still abundant in the local remnants of Caatinga and maximum amplitude and continuing to descend further to
‘mata de cipó’ in protected private areas and in the Boa a lower frequency (~580 Hz), which confers to it a sigmoid
Nova National Park. shape on spectrograms (Figure 2). Additionally, the loud-
Once we defined S. cristatus as the applicable name for song of S. niedeguidonae differs from that of S. cristatus by
clade 2 and no taxon name is available for clade 1 birds, a significantly greater number of notes, longer overall du-
herein we describe the latter group as a new species: ration, and slower pace in total song (see Table 3). Female
Sakesphoroides niedeguidonae, sp. nov. Cerqueira, recordings present noticeable higher pitched songs than
Gonçalves, Quaresma, Silva, Pichorim & Aleixo males (ML579640291; Macaulay Library of Natural Sounds,
Northern Silvery-cheeked Antshrike (English). Cornell Laboratory of Ornithology, Ithaca).
Choca-do-nordeste-de-cauda-barrada (Portuguese).
urn:lsid:zoobank.org:act:2B9FC637-008A-4E9A-B92B-
69ED7FE23823 3.7.3 | Description of holotype
See cover plate (Supplementary Material). Crown, forehead,

3.7.1 | Holotype and nape Amber Colour (36), forming a crest. Sides of head,
neck, and throat pale horn Colour (92). Back/mantle olive
MPEG 84680, field number PVC 275, adult female from Serra brown (28), presence of weak white interscapular patch.
Vermelha, surroundings of Serra da Capivara National Park, Rump and uppertail coverts olive brown, as in the back/man-
São Raimundo Nonato, Piauí (08°52′47.5”S, 42°43′32.8”W); tle. Breast, belly, and undertail coverts Buff (124), slightly
elevation c. 525 m; collected on 11 January 2019 by P. V. contrasting with upperparts. Primary and secondary feath-
Cerqueira, G. R. Gonçalves and A. Aleixo; study skin and ers olive brown colour (28), tertials olive brown with white
tissue prepared by P. V. Cerqueira (Tissue accession number on outer vane edge. Secondary coverts Dusky Brown colour
PVC275); audio recorded by P. V. Cerqueira (ML579640291; (19) with white and Cinnamon colour (123A) on distal por-
Macaulay Library of Natural Sounds, Cornell Laboratory of tion. Greater primary coverts Dusky Brown colour (19) with
Ornithology, Ithaca, New York). white and Cinnamon colour (123A) on outer vane edge and
distal portion of feathers. Central rectrices darker than outer
rectrices, showing marginal black and white spots some-
3.7.2 | Diagnosis times washed Amber (36) close to the five bars formed by
white spots in both vanes. Soft parts in life: iris dark brown;
Morphology: The new species is assigned to the genus maxilla black; mandible with a grey proximal portion and a
Sakesphoroides based on its morphological diagnostic black distal portion; tarsus and feet dull bluish grey.
|
CERQUEIRA et al. 13
3.7.4 | Measurements of holotype at anterior edge of nostrils 4.21 mm; bill depth at anterior
edge of nostrils 5.74 mm; wing chords 68 mm (left) and
Exposed culmen from cranium insertion 17.83 mm; bill 67 mm (right); tail 59.11 mm; tarsus 26.91 mm (left) and
length from anterior edge of nostrils 9.85 mm; bill width at 27.29 mm (right); body mass 15.0 g; Skull 100% ossified;
anterior edge of nostrils 3.87 mm; bill depth at anterior edge of left testis 4.0 × 2.0 mm.
nostrils 4.83 mm; wing chords 64.42 mm (left) and 64.13 mm
(right); tail 54.8 mm; tarsus 24.18 mm (left) and 24.37 mm
(right); body mass 15.0 g; Skull 100% ossified; ovary 6 × 3 mm. 3.7.8 | Distribution
The new species is almost restricted to the left/northern

3.7.5 | Allotype bank of the São Francisco River, with one population oc-
curring on the right bank at ‘Raso da Catarina’ region in
MPEG 84480, field number PVC 272, adult male from Serra Bahia state (Figure 5). This population found on the right/
Vermelha, surroundings of Serra da Capivara National Park, southern bank of the São Francisco River could have been
São Raimundo Nonato, Piauí (08°52′47.5”S, 42°43′32.8”W); ‘transferred’ across the river due to paleocourse shifts, as
elevation c. 525 m; collected 09 January 2019 by P. V. discussed in detail below in discussion section.
Cerqueira, G. R. Gonçalves and A. Aleixo; study skin and
tissue prepared by P. V. Cerqueira (Tissue accession number
PVC272); audio recorded by P. V. Cerqueira (ML579595301; 3.7.9 | Specimens examined (skins)
Macaulay Library of Natural Sounds, Cornell Laboratory of
Ornithology, Ithaca, New York). See Table S1 for detailed information on the specimens
examined below.
Sakesphoroides niedeguidonae: Brazil; Piauí: 10 females
3.7.6 | Description of allotype (MPEG 68001, 75,501, 84,680 [holotype], PVC285, 84,481,
UFRN 410, 546, 550, 567, UFPE 4936), 10 males (MPEG
See cover plate (Supplementary Material). Crown, forehead, 68000, 68,002, 75,506, 76,052, 76,752, 76,753, 84,466, 84,480
and central portion of nape Jet Black Colour (89), forming [allotype], UFRN 376, 521); Ceará: 5 females (MPEG 82876,
a crest. Supercilium and lores Jet Black. Sides of head Pale 82,885, 82,886, UFPE 4988, 4998), 6 males (MPEG 72194,
Neutral Grey (86) with Jet Black just behind the eyes, and 82,890, 82,893, 82,894, UFPE 4953, 4993); Rio Grande do
black in the anterior portion of the malar region. Back/ Norte: 1 female (UFRN 167), 7 males (UFRN 160, 166,
mantle Raw Umber colour (23), with presence of a weak 252, 275, 296, 302, 321); Paraíba: 1 female (UFPE 4546);
white interscapular patch. Rump and uppertail coverts Raw Pernambuco: 10 females (UFPE 934, 943, 972, 1228, 1301,
Umber, as the back/mantle. Throat and breast black as crest 4162, 4748, 4749, 4750, 4751), 11 males (UFPE 213, 214, 936,
and lores; the black colour is restricted to the central por- 969, 1252, 4223, 4691, 4744, 4745, 4746, 4747); Bahia: 8 fe-
tion of breast, continuous from the throat and narrowing males (MPEG 83581, 83,590, 83,592, 83,852, UEFS DA220,
towards the belly. Belly and undertail coverts White, flanks DA336, DA443, UFPE 4844), 14 males (MPEG 83583,
Light Neutral Grey (85). Primary and secondary feathers 83,584, 83,586, 83,589, 83,591, 83,615, 83,616, UEFS DA675,
olive brown (28), tertials olive brown with white on the DA238, DA347, DA577, DA497, DA240, UFPE 4848).
edges of outer vanes. Marginal and lesser secondary coverts Sakesphoroides cristatus: Brazil; Bahia: 2 females
Jet Black with white distal portions. Median and greater (MPEG 83595, 83,620), 17 males (MPEG 83596, 83,597,
secondary coverts Jet Black with white distal portions. 83,598, 83,599, 83,602, 83,603, 83,611, 83,612, 83,613,
Greater primary coverts Jet Black with white on outer vane 83,617, 83,618, 83,619, UEFS DA028, DA039, DA835,
edges and distal portion of feathers. Some secondary and DA995, DA254).
primary coverts present Raw Umber colour, probably re-
lated to moult limit. Tail Jet Black with white spots on each
vane of rectrices. Soft parts in life: iris dark brown; maxilla 3.7.10 | Specimens examined (tape
black; mandible black; tarsus and feet dull bluish grey. recordings)
See Table S2 for detailed information of examined tape

3.7.7 | Measurements of allotype recordings.
Sakesphoroides niedeguidonae: Brazil; Piauí (3), Ceará
Exposed culmen from cranium insertion 17.65 mm; bill (18), Rio Grande do Norte (1), Paraíba (1), Pernambuco
length from anterior edge of nostrils 10.45 mm; bill width (7), Bahia (6).
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14 CERQUEIRA et al.
F I G U R E 5 Geographic distributions of Sakesphoroides niedeguidonae sp. nov. (green) and Sakesphoroides cristatus (blue). The current
São Francisco River course is depicted by a black line, whereas its inferred paleocourse is shown as a dashed red line.
Sakesphoroides cristatus: Brazil; Bahia (41), Minas Silvery-cheeked Antshrike). The Portuguese name (Choca-
Gerais (18). do-nordeste-de-cauda-barrada) highlights the main plum-
age feature in females distinguishing the new species from
S. cristatus (Choca-do-nordeste in Portuguese). Both names
3.7.11 | Etymology maintain in its composition the name used in S. cristatus for
an easy reference to their close phylogenetic relationship.
It is our pleasure to name this species in honour of Niède
Guidon, a Brazilian archaeologist who in 1970's explored
the largest and oldest concentration of prehistoric sites in 3.7.12 | Habitat, ecology, and behaviour
the Americas. Niède's efforts helped to create the Serra da
Capivara National Park in Piauí state, where her research Sakesphoroides niedeguidonae is restricted to Caatinga
found evidence of artefacts that have provoked a re- vegetations, occurring in woody deciduous forest, arid
evaluation of the traditional theories about human settle- lowland scrub Caatinga, coastal sandy scrub vegetation in
ment in the Americas. Moreover, she is a symbol of power Ceará and Rio Grande do Norte states and in enclaves of
and persistence in preserving the Caatinga environment ‘cerradão’ (forested savannas) in the Chapada do Araripe
not just for archaeological purposes but also considering region. Sometimes it occurs close to anthropogenic
biodiversity and local human communities, highlighting landscapes whenever natural ecosystems are available
the Caatinga as a singular place in the world, which has nearby, supporting a medium adaptative capacity status
inspired us every day. to human impacts like the sister S. cristatus (De Araujo &
The English name (Northern Silvery- cheeked Da Silva, 2017).
Antshrike) is a reference to its range in the Caatinga do- Commonly seen in pairs or small familiar groups. In
main and the sister taxon S. cristatus (now called Southern March 2018 during one of our collecting expeditions at
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CERQUEIRA et al. 15
Formosa do Rio Preto (western Bahia), we observed a fa- climate change exacerbate the impacts of main threats
miliar group with two adults (male and female) and two ju- in this region.
veniles with incomplete male plumage. It forages in pairs Sakesphoroides niedeguidonae is a common species in
or sometimes in mixed-species flocks in low and middle most of its distribution, occurring in different habitats in
strata of woodland and scrub vegetation, catching insects the Caatinga, including secondary growth vegetations but
in short flights between close branches, dropping to the the capacity to tolerate anthropogenic landscape without
ground where it scans for prey while tossing dry leaves. natural vegetation nearby seems to be very low (De Araujo
Ants, spiders, termites, beetles, and caterpillars are the & Da Silva, 2017). The extent of occurrence (EOO) is esti-
main food items in Sakesphoroides niedeguidonae, and we mated at 630,000 km2, not approaching the threshold for
expect a similar diet to that reported for S. cristatus, which a Vulnerable status under the IUCN range size criterion
includes Formicidae, Isoptera, and Coleoptera during dry (<20,000 km2). Although no data on population size is
season, with the proportion of Isoptera, Coleoptera and available, we expect a number greater than 10,000 mature
Lepidoptera (larvae) increasing and that of Formicidae individuals, so this species is best placed as Least Concern
decreasing during the rainy season (Buainain et al., 2016). (LC). However, new approaches using niche modelling for
Additional studies focused on feeding biology of S. nied- future climate scenarios support a significant reduction in
eguidonae are needed to corroborate similar patterns re- range size of S. cristatus (including S. niedeguidonae, as
ported for S. cristatus. treated before this study); applying IUCN criteria to these
Similar to most antshrikes, Sakesphoroides niedegui- future estimated range sizes can change its threat cate-
donae is a territorial species. During collecting expeditions gory to Near Threatened (NT) and even Vulnerable (VU)
at the type locality and in western Bahia, males were com- depending of the future global change scenario (i.e. RCP
monly observed on top of vegetation and exposed branches 4.5 and RCP 8.5, respectively; Gonçalves et al., 2023). To
showing a clear territory defence behaviour in response to improve extinction risk evaluations for this species, more
playback. Females in contrast were more secretive in low precise estimates of population sizes and a better under-
strata of dense vegetation, perching on horizontal and ver- standing of landscape fragmentation and climate change
tical branches when stimulated by playback or scanning effects on the species' range are needed.
for prey, with discrete tail movements. We located sev-
eral adjacent territories during linear transects distancing
at least 50 m from each other, with similar observations 4 | DISC USSION
made in the Serra Vermelha area (Santos et al., 2012), sug-
gesting small territories in S. niedeguidonae. 4.1 | The role of São Francisco River
Little is known about the breeding biology of driving bird evolution in the Caatinga
Sakesphoroides niedeguidonae. Lima et al. (2010) described domain
a nest in the Raso da Catarina region found in April (see
WA1109656 and WA1109655 at www.wikiaves.com.br) We confirmed that the SFR coincides for the most part
as an open cup c. 45 mm deep, 50 × 60 mm diameter and with a major phylogeographic break in Sakesphoroides,
50 mm in total height. Clutch size consisted of 2 eggs with essentially delimiting the ranges of two evolutionary
creamy colour variably blotched with vinaceous-brown distinct units recognized as species herein. Although
markings concentrated on the larger pole of eggshell river-
delimited ranges such as that documented for
(based on photographs provided in Lima et al., 2010). Eggs Sakesphoroides have been recovered for multiple humid
weighed 3 g and measured 22 × 17 mm. Both male and fe- forest bird lineages across major Amazonian rivers
males incubate the eggs (Lima et al., 2010). (Carneiro et al., 2012; Silva et al., 2019), this is the first
documented instance of a major river playing a role in
structuring bird lineages in a SDTF area. The average
3.7.13 | Conservation pairwise genetic divergence (p-distance) between S. crista-
tus and S. niedeguidonae (1.8 ± 0.3%), is even higher than
The Caatinga region has been strongly disturbed by those separating at least two other sister species pairs of
human activities and main threats are large-scale conver- Suboscine passerines whose ranges are separated by wide
sion of native vegetation areas for commercial agricul- Amazonian rivers: Rhegmatorhina berlepschi/hoffmannsi
ture, livestock farms, and continuous overexploitation of (1%; Ribas et al., 2018), and Hylopezus paraensis/whittakeri
native vegetation by firewood collecting and slash-and- (0.77%; Carneiro et al., 2012; see Table S7).
burn agriculture (Da Silva et al., 2017). Moreover, the The estimated divergence time for the separation be-
absence of a large protected area system connected by tween S. cristatus and S. niedeguidonae was most likely
corridors, plus the intensification of desertification and during the Middle Pleistocene (around 1 Mya), but
|
16 CERQUEIRA et al.
confidence intervals do not rule out a Late Pleistocene ancestral polymorphism due to incomplete lineage sorting
date (1.313–0.366 Mya), particularly that obtained with (Dias et al., 2018; Ferreira et al., 2018). We did not obtain
the coalescent species tree approach (Figure 3). During any evidence of actual gene flow analysing plumage and
the Middle Pleistocene, the SFR course became inter- vocal patterns, but either a former selective mitochon-
rupted to the north, where it connected to the Piauí and drial sweep or past/current gene flow might explain the
Parnaíba rivers, and turned into a lacustrine/palustrine observed mismatch between genetic and phenotypic char-
system (Mabesoone, 1994; Potter, 2003). According to the acters in S. niedeguidonae Raso da Catarina populations.
paleolacustrine hypothesis, during the Middle Pleistocene, Future studies should address the dynamics of this con-
the SFR became an endorheic drainage emptying into tact zone by increasing the number of molecular loci and
a fluvio-lacustrine environment in its middle portion individuals sampled.
between Remanso and Petrolina (Mabesoone, 1994; Both S. cristatus and S. niedeguidonae are also in
Rodrigues, 1986, 2006; Tricart, 1974). The changing pa- apparent contact in Western Bahia Region (WBR, see
leo-SFR course and the large swampy environment in the Figure S6 for detailed views), where S. cristatus has
Remanso – Petrolina area could have played a role in isolat- ‘crossed’ onto the left bank of the middle SFR probably
ing the ancestor of S. cristatus and S. niedeguidonae in two mediated by changes in river flow caused by erosion and
populations. Subsequently, during the Mindel glaciation sedimentation (Barreto, 1996; Tricart, 1974). This process
(~450 kya), the paleo-SFR course shifted radically south- created forsaken meanders that transferred populations
eastward into the Atlantic Ocean, eventually reaching its across riverbanks or even connected both banks in drier
current position after some extensive meandering south of periods during the establishment of a lacustrine system
the modern mouth, as revealed by several surfaces inter- downriver in the Middle Pleistocene (Barreto, 1996; see
preted as paleochannels (King, 1956; Mabesoone, 1994). also below). Our sampling from this particular area in-
The confidence intervals of our estimate for the splitting dicates congruency between molecular and phenotypic
times between both Sakesphoroides species match this pe- variation in both S. cristatus and S. niedeguidonae, in
riod of extensive changes in the SFR course and could be contrast to what we documented in the Raso da Catarina
related to their origin. region. Additionally, we found that S. cristatus and S.
Indeed, the configuration of the lower SFR Late niedeguidonae are not syntopic in WBR, but instead sep-
Pleistocene course south of its modern position is congru- arated by ca. 40 km across an area dominated by Cerrado
ent with the geographic variation in phenotypic characters and an anthropogenic landscape resembling a former ri-
documented between S. cristatus and S. niedeguidonae, parian vegetation. Cerrado /Caatinga ecotones are com-
such as loudsong and plumage, known to be import- mon in this region, and we could not find any suitable
ant features in mate recognition (Isler et al., 1998, 2007; habitat and individuals of both species on this narrow
Remsen Jr., 1984). Subsequent to its major shift southeast- Cerrado patch, concluding that it could work as an en-
ward, a second drainage shift event probably mediated by vironmental barrier preventing their direct contact and
erosional processes and climate change moved the SFR possibly gene flow. Nevertheless, occurrence of hybrids
lowermost course northward to its current position; evi- or introgression events in these regions cannot be com-
dence found in sediments suggests the estimated position pletely ruled out.
of this recent paleocourse between the modern courses of Recently, some studies have found patterns congru-
the Itapicuru and Vaza-Barris rivers (King, 1956). This led ent with genetic structure mediated by the SFR in mam-
to the establishment of a rather recent secondary contact mals, lizards, insects, and frogs (Bruschi et al., 2019;
zone between S. niedeguidonae populations from ‘Raso da Coutinho-Abreu et al., 2008; Faria et al., 2013; Werneck
Catarina’ region (which could have been passively trans- et al., 2015). The role of rivers as barriers and speciation
ferred from the left to the right bank of the SFR) and those drivers is better understood in the Amazon Forest, and
of S. cristatus, which were prevalent on the right bank the SFR represents a new system to study complex pat-
of the river (for similar pattern in other species see also terns of speciation with mechanisms including multiple
Werneck et al., 2015). This pattern of putative recent sec- changes in river courses, Quaternary climate change,
ondary contact of the two Sakesphoroides species is cor- and neo tectonism. In our comparisons using the Hand-
roborated by our molecular phylogeny (mainly based on wing index as a proxy for dispersal ability, we verified
the mitochondrial dataset, with the only nuclear gene se- that Sakesphoroides spp. are low vagility antbirds like
quenced providing inconclusive evidence), which groups other species living in the understory of Amazonian
specimens S. niedeguidonae from Raso da Catarina in the forest, which is congruent with their habitat selection
S. cristatus clade. The incongruence between genotype and behaviour patterns (e.g. small territories and move-
and phenotype can result either from mitochondrial in- ments through short flights in middle of dense low veg-
trogression due recent secondary contact or retention of etation). The genetic, phenotypic, and morphometric
|
CERQUEIRA et al. 17
evidence shown here support the hypothesis that the Oliveira et al., 2015; Thomé et al., 2016). Recently, Gehara
SFR establishment played a very important role in the et al. (2017) recovered a strong signal of synchronous pop-
diversification of Sakesphoroides by acting as a strong ulation expansion in the Late Pleistocene for reptiles and
barrier for dispersion, whose effects varied in time and amphibians with quite different ecological adaptations
different locations. and life history strategies, indicating a strong role of cli-
matic oscillations in promoting effective population size
changes after diversification events in the Caatinga fauna.
4.2 | The influence of quaternary Both Sakesphoroides species have also expanded at con-
climatic oscillations gruent times in the Late Pleistocene, as found in Gehara
et al. (2017).
We also evaluated the influence of Quaternary climatic Future studies should test these same spatio-temporal
oscillations in Sakesphoroides using paleodistribution patterns of expansion in other Caatinga birds to evaluate
modelling and inferences of past population sizes through for shared responses to climate change among Caatinga
time (EBSP). Our results are partially consistent with organisms.
predictions of the Refugia Hypothesis for Dry Forests
(Haffer, 1969; Haffer & Prance, 2001), supporting both de- ACKNOWLEDGEMENTS
mographic and paleodistribution range expansions during We thank the curators and curatorial assistants for al-
and shortly after the LGM (~22 kya), preceded by smaller lowing us the use of skins and tissues held in the fol-
population sizes and more restricted distribution ranges lowing institutions: Museu Paraense Emílio Goeldi
during LIG (~120 kya; Figure 4). The recovered pattern (MPEG), Universidade Federal de Pernambuco (UFPE),
of sympatry between both species in WBR was likely a Universidade Federal do Rio Grande do Norte (UFRN),
secondary contact as supported by paleoenvironmen- Universidade Estadual de Feira de Santana (UEFS)
tal evidence indicating a reduction in water levels of the and Museu de Zoologia da Universidade de São Paulo
middle SFR resulting in increasing sand deposition dur- (MZUSP). We thanks Paul R. Sweet and Thomas J.
ing hot and dry periods that could have promoted con- Trombone from the American Museum of Natural
nections and a stepping-stone dispersion across opposite History (AMNH) for providing high quality photo-
banks (Barreto, 1996). So, we hypothesized that climatic graphs of the type specimens. Instituto Chico Mendes
oscillations after diversification events mediated by the de Conservação da Biodiversidade (ICMBio) issued col-
SFR could have shifted the ranges of both Sakesphoroides lecting permits (#52349-3). We are especially grateful
species, possibly promoting secondary contact by expan- to Gustavo Gonsioroski, Ciro Albano, Cristine Prates,
sion during drier periods and isolation again by retraction Chico Rasta, Firmino Filho, André Adeodato and
during mesic periods, reinforcing their genetic and phe- ‘Seu' Carmelo for all support during our fieldwork. To
notypic differentiation in areas of sympatry, as in WBR Fernando Pacheco for the valuable information and rich
(Bruschi et al., 2019). discussion about Wied's itinerary and Gustavo Bravo
The Quaternary climatic oscillations promoted niche for discussions and sharing unpublished data during
evolution in some species, mainly in organism with fast the early stages of this project. Thanks to Marcos Pérsio
life cycles like bees (Silva et al., 2014). But we did not find for the valuable material collected in the Caatinga dur-
evidence for Grinnellian niche evolution or divergence ing many years and all support before and during the
between both species of Sakesphoroides, the small niche project and to Andy and Gill Swash for discussions and
overlap recovered was not statistically significant indicat- suggestion of English names. We also thank field re-
ing that both species cannot be differentiated by occupied cordists and birders who shared sound recordings and
niche. Speciation is not commonly associated with an eco- photographs in the following citizen science platforms:
logical innovation or niche evolution and appears to be WikiAves, Xenocanto and Macaulay Library (Cornell
more noticeable farther into the past as in genera or family University). PC warmly thanks all bird watchers that
levels (Peterson, 2011; Peterson et al., 1999). acquired his birding tours services making possible the
The Refugia Hypothesis has been tested in the realization of fieldwork, as they indirectly funded this
Neotropical region mostly using humid forest taxa research. Laboratory work related to this paper was
(Batalha-Filho et al., 2013; Batalha-Filho & Miyaki, 2016; funded by a FINEp grant (# 0118003100). PC and GRG
Menezes et al., 2017; Santos et al., 2018; Solomon were supported by Coordenação de Aperfeiçoamento
et al., 2008). Studies focused on Caatinga taxa found ev- de Pessoal de Nível Superior – Brasil (CAPES) Doctoral
idence of population and/or range expansions as we did fellowships (#1537057 and #1537056, respectively). AA
for Sakesphoroides (Caetano et al., 2008; Coutinho-Abreu is supported by a productivity fellowship from CNPq
et al., 2008; Faria et al., 2013; Nascimento et al., 2013; (309243/2023-8).
|
18 CERQUEIRA et al.
spatial environmental data. Global Ecology and Biogeography,

ORCID
21, 481–497. https://doi.org/10.1111/j.1466-8238.2011.00698.x
Pablo Cerqueira https://orcid.org/0000-0002-7311-6229 Brumfield, R. T., & Edwards, S. V. (2007). Evolution into and out of
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Tânia F. Quaresma https://orcid. doi.org/10.1111/j.1558-5646.2007.00039.x
org/0000-0001-8192-064X Bruschi, D. P., Peres, E. A., Lourenço, L. B., De Macedo Bartoleti, L.
Marcelo Silva https://orcid.org/0000-0003-0642-8246 F., Sobral-Souza, T., & Recco-Pimentel, S. M. (2019). Signature
of the paleocourse changes in the São Francisco river as
Mauro Pichorim https://orcid.
source of genetic structure in neotropical Pithecopus nordesti-
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SUPPORTING INFORMATION
dry forests. Zoologica Scripta, 00, 1–22. https://doi.
Additional supporting information can be found online
org/10.1111/zsc.12672
in the Supporting Information section at the end of this
article.

Zoologica Scripta - 2024 - Cerqueira - A New Antshrike Aves Thamnophilidae Endemic To The Caatinga and The Role of

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Zoologica Scripta - 2024 - Cerqueira - A New Antshrike Aves Thamnophilidae Endemic To The Caatinga and The Role of

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Received: 18 December 2023

| Revised: 19 March 2024

A new antshrike (Aves: Thamnophilidae) endemic to the

Pablo Cerqueira1,2,3 | Gabriela R. Gonçalves1,2 | Tânia F. Quaresma3,4 |

Zoologica Scripta. 2024;00:1–22. wileyonlinelibrary.com/journal/zsc | 1

T A B L E 1 Diagnostic body-part colour

TaiL 57.03 (b) 4.04 48–68 61.74 (a) 2.86 56–67

Measurementa Mean SD Min–Max Mean SD Min–Max

Extended Bayesian Skyline Plots (EBSPs) indicated a post-

See cover plate (Supplementary Material). Crown, forehead,

The new species is almost restricted to the left/northern

See Table S2 for detailed information of examined tape

spatial environmental data. Global Ecology and Biogeography,

Zimmer, K., Isler, M. L., & de Juana, E. (2003). Silvery-cheeked

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Zoologica Scripta - 2024 - Cerqueira - A New Antshrike Aves Thamnophilidae Endemic To The Caatinga and The Role of

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Zoologica Scripta - 2024 - Cerqueira - A New Antshrike Aves Thamnophilidae Endemic To The Caatinga and The Role of

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Zoologica Scripta - 2024 - Cerqueira - A New Antshrike Aves Thamnophilidae Endemic To The Caatinga and The Role of

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Available Formats

Received: 18 December 2023

| Revised: 19 March 2024

A new antshrike (Aves: Thamnophilidae) endemic to the

Pablo Cerqueira1,2,3 | Gabriela R. Gonçalves1,2 | Tânia F. Quaresma3,4 |

Zoologica Scripta. 2024;00:1–22. ﻿ wileyonlinelibrary.com/journal/zsc | 1

T A B L E 1 Diagnostic body-­part colour

TaiL 57.03 (b) 4.04 48–68 61.74 (a) 2.86 56–67

Measurementa Mean SD Min–Max Mean SD Min–Max

Extended Bayesian Skyline Plots (EBSPs) indicated a post-­

See cover plate (Supplementary Material). Crown, forehead,

The new species is almost restricted to the left/northern

See Table S2 for detailed information of examined tape

spatial environmental data. Global Ecology and Biogeography,

Zimmer, K., Isler, M. L., & de Juana, E. (2003). Silvery-­cheeked

You might also like

Zoologica Scripta. 2024;00:1–22. wileyonlinelibrary.com/journal/zsc | 1

T A B L E 1 Diagnostic body-part colour

Extended Bayesian Skyline Plots (EBSPs) indicated a post-

Zimmer, K., Isler, M. L., & de Juana, E. (2003). Silvery-cheeked