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Revista Brasileira de Ornitologia 26(2): 151–186.
June 2018 ARTICLE
INTRODUCTION
Accipitriformes (osprey, kites, hawks, and eagles; families
Pandionidae and Accipitridae) is an extremely diversied
and successful clade of diurnal raptors (Ferguson-Lees &
Christie 2001, Márquez et al. 2005, Amaral et al. 2009,
Dickinson & Remsen-Jr. 2013). ese predators have a
noteworthy participation in trophic webs, being able to
mediate the whole structure and diversity of a community
(Bierregaard-Jr. 1995, Touchton et al. 2002), and are also
relevant indicators of environmental quality (Jullien &
iollay 1996, Blendinger et al. 2004, iollay 2007)
and providers of important environmental services (Estes
et al. 2011). Breeding biology of this clade is widely
varied (Newton 2010, Whitacre & Burnham 2012), and
knowledge about the breeding patterns of each species
and subspecies plays a central role in their eective
conservation (de Labra et al. 2013).
Many breeding aspects of Accipitriformes are in fact
important parameters for management and conservation
Breeding biology of Neotropical Accipitriformes: current
knowledge and research priorities
Julio Amaro Betto Monsalvo1,3, Neander Marcel Heming2 & Miguel Ângelo Marini2
1 Programa de Pós-graduação em Ecologia, IB, Universidade de Brasília, Brasília, DF, Brazil.
2 Departamento de Zoologia, IB, Universidade de Brasília, Brasília, DF, Brazil.
3 Corresponding author: juliobetto@yahoo.com.br
Received on 08 March 2018. Accepted on 20 July 2018.
ABSTRACT: Despite the key role that knowledge on breeding biology of Accipitriformes plays in their management and conservation,
survey of the state-of-the-art and of information gaps spanning the entire Neotropics has not been done since 1995. We provide
an updated classication of current knowledge about breeding biology of Neotropical Accipitridae and dene the taxa that should
be prioritized by future studies. We analyzed 440 publications produced since 1995 that reported breeding of 56 species. ere is a
persistent scarcity, or complete absence, of information about the nests of eight species, and about breeding behavior of another ten.
Among these species, the largest gap of breeding data refers to the former “Leucopternis” hawks. Although 66% of the 56 evaluated
species had some improvement on knowledge about their breeding traits, research still focus disproportionately on a few regions and
species, and the scarcity of breeding data on many South American Accipitridae persists. We noted that analysis of records from both
a citizen science digital database and museum egg collections signicantly increased breeding information on some species, relative
to recent literature. We created four groups of priority species for breeding biology studies, based on knowledge gaps and threat
categories at global level. Group I (great scarcity of information, plus higher categories of threat): Leptodon forbesi, Cryptoleucopteryx
plumbea, and Buteogallus lacernulatus; Group II (breeding data have recently increased, but threat categories are high): Spizaetus
isidori, Accipiter gundlachi, Buteogallus coronatus, Pseudastur occidentalis, and Buteo ventralis; Group III (“Near reatened” species
with still scarce breeding information): Accipiter poliogaster, Accipiter collaris, Buteogallus aequinoctialis, and Pseudastur polionotus;
and Group IV (other priority cases): Buteo ridgwayi, Buteo galapagoensis, four eagles (Morphnus guianensis, Harpia harpyja, Spizaetus
ornatus and Buteogallus solitarius), Leptodon cayanensis, Accipiter superciliosus, Buteogallus schistaceus, and the three Leucopternis hawks
(L. semiplumbeus, L. melanops and L. kuhli). We also discuss the way that novel breeding data can show in what manners dierent
species and populations are responding to environmental changes.
KEYWORDS: eagles, hawks, information gaps, life history, raptors, reproduction.
programs. For instance, clutch size is directly related to
population size, and this is related with extinction risk
of species (Krüger & Radford 2008). Conversely, their
reproductive rates are related to population density
(Krüger 2000). Also, nest site choices reveal habitat
selection by these raptors (Ferguson-Lees & Christie
2001), and therefore make evident their sensitivity to
environmental changes (Trejo 2007a).
According to the latest classication adopted by the
American Ornithologists' Union (NACC 2017, Remsen-
Jr. et al. 2018 – therefore, AOU), there are 28 genera and
67 species of Accipitriformes occurring in the Neotropical
region. Nonetheless, most Neotropical breeding data
presented in some key references on diurnal raptors (e.g.,
Ferguson-Lees & Christie 2001, del Hoyo et al. 2016)
have important limitations. Information often consist of
no more than anecdotal breeding records coming from
scattered studies, or are generalizations based other tropical
regions of the world (e.g., Newton 2010), largely unveried
to occur in the Neotropics (Whitacre & Burnham 2012).
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Bierregaard-Jr. (1995) reviewed the state of the
knowledge available addressing various aspects of the
biology of 81 diurnal raptors that breed mainly in Central
and South America. Regarding the breeding biology, the
author showed that nests of 11 species of Accipitriformes
and breeding behavior of 15 were not described. Moreover,
most research concentrated on a few regions, such as
further north of the Neotropics (e.g., southern part of
North America, and Guatemala). He also mentions that
breeding data on most of South American species and
subspecies of raptors is lacking (Bierregaard-Jr. 1995).
More recently, similar reviews were done only on
a few South American countries (Pardiñas & Cirignoli
2002, Trejo et al. 2006, Trejo 2007a, b, Raimilla et al.
2012, Cortés et al. 2013). ese studies assessed from
four to 28 species, and just two reviews (Trejo 2007a, b)
dealt with a larger amount (55 species). All these analyses
comprised only studies conducted in the specic country
(ies) (i.e., Argentina, Chile and Uruguay), and so none
included raptors that occur north of the Southern Cone
of South America. Consequently, these surveys left out
the Amazon Basin, one of the world's most decient
areas on bird breeding data, and around 20 species of
Accipitriformes (Whitacre & Burnham 2012, del Hoyo
et al. 2016, Xiao et al. 2016).
Countries that produce most scientic publications
on breeding biology of Neotropical birds do not have
english as their native language (Heming et al. 2013,
Freile et al. 2014). For instance, all recent reviews on
South American raptor research were written in Spanish
(save their abstracts), with the exception of Trejo et al.
(2006). Yet, there is still a visibility bias aecting science
made in such countries (Cabot & de Vries 2004, Lortie et
al. 2007), making such publications not easily accessible
for researchers that do not read spanish or portuguese (see
Bierregaard-Jr. 1995).
Moreover, many information on the natural history
of Neotropical raptors come from studies not specically
designed for this aim (Cortés et al. 2013). Such studies
often are published at small, local journals or bulletins
(Figueroa, in litt.). us, important advances in
knowledge are hardly visible to ornithologists from other
countries. Indeed, Bierregaard-Jr. (1995) mentioned
that “inaccessibility” of certain Latin American journals
may have prevented him from collecting information
from them. However, since then, internet access to
many of these journals greatly improved (e.g., Hornero,
from Argentina; http://digital.bl.fcen.uba.ar), allowing
more complete reviews to be made. Also, during the last
two decades, the ornithological community of South
America increased considerably, boosting the number of
publications (Vuilleumier 2004, Freile 2005, Freile et al.
2014).
Scrutiny of oological (egg) collections from museums
could also be useful for avian breeding biology research
(McNair 1987). Yet, very few researchers in the Neotropics
used museum eggs for analyzing breeding traits of diurnal
raptors (e.g., Denis et al. 2013, Hayes 2014), the most
frequent approach being the presentation of revised
summaries of some specic collections (e.g., Román &
Wiley 2012). Also, Bierregaard-Jr. (1995) did not provide
information on museum eggs when evaluating knowledge
on breeding biology of diurnal raptors, although such
data is to some extent included in past literature (e.g.,
Belcher & Smooker 1934). e amount of information
(unpub. data) that we and other authors (Murphy 1989,
Olsen & Marples 1993) obtained from museum egg sets
strongly suggests that such sources could provide data not
easily obtainable from other sources.
Considering the above, there is a need for a new
comprehensive survey to access the state of the knowledge
on the breeding biology of Neotropical Accipitriformes,
and an update on research priorities. So, our main objective
was to make a comprehensive analysis of the literature
on breeding biology of Neotropical Accipitriformes
produced after Bierregaard-Jr.'s (1995) review, and thus,
to dene the taxa that should be prioritized by future
studies. We created an updated classication of current
levels of knowledge of the breeding biology of these
raptors, evaluating the progress made in the last decades.
We also discuss the information gaps, ponder on their
possible causes, implications, and potential solutions
to the lack of breeding data, and present additional
information obtained from alternative sources such as
a citizen science database and museum collections. To
conclude, we briey exemplify how breeding data can
show the ways that dierent species and populations are
responding to environmental changes.
METHODS
Taxa
We follow Bierregaard-Jr.'s (1995) criteria by not
including species with centers of distribution outside the
Neotropics (see below), and Nearctic taxa that do not
breed in there (which excluded the family Pandionidae
from the analysis). us, we perform a comprehensive
recent review of Neotropical raptors, including 56 species.
Our subspecies division follows Dickinson & Remsen-Jr.
(2013).
Categories and scoring criteria, and major
changes in classication
We used two categories concerning reproduction, largely
based on Bierregaard-Jr. (1995) and Trejo (2007a). Under
“nest”, the information that we analyzed includes the
physical description of the nest, as well as its seasonality
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and location, clutch size, and description of eggs. at is,
all aspects, mostly “physical”, related to the early nesting
stage. Under “breeding behavior”, we included breeding
displays of adult birds; descriptions of copulating and
parental behaviors; incubation and edging times;
development of the young (both morphological and
behavioral); the period of dependence of juvenile(s)
after its rst ights (post-edging dependency period);
and more detailed information – provided by relatively
few studies – such as spatial distribution of breeding
pairs, rate of reproductive success, nest productivity, and
subsequent dispersal and survival of juveniles.
e numerical scores assigned in the classicatory
scale of knowledge also follow the criteria of Bierregaard-
Jr. (1995) and Trejo (2007a): (0) no information; (1)
only anecdotal/scattered reports; (2) detailed study of
one breeding pair or event; (3) study of more than one
pair in the same population, and/or a substantial amount
of anecdotal reports of representative areas of the species
range; (4) detailed studies of separate populations in
dierent portions of the species range; and (5) detailed
information from the entire range of the species.
Besides producing an updated classication of
current levels of knowledge about the breeding biology
of these raptors, these scores act as an intuitive measuring
scale to signal whether some reproductive aspects
and taxa still need more studies (see also “Research
recommendations and conservation relevance”). More
importantly, they allowed a comparison between our
scores and those reported by Bierregaard-Jr. (1995), to
assess whether levels of knowledge changed in the last
decades, and thus identify persistent gaps.
Classication had to be evaluated and updated, due
to changes since 1995. Two of these changes were the
recent splits of the “Gray Hawk” complex (Buteo nitidus/
Buteo plagiatus; Millsap et al. 2011), and of Cuban Black
Hawk Buteogallus gundlachii and Common Black Hawk
Buteogallus anthracinus (Wiley & Garrido 2005). On the
rst case, the split of the taxon into southern and northern
forms facilitates the evaluation of its case, and we chose
to consider the scores attributed to “Buteo nitidus” by
Bierregaard-Jr. (1995), as default for both B. nitidus and
B. plagiatus. For the Black Hawks, Bierregaard-Jr. did not
report a separate score for the then subspecies gundlachii,
what prevented us from making a comparison of levels of
knowledge about this taxon then and now. Nevertheless,
we briey discuss the status of Cuban Black Hawk on
Appendix III.
Bierregaard-Jr. (1995) reported dierent scores
for the taxa Accipiter ventralis, Accipiter chionogaster and
Accipiter erythronemius, but these are currently classied
as subspecies of the Sharp-shinned Hawk Accipiter striatus
(Remsen-Jr. et al. 2018). In turn, Sharp-shinned Hawk
was not included in Bierregaard-Jr.'s review, for having a
center of distribution outside Central and South America.
Ferguson-Lees & Christie (2001) already argued that this
so-called “Central and South American group” of Sharp-
shinned Hawk's subspecies (that is, A. s. ventralis, A. s.
chionogaster and A. s. erythronemius) is so divergent, that
treatment at species level should be considered for at least
some of these, but not for other groups of subspecies
such as the Caribbean. Since Remsen-Jr. et al. (2018)
acknowledge that the taxonomic status of A. striatus still
needs clarication, we comment on the knowledge on
those three subspecies on Appendix III.
Other hawk species with some breeding populations
in the Neotropics (mostly in the Caribbean) but centers
of distribution in the Nearctic were excluded from
our analysis. We based such decision not only because
comparing scores of knowledge then and now was
impossible, since Bierregaard-Jr. (1995) also excluded
those from his assessment. Most importantly, we rely
on evidence of little divergence between some of such
disjunct populations and its Nearctic counterparts, on
respect of phenotypic traits (Ferguson-Lees & Christie
2001), especially most breeding aspects (e.g., Santana &
Temple 1988). Likewise, such ndings are being further
supported by an ongoing meta-analysis of geographical
variation on these species breeding patterns (author's
unpub. data).
Other splits adopted by Bierregaard-Jr. (1995),
but not maintained on current classication, are
“Accipiter chilensis” (subspecies of Bicolored Hawk A.
bicolor), “Buteogallus subtilis” (included three subspecies
of Common Black Hawk) and “Buteo poecilochrous”
(subspecies of Variable Hawk Geranoaetus [Buteo]
polyosoma). We ignored the scores that Bierregaard-Jr.
separately assigned to each of these taxa, and analyzed
only those ascribed to the currently recognized species.
Yet, we commented on the status of some of these
subspecies when relevant.
Literature search methods and sources
We screened the Global Raptor Information Network
(GRIN; http://www.globalraptors.org/grin/indexAlt.asp)
until October 2016. is database focus only on raptors,
concentrating information on diurnal species from around
the world and includes bibliography of other renowned
databases on raptors such as e Peregrine Fund and
Raptor Information System. We analyzed the literature
on reproduction of the 56 species after 1994, indicated
in the section “Breeding” in the species accounts. We
also searched for other studies whose titles refer to
reproductive aspects, mainly the bibliography contained
in the topic “Breeding biology”. In some isolated cases,
we considered in this review breeding data not published
in other sources and made available by researchers in the
GRIN database.
We chose to use Google Scholar (http://scholar.
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Revista Brasileira de Ornitologia 26(2): 2018 Revista Brasileira de Ornitologia 26(2): 2018
google.com/) as the main tool to complement GRIN
reference search because we noted it was able to locate
the same references found with Scopus and Searchable
Ornithological Research Archive (SORA; http://elibrary.
unm.edu/sora), search tools also chosen by almost all
recent revisions (Trejo 2007a, b, Raimilla et al. 2012,
Cortés et al. 2013). e search terms we used were all
possible scientic names recently assigned for these
species (except for those variables only in the sux,
which were already supplied by the search heuristic),
combined with each of the following terms: nest, ninho,
nido, nidicação, anidamiento, anidación, reprodução,
reproducción, breeding, and biologia reprodutiva. e
great redundancy of results when using somewhat similar
terms indicated the eectiveness of the choices, and terms
like “nesting” and “biología reproductiva” were discarded.
We searched for all kinds of references, from articles
in any category of scientic journal, through monographs,
conference abstracts and posters, to technical reports
and unpublished manuscripts. We reviewed citations
contained in the references, even though most were
already found in key word searches. Yet, we could not
retrieve 19 (4.1%) of the 459 references produced
between 1995–2016 (Appendix IV), neither through
requesting directly from their authors nor from databases
such as e Peregrine Fund.
We also screened and retrieved information from a
bibliographical review of Brazilian birds (Oniki & Willis
2002), and the following books: Bird et al. (1996), Sick
(1997), Machado et al. (1998), Arballo & Cravino (1999),
Naka & Rodrigues (2000), Höing & Camargo (2002),
Fontana et al. (2003), Reichle et al. (2003), Wheeler
(2003), Willis & Oniki (2003), Antas (2004), Mikich &
Bérnils (2004), de la Peña (2005), Márquez et al. (2005),
Angehr (2006), Sigrist (2006), Eisermann & Avendaño
(2007), Gussoni & Guaraldo (2008), Whitacre (2012),
Santos (2014), Straube et al. (2014), and Alvarado et al.
(2015).
Exclusion and inclusion search criteria
As previously mentioned, Bierregaard-Jr. (1995)
claimed that antiquity or “obscurity” of certain journals,
particularly Latin American's, prevented him from
gathering information from them. Yet, he did include
some of these studies that were cited in more broadly
distributed journals. We veried that some of these
Latin American journals (e.g., Hornero) were already
scrutinized by recent reviews (Trejo 2007a, b, Raimilla
et al. 2012). Notwithstanding, we could not determine
with certainty which studies prior to 1995 were not
included by Bierregaard-Jr., given that his study lacks a
complete list of references. So, we opted to consider only
papers published from 1995 on, to avoid repeating data
already collected. After all, one of our aims was to get a
clear picture of the amount of research done in the last
decades, and not previously.
We also assume that papers from 1995 would not
have been included by Bierregaard-Jr. Depending on
the date of completion of his search (not stated in the
paper), the author could have included at least some of
these studies, but information contained in such papers
is not consistent with certain scores assigned by him [e.g.,
the Gray-backed Hawk Pseudastur occidentalis, studied
by Vargas (1995)]. is fact suggests that in most cases
the inclusion of these papers in that review may not have
occurred. Nevertheless, only a few studies from 1995
were found in our review, suggesting that the inuence
of possible duplicate data on the dierent species would
be irrelevant.
Some books contain secondary information often
without direct citation of the original data (e.g., Ferguson-
Lees & Christie 2001, Márquez et al. 2005, Sigrist 2006).
Because of lack of clear indication of each of their sources
in the text, we could not retrieve the original studies year
or sometimes even the geographic region. us, we also
chose to not include such breeding reports, except when
it was clearly indicated in the text that it was an original
data.
Research recommendations and
conservation relevance
We created a four-group classication of research
priorities on species breeding aspects, based mostly on
knowledge gaps (by means of the assigned numerical
scores), but also considering current threat categories at
the global level (IUCN 2017). Group I includes species
with great scarcity of available information about their
reproduction, combined with higher categories of threat.
Group II comprises species whose studies have advanced,
although very little since Bierregaard's (1995) review,
but which are at some higher threat category. Group III
includes species whose knowledge is still scarce and are
currently “Near reatened” according to IUCN. Finally,
Group IV represents species framed in three possible
situations: i) the knowledge about their breeding has not
increased (although it was already very high, i.e. scores
of 4 or 5) and also are in some greater category of threat;
ii) the remaining species considered “Near reatened”;
or iii) species not threatened, but of which nothing or
practically nothing is known about their reproduction
and/or have at least one of the topics of breeding aspects
classied as 1 (see “Categories and scoring criteria, and
major changes in classication”).
Screening of the Handbook of Birds of the
World and WikiAves
e Handbook of Birds of the World (HBW) was the
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baseline for Bierregaard-Jr.'s (1995) gap analysis and until
today is considered a reference for current knowledge
about biology of bird species (e.g., Trejo et al. 2006, Xiao
et al. 2016). us, we opted to review information in
the online version “HBW Alive” (http://www.hbw.com).
Our purpose was to determine if data available regarding
reproductive aspects (topic “Breeding”, in each species
account) were commensurate with the actual state of
knowledge about these subjects.
e online database WikiAves (www.wikiaves.
com) is a collaborative tool launched in 2008 that allows
posting of photographic records of bird species that occur
in Brazil. is initiative has a great advantage over other
popular citizen science platforms, such as eBird (ebird.
org), by working with digital records and not lists. Also,
we are not aware of initiatives from other Neotropical
countries (e.g., http://www.wikiaves.com.ar/inicio.php)
that are equally reliable and allow similar content-based
searches of their records.
Considering the enduring scarcity of avian breeding
records from South American mid-latitudes (Baker 1938,
Heming et al. 2013), the fact that WikiAves focus on
Brazil is particularly convenient. We searched for breeding
records of 25 species in this database. e low number of
species was due primarily to the scope of WikiAves, which
only contains species recorded in Brazil. In addition, we
chose to review only species that obtained scores less than
3 in at least one of the categories, or those with values
equal to or greater than that, but for which there was a
marked relative scarcity of South American data. In the
“Advanced Search” tool for photos, we used (separately)
the lters: egg, nest, juvenile, copulating, incubating,
courting, caring/feeding its chick(s), and making nest.
e search was made in October 2016 and we included
only records whose identication was considered secure
– both at specic level and, in the case of breeding
behaviors and/or stages that were clearly illustrated in the
photographic records. Records already present in papers
located in the survey were discarded.
Museum egg records
Eggs and labels were photographed in the following egg
collections between 2014–2017 at Western Foundation
of Vertebrate Zoology - WFVZ (Camarillo, USA),
Natural History Museum - NHMUK (Tring, UK),
National Museum of Scotland - NMS (Edinburgh, UK),
Muséum national d'Histoire Naturelle - MNHN (Paris,
France), Naturhistoriches Museum - NMW (Wien,
Austria), Instituto de Investigación de los Recursos
Biológicos “Alexander von Humboldt” - IAVH (Villa
de Leyva, Colombia), Museo Argentino de Ciencias
Naturales “Bernardino Rivadavia” - MACN (Buenos
Aires, Argentina), Museo de Ciencias Naturales de La
Plata - MLP (La Plata, Argentina) and in Brazil, Museu
de Zoologia da Universidade de São Paulo - MZUSP
(São Paulo), Museu Nacional do Rio de Janeiro - MN
(Rio de Janeiro), Museu Paraense “Emilio Goeldi” -
MPEG (Belém), and Coleção Ornitológica “Marcelo
Bagno” - COMB (Brasília). We also visited the online
egg collections of the Field Museum of Natural History
- FMNH (Chicago, USA), and the Arctos Collaborative
Collection Management Solution (arctos.database.
museum), and had access to data of the egg collection of
the Smithsonian Institution (USNM, Washington, D.C.,
USA), and the American Museum of Natural History
(AMNH, New York, USA). Finally, we consulted the
catalog of the Cris-Rivers Region Museum (CRRM,
Oradea, Romania; Béczy 1971).
ese author's previous experience suggests that
diurnal raptor's eggs collected in the United States can
outnumber those from all other new world countries
together, on a ratio of roughly nine to one (authors'
unpub. data). Also, Bierregaard-Jr. (1995) veried that
when the distribution of a species reaches the southern
part of North America, it tends to be much more studied
there than in the rest of its range. Considering the above,
we opted to not include museum data from eggs collected
in the USA in this analysis. Breeding information from
that country certainly is already overly represented in
literature, and augmenting it with museum records would
only exacerbate this bias.
Museum egg sets are a proven reliable source
(McNair 1987), but a few inconsistencies in the records
of certain collectors have been reported (Hellmayr &
Conover 1949, orstrom & Ki 1999). us, we
carefully validated species identication based on our
own experience, on remarks from other researchers, and
also resorting on other references that provide clutch
sizes, egg measurements and descriptions (e.g., the GRIN
database). A few species suer from faulty information
about their eggs and clutches in the literature, and these
cases are still being validated by us. Such egg sets are not
assigned to any species here but are included in the total
number of sets we found from the Neotropics. In the
process of validating eggs' identication, measurements
were standardized using the software ImageJ (Bridge et al.
2007, Troscianko 2014).
RESULTS
Bierregaard-Jr.'s (1995) review found 431 references
of 81 species and included information about various
aspects of Neotropical raptor biology. Meanwhile, our
research found 440 references exclusively about breeding
biology of 54 species (out of 56 studied taxa – as we
did not nd any published records for two species).
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Such results are presented in Appendix I, with complete
reference list on Appendix II. is represents an increase
in the number of published references since Bierregaard-
Jr.'s review, especially since he covered many other aspects
of biology, included also Falconidae, and had no date
limitation (unlike our scope of 22 years). We found 11
references and citations referring to data from captive
birds, but these were not included in our review given
the uncertainty involving raptor's breeding aspects in
unnatural conditions (Cabot-Nieves et al. 2013).
Much of the breeding data we found came from
inventories that provide a list of species for one or
more localities, often highlighting new occurrences or
noteworthy records (e.g., Bodrati et al. 2010), or research
addressing ecological aspects of bird communities of a
given region (e.g., Cintra & Naka 2012). Observations
on the breeding activity of some species are frequently
included in such studies (e.g., Hennessey et al. 2003),
and it is common for raptors to receive some prominence
(e.g., Greeney & Nunnery 2006). However, such reports
still remain mostly anecdotal (e.g., Ruvalcaba-Ortega &
González-Rojas 2009). For instance, nest records often
do not provide any information on nest content or stage
(e.g., Bodrati et al. 2010), frequently because the nest was
presumably inaccessible to the researchers (e.g., Bellatti
2000). Many times all that can be concluded is that the
species was “nesting” in a given locality, during a quite
long period of time (e.g., Cavicchia & García 2012).
Of the 11 species of Neotropical accipitrids for
which the nest had not been described prior to 1995,
six remain undescribed and two present only anecdotal/
scattered reports (Table 1). Of 15 species with no
information about their breeding behavior (e.g., Leptodon
cayanensis, Cryptoleucopteryx plumbea, Leucopternis
melanops) in 1995, little or no additional information is
still not available for 10. Also, in 1995 only anecdotal
descriptions were available for the nests of 15 species,
and breeding behaviors of another 14 species. is case
remains the same for the Tiny Hawk Accipiter superciliosus
and Rufous Crab Hawk Buteogallus aequinoctialis, which
have no recent published information. Yet, 66% of the
analyzed species (n = 37) showed an increase in knowledge,
of these, nearly half (n = 19) showed an increase in only
one of the categories, and the remaining in both.
Probably the most signicant increases in knowledge
were for Barred Hawk Morphnarchus princeps and White-
throated Hawk Buteo albigula, followed by Gray-bellied
Hawk Accipiter poliogaster, Chaco Eagle Buteogallus
coronatus, Gray-backed Hawk and Rufous-tailed Hawk
Buteo ventralis, and also Rufous-thighed Kite Harpagus
diodon. e following species also had a signicant increase
in knowledge about the two breeding categories: Black-
and-white Hawk-Eagle Spizaetus melanoleucus, Black-
collared Hawk Busarellus nigricollis, Long-winged Harrier
Circus buoni, Crane Hawk Geranospiza caerulescens,
Solitary Eagle Buteogallus solitarius and Short-tailed
Hawk Buteo brachyurus. On the other hand, very scant
information was found for the former “Leucopternis”
hawks, currently classied in ve genera. Even the best-
known species in this polyphyletic group of 10 species
(Amaral et al. 2009), the Barred Hawk and the White
Hawk Pseudastur albicollis, either have only anecdotal
reports in distinct areas of the species distribution range,
or detailed studies of nests from just one population (e.g.,
Muela & Valdez 2003, Cisneros-Heredia 2006, Gelis &
Greeney 2007, Draheim 2012).
As Bierregaard-Jr. (1995) also noted, we found a
longstanding concentration of studies further north of the
Neotropics (i.e., southern United States), as well as in the
northern portion of this region. For instance, Guatemala
still stood out due to the quantity and quality of research
developed by the Peregrine Fund's Maya Project, which
resulted in a large number of published studies on raptor
biology (e.g., Seavy & Gerhardt 1998, Seavy et al. 1998,
orstrom & Quixchán 2000, Sutter et al. 2001, Panasci
& Whitacre 2002), ultimately leading to the publication
of a book (Whitacre 2012). e Southern Cone of South
America also have a large amount of research developed in
Chile, already emphasized by Bierregaard-Jr. (1995), and
Argentina (e.g., Jiménez 1995, Trejo et al. 2001, Ojeda et
al. 2003, Medel-Hidalgo et al. 2015, Pérez 2015, Rivas-
Fuenzalida et al. 2015).
Even for species considered already relatively
well known, with both categories scoring 3 or 4, there
is a lasting shortage of research on South American
populations or subspecies. is was the case for the
White-tailed Kite Elanus leucurus, the Swallow-tailed
Kite Elanoides forcatus, and the Zone-tailed Hawk
Buteo albonotatus, among others. We also found little
or no information about the breeding biology of some
subspecies of some polytypic species, including the Cuban
Kite (Chondrohierax uncinatus wilsonii), considered a full
species and “Critically Endangered” by IUCN (2017);
Mangrove Black Hawk (Buteogallus anthracinus subtilis),
included in a separate species by Bierregaard-Jr. (1995);
Pearl Kite (Gampsonyx swainsoni magnus); and Snail Kite
(Rostrhamus sociabilis major). Additional comments in
Table 1 are given to indicate taxa and/or regions in which
research is critically needed.
Although incomplete, some sets of new studies
revealed both similarities and divergences in breeding
behavior between dierent populations. For instance,
the cooperative behavior of Harris's Hawks Parabuteo
unicinctus, well known for the subspecies P. u. harrisi in
the United States, at the time of Bierregaard-Jr.'s (1995)
review was not reported anywhere else in the species range.
ere is now good evidence that cooperative breeding
also occurs in at least one population of the nominate
subspecies in southeastern Brazil (Silva & Olmos 1997),
hence this behavior is not restricted to North America.
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Table 1. Assessment of current knowledge on the breeding biology of 56 species of Neotropical Accipitriformes.
Species Bierr.
Nest
Bierr.
Behav Nest Breeding
behavior
Research
priority Comments
Elanus leucurus 4 4 4 4 No Lack of more detailed data from most regions, mainly
South America.
Gampsonyx swainsonii 3 3 3 3 No Still a lack of behavioral data from most regions,
particularly later stages.
Chondrohierax uncinatus 4 3 4 4 No Most data missing from South America; nothing from
subspecies wilsonii.
Leptodon cayanensis 1 0 3 1 IV Detailed data from only two areas; very few behavioral
data, particularly later stages.
Leptodon forbesi 0 0 0 1 I Only breeding displays.
Elanoides forcatus 3 3 4 4 No Many detailed studies, but there is still a lack of detailed
data from other areas.
Morphnus guianensis 2 2 3 3 IV Some detailed studies, but still a lack of behavioral data
in many regions.
Harpia harpyja 4 3 4 4 IV Still a lack of detailed data from some portions of the
range (e.g., Atlantic Forest).
Spizaetus tyrannus 3 3 3 4 No Still a lack of detailed data from many regions.
Spizaetus melanoleucus 1 1 3 3 No Isolated cases and incomplete observations.
Spizaetus ornatus 4 4 4 4 IV New data did not change status.
Spizaetus isidori 3 2 3 3 II Still a lack of detailed data from many regions.
Busarellus nigricollis 1 1 3 3 No Still a lack of detailed data from many regions.
Rostrhamus sociabilis 4 4 4 4 No Many detailed studies, but still missing data from most
regions/subspecies.
Helicolestes hamatus 3 3 3 3 No New data did not change status; only one population
studied in detail.
Harpagus bidentatus 3 1 3 3 No Only one population studied in detail; still a lack of
behavioral data.
Harpagus diodon 1 0 3 3 No Isolated cases and incomplete observations; still a lack
of behavioral data.
Ictinia plumbea 3 3 4 3 No Still a lack of more behavioral data from many regions.
Circus cinereus 3 1 3 3 No Lack of more detailed data from many regions.
Circus buoni 1 1 3 3 No Lack of more detailed data from many regions.
Accipiter poliogaster 0 0 2 3 III Basically, just one or two pairs studied in detail.
Accipiter superciliosus 1 1 1 1 IV Still very little information.
Accipiter collaris 0 0 0 1 III Only information of specimens on breeding condition.
Accipiter gundlachi 3 1 3 3 II Some detailed studies, but coming from a few areas.
Accipiter bicolor 3 3 3 3 No Most data missing for two subspecies; new data but
several old ones discarded.
Geranospiza caerulescens 1 1 3 3 No Only one population studied in detail.
Cryptoleucopteryx plumbea 0 0 0 0 I No new data.
Buteogallus schistaceus 0 0 0 0 IV No new data.
Buteogallus anthracinus 4 4 4 4 No Still a lack of South American data, especially from
subspecies subtilis.
Buteogallus aequinoctialis 1 1 1 1 III Still very little information.
Buteogallus meridionalis 4 3 4 3 No New data did not change status; still a lack of detailed
data from many regions.
Buteogallus lacernulatus 0 0 0 1 I Only displays.
Buteogallus urubitinga 3 3 4 3 No Still a lack of more behavioral data from most regions.
Buteogallus solitarius 1 1 3 3 IV Data on nests or late stages (nothing in between); lack of
data from most regions.
Buteogallus coronatus 1 1 4 3 II Many detailed studies, but there is still a lack of more
behavioral data.
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Species Bierr.
Nest
Bierr.
Behav Nest Breeding
behavior
Research
priority Comments
Morphnarchus princeps 0 0 3 3 No Most data missing from many regions.
Rupornis magnirostris 3 3 4 3 No Some detailed studies, but still a lack of behavioral data
from most regions/subsp.
Parabuteo unicinctus 4 4 4 4 No New data did not change status; but evidence of
cooperative behavior in Brazil.
Parabuteo leucorrhous 1 1 2 3 No Isolated cases and incomplete observations.
Geranoaetus albicaudatus 3 3 3 3 No Detailed data only of two subspecies; lack of detailed
data from many regions.
Geranoaetus polyosoma 3 3 4 3 No Still a lack of more behavioral data.
Geranoaetus melanoleucus 3 3 4 3 No Some detailed studies, but still a lack of more behavioral
data from many regions.
Pseudastur polionotus 0 0 1 1 III Very little information.
Pseudastur albicollis 3 3 3 3 No New data did not change status; only one population
studied in detail.
Pseudastur occidentalis 0 1 3 3 II Only one population studied in detail.
Leucopternis semiplumbeus 1 0 1 1 IV No signicant advances.
Leucopternis melanops 0 0 ? ? IV No real advances.
Leucopternis kuhli 0 0 1 0 IV Only one nest.
Buteo plagiatus 3 3 4 3 No Still a lack of detailed data from most regions.
Buteo nitidus 3 3 3 3 No New data did not change status; many missing data,
incl. more egg descriptions.
Buteo ridgwayi 5 4 5 4 IV New data did not change status; still a lack of more
behavioral data.
Buteo albigula 1 0 4 3 No Breeding status in northern range still uncertain; many
missing data, incl. on eggs.
Buteo brachyurus 1 1 3 3 No Lack of more detailed data from most regions, mainly
South America.
Buteo galapagoensis 5 5 5 5 IV -
Buteo albonotatus 3 2 3 3 No Still limited to the northern range.
Buteo ventralis 1 0 3 3 II Still limited to Chile; many missing data, including
more egg descriptions.
Bierr. Nest and Bierr. Behav = scores assigned by Bierregaard-Jr. (1995), on Nest and Breeding behavior respectively; Nest and
Breeding behavior = scores assigned by this study. Scores: (0) no information; (1) only anecdotal/scattered reports; (2) detailed study
of one breeding pair or event; (3) study of more than one pair in the same population, and/or substantial amount of anecdotal reports
of representative areas of the range; (4) detailed studies of separate populations in dierent portions of the range; and (5) detailed
information from the entire range. Shaded cells denote improvements on knowledge in the last decades. Research priority = whether
species should be prioritized by future studies on breeding biology, and for those that should, the priority group (I-IV) to which it
was assigned; names of such species are also given in bold letters. Further explanations on the main text. Taxonomic ordering follows
AOU (2018).
On the other hand, Short-tailed Hawk's breeding traits
such as duration of the post-edging dependency period
and nest defense behaviors diverge not only among the
dierent subspecies but even within the same country
(Monsalvo 2012).
e species formerly called the Gray Hawk was
separated into two species by Millsap et al. (2011),
amendment accepted by the AOU (Remsen-Jr. et al.
2018). However, most recent studies of Buteo nitidus,
all published prior to this split (e.g., Patrikeev 2007,
Ruvalcaba-Ortega & González-Rojas 2009), focused on
the current northern species (Gray Hawk, B. plagiatus).
us, the status of the Gray-lined Hawk (B. nitidus
sensu AOU) remains the same. Although the number of
references found was similar (ten and seven, respectively;
Appendix I), information about Gray Hawks comes
from almost 100 breeding events, at about ten dierent
locations. Whereas for Gray-lined Hawks, only six
records were found, and some of these informations
could not have their localities conrmed. Such lack of
detail prevented us from determining if data on the eggs
of the latter species provided in recent literature (Sick
1997, Reichle et al. 2003) do not, in fact, refer to the
northern species eggs.
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Based on the criteria put forward before (see
“Categories and scoring criteria, and major changes
in classication” in the Methods), the highest priority
species for research on their breeding aspects are, as
follow: White-collared Kite Leptodon forbesi, Plumbeous
Hawk Cryptoleucopteryx plumbea, and the White-necked
Hawk Buteogallus lacernulatus (Group I); Black-and-
chestnut Eagle Spizaetus isidori, Gundlach's Hawk
Accipiter gundlachi, Chaco Eagle, Gray-backed Hawk,
and Rufous-tailed Hawk (Group II); Gray-bellied Hawk,
Semicollared Hawk Accipiter collaris, Rufous Crab Hawk
and Mantled Hawk Pseudastur polionotus (Group III);
and the two island species of Buteo hawks (Ridgway's
B. ridgwayi and Galapagos B. galapagoensis), four eagles
(Crested Morphnus guianensis, Harpy Harpia harpyja,
Ornate Hawk-Eagle Spizaetus ornatus and Solitary Eagle),
Gray-headed Kite Leptodon cayanensis, Tiny Hawk,
Slate-colored Hawk Buteogallus schistaceus, and the three
Leucopternis hawks (Group IV).
Despite recent reviews considered HBW as
informative of the state-of-the-art (Trejo et al. 2006, Xiao
et al. 2016), we concluded that information provided
in the “Breeding” topic in this reference is outdated
for at least 18 of the 56 species that we analyzed. In
the WikiAves database, we compiled a total of 174
photographic records representing breeding aspects, for
18 of the 25 species surveyed (Appendix V). No reliable
records were available for the remainder of the species.
For one of these 18 species, Gray-bellied Goshawk, which
had detailed literature records of only one or two breeding
pairs (de Vries & Melo 2000, orstrom 2002, Boesing et
al. 2012), inclusion of data from WikiAves augmented its
assessment score (Table 2).
Another species for which WikiAves allowed a
change in the assigned score was the White-collared Kite,
whose only nesting record (Brito 2013, also quoted by
HBW) is posted on that platform. It is also noteworthy
the case of the Rufous-thighed Kite, for which WikiAves
provides 42 records of at least 15 distinct breeding events
in six dierent states of Brazil, including pairs with nesting
accompanied throughout, and even in consecutive years.
In addition to these three species, another ve showed
a signicant increase in breeding records from South
America, although these not have allowed an eective
change in their scores (Table 2).
We located 729 egg sets from the Neotropical
Table 2. Results of the search for photographic breeding records from the WikiAves database, for 25 species of Neotropical
Accipitriformes.
Species Change in
score(s) Comments
Elanus leucurus No Many records of dierent stages and populations, but did not change
status.
Chondrohierax uncinatus No Only three or four breeding pairs; always more southernly records.
Leptodon cayanensis No Only one nest, not monitored.
Leptodon forbesi Nest = 1 e rst nest of the species, also cited in HBW.
Spizaetus melanoleucus No Little informative and poorly distributed records.
Rostrhamus sociabilis No Many records of dierent stages and populations, but did not change
status.
Helicolestes hamatus No Only two breeding localities, records of later breeding stages.
Harpagus bidentatus No ree records from the same locality, presumably of the same pair.
Harpagus diodon No Some breeding events monitored thoroughly, including same pair in
dierent years.
Accipiter poliogaster Nest = 3 Little informative and always more southernly records.
Accipiter superciliosus No Nothing.
Accipiter bicolor No Only three records, with no new information on subspecies.
Geranospiza caerulescens No Very diverse breeding stages, especially of the subspecies exipes.
Buteogallus schistaceus No Nothing.
Buteogallus anthracinus No Only one nest, no new information.
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Species Change in
score(s) Comments
Buteogallus aequinoctialis No One copulation record.
Buteogallus lacernulatus No No reliable records.
Parabuteo leucorrhous No Nothing.
Pseudastur polionotus No Only one nest, not monitored.
Pseudastur albicollis No Only two nests, no new information.
Leucopternis melanops No Nothing.
Leucopternis kuhli No Nothing.
Buteo nitidus No Some poorly distributed records.
Buteo brachyurus No Many records of dierent stages and populations, but did not change
status.
Buteo albonotatus No No reliable records.
Change in score(s) = whether scores assigned previously in our review, for the two categories concerning reproduction (“Nest” and “Breeding
Behavior”, see Table 1) augmented with inclusion of data from WikiAves. Shaded cells denote any substantial addition of new information, relative
to recent literature.
region in egg collections, besides six records of eggs laid
in captivity in this same region. Of these 729, 706 could
be soundly assigned to some species (Table 3), from
which 58% pertain to only four species: White-tailed
Kite, Common Black Hawk, Roadside Hawk Rupornis
magnirostris, and Gray Hawk. Around 88% of the total
of clutches of these four species were collected in Mexico.
is country is also the origin of almost two-thirds of the
egg sets of all 31 species reliably identied in museum
collections. Argentina and Chile are respectively the
second and third countries with more collected clutches,
but each represents less than 10% of the total.
We propose a correction in the identication of four
clutches, all in the WFVZ collection and all previously
recognized as misidentied by L. Ki (Appendix VI).
From our analyses, we conclude that their correct
identications probably agree with those tentatively
suggested by him in the data slips accompanying these
egg sets. We highlight the relevance of the egg sets
assigned to White-rumped and Gray-lined Hawks, as
they almost doubled the number of breeding reports for
each of these species. Overall appearance and dimensions
from the former's eggs are similar to those reported by
Zilio & Mendonça-Lima (2012), the only other clutch
known for the White-rumped Hawk, but museum eggs
are slightly larger.
Unfortunately, the clutches of Gray-lined Hawk
that we located are essentially the same widely used as
reference for this species (Belcher & Smooker 1934), yet
their measurements are within the range described for the
allospecies Gray Hawk B. plagiatus (del Hoyo et al. 2016).
Also relevant are egg sets from the subspecies
Gampsonyx swainsoni magnus (n = 1) and Rostrhamus
sociabilis major (n = 7), both largely absent in recent
literature. We also located ve clutches of the Mangrove
Black Hawk (former Buteogallus subtilis), for which
Bierregaard-Jr. (1995) found no breeding information
in literature (but see Wetmore 1965). Likewise, in our
literature review we located only poorly detailed, scattered
reports of nesting in a few localities of its range (Barrantes
1998, Pérez-León 2007, Alava et al. 2011). Relative to
recent literature, museum eggs allowed a substantial
increase in breeding information for a total of six species.
DISCUSSION
Breeding knowledge is not yet uniformly distributed
across dierent regions for most species of Neotropical
Accipitridae, with many areas lacking more studies about
their populations or subspecies. e main evidence of this
poor distribution of breeding data is the fact that we have
not assigned any new score of 5 (i.e., detailed information
coming from the entire range). Information on many
South American Accipitridae is still scant, even after
two decades (Bierregaard-Jr. 1995). With exception of a
few restricted-range subspecies, most of the least-studied
populations occur in mid-latitudes of South America or
in the Amazon Basin, a situation that barely improved in
the last eight decades (Baker 1938, Xiao et al. 2016).
e regions where most quality-research are still
concentrated are near the limits of many species ranges.
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Table 3. Results of the search for museum egg records of Neotropical Accipitriformes.
Species No. of sets Comments
Elanus leucurus 65 Mostly from Mexico; also southern South America.
Gampsonyx swainsonii 2 From Colombia and Peru; the latter of subspecies G. s. magnus.
Chondrohierax uncinatus 8 All from Mexico; eggs from Trinidad were misidentied.
Leptodon cayanensis 5 ree of these were misidentied as other species.
Elanoides forcatus 4 From Brazil and Venezuela.
Morphnus guianensis 1From Panama; presumably from the wild but no further details
known.
Harpia harpyja 1 From Amazon Basin; plus 6 clutches laid in captivity.
Spizaetus ornatus 1 From Guatemala, at the same site of Peregrine Fund's Maya Project.
Busarellus nigricollis 4 All sets but one from Paraguay.
Rostrhamus sociabilis 34 Most from South American countries; seven clutches of R. s. major.
Ictinia plumbea 18 Records from throughout the species' range.
Circus cinereus 7 All sets from Chile.
Circus buoni 6 All sets but one from Argentina.
Accipiter bicolor 3 One misidentied clutch was discarded (Lloyd & Ki 1999).
Geranospiza caerulescens 5 All sets from Mexico.
Buteogallus anthracinus 100 90% from Mexico; ve clutches of “Mangrove Black Hawk”.
Buteogallus meridionalis 25 Around half from Mexico and the other half from South America.
Buteogallus urubitinga 14 Mostly from Mexico; also northern South America.
Buteogallus solitarius 1 From Mexico.
Rupornis magnirostris 142 Mostly from Mexico; others scattered throughout the species' range.
Parabuteo unicinctus 43 Mostly from Mexico.
Parabuteo leucorrhous 4 Largely increased the total number of breeding reports.
Geranoaetus albicaudatus 10 Records scattered through the species' range.
Geranoaetus polyosoma 43 Only one set from its northern range; 11 from the Falkland Islands.
Geranoaetus melanoleucus 23 All sets from its southern range.
Pseudastur albicollis 1From Trinidad.
Buteo plagiatus 104 All sets but one from Mexico.
Buteo nitidus 3All from Trinidad; seemingly no other eggs of the species are known.
Buteo brachyurus 13 All sets but one from Mexico.
Buteo galapagoensis 5 No new information added.
Buteo albonotatus 10 From its northern range.
No. of sets = number of soundly identied egg sets. Shaded cells denote any substantial addition of information, relative
to recent literature. Further explanations on the main text.
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Some aspects of the behavior of a species could be
geographically restricted (iollay 1989), and its breeding
aspects can be distinct at extreme limits of its geographical
distribution (Kennedy et al. 1995). us, generalizations
about the breeding biology of raptors become highly
susceptible to errors (Bierregaard-Jr. 1995, Trejo 2007a).
Albeit results show that the informative potential of
geographically isolated data and anecdotal descriptions
may be important contributions to our knowledge on
raptors breeding ecology (Whitacre & Burnham 2012),
we emphasize the importance of conducting detailed
studies with dierent populations.
Most recent studies that provide some new
information on breeding aspects of Neotropical
Accipitriformes are generalist in nature. e lack of detail
of anecdotal reports may be due to logistical limitations
during eld work and to the studies scope, but it is
also likely that it is often due to unawareness by local
researchers of the relevance of the material. Whichever the
reason, an emblematic outcome of this, is one occasional
report of “breeding” that, if well described, would be
the rst description on any reproductive aspect of the
Black- faced Hawk Leucopternis melanops (Cintra & Naka
2012). Because of the lack of detailed information, this
report could not be properly attributed by us to any of the
categories assessed (Table 1). Additionally, it is possible
that such lack of detail may be caused by imperfections
in the peer-review system (Figueroa, in litt.), or in
publication policies of the journals, that does not give the
opportunity to the publishing of complete information
on natural history, or disregard the value of local breeding
data.
A few of the less abundant and restricted-range
species still attract most of the attention of eld
ornithologists. Bierregaard-Jr. (1995) already remarked
on the oddness of a scarcity of breeding information for
some common species, while a few, and not necessarily
common ones (e.g., Harpy Eagle), are increasingly well
studied. For example, knowledge about the breeding
behavior of the Gray-headed Kite, a conspicuous and
widespread species (orstrom et al. 2012), is still mostly
anecdotal (Table 1, Appendix I). Figueroa (2015) stated
that among potential causes for these information gaps of
common raptors, may be the species own “commonness”,
associated with a number of other biases of research focus
in ornithology. On the other hand, knowledge of all
the former “Leucopternis” species still can be considered
the largest gap of breeding data among Neotropical
Accipitridae, from Bierregaard-Jr.'s 1995 review up to
date.
We noted that records posted in the WikiAves
database could attenuate gaps in knowledge about some
raptors in middle latitudes of South America. However,
possibly the weakest point of this database is precisely
its geographical limitation to Brazil. We believe that the
development of similar initiatives in other Neotropical
countries should be helpful as a complementary measure
to elucidate diverse information on the biology of this
region's avifauna (Lees & Martin 2014). We also stress
the importance of the use of digital records in such citizen
science tools, making possible for the researchers the
correction of misidentications. It is particularly relevant
when it comes to diurnal raptors, a group renowned for
having problematic identication in the eld (Griths
& Bates 2002, Seipke et al. 2006, 2011), leading to
errors in citizen science records (Bailey 2015) and even in
published peer-reviewed studies (de Vries & Melo 2002,
Alves et al. 2017).
We also reinforce the importance of “conventional”
records in museums (McNair 1987), as they oer the same
benets as exposed above. ey make possible to verify
previous identications (e.g., Griths & Bates 2002,
Appendix VI) and therefore prevent the perpetuation of
cascading errors. By using museum egg sets, this study
and others (Murphy 1989, Olsen & Marples 1993, Hayes
2014) also gathered breeding data that could not be
obtained from other sources, such as literature. Such fact
is clearly illustrated in the cases of taxa with substantial
increases in number of breeding records after the scrutiny
of oological collections (see Table 3).
Museum data on some diurnal raptors can yet
be very limited. For instance, we stress the need for
collecting additional information on eggs of both White-
rumped and Gray-lined Hawks, since our validation of
the identication of their museum sets must be seen as
conditional. In fact, sometimes the very same egg sets
we analyzed are the only (or at least the major) source
for egg measurements of a species provided by any
reference. In such cases, only by carefully scrutinizing all
references ever produced on a given species, and also by
examining closely-related species, it is possible to avoid
circular reasoning in validating the identication of these
eggs. Perhaps some species' eggs still are unknown, if
literature information are based in sets with questionable
identication.
We also veried that oological collections undergo
the same geographic bias found in both recent and
former (Bierregaard-Jr. 1995) literature breeding records.
Essentially the same regions (i.e., northernmost and
southernmost Neotropical countries, and the United
States) predominate with respect to amount of breeding
data. Trinidad and Tobago is an exception to this pattern,
because the work of egg collectors (e.g., Belcher &
Smooker 1934) seems to be the ultimate source of almost
all reproductive information on its raptors (Herklotts
1961, Ffrench 1991). In fact, no recent literature reference
was found for this country.
Adequate knowledge of breeding parameters is
necessary to better understand how dierent species and
populations respond to environmental changes (Marini
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et al. 2010, D'Elia et al. 2015). Such information is
particularly relevant for diurnal raptors, as they: provide
important environmental services, preying upon
potential pests and invasive species (Estes et al. 2011,
Speziale & Lambertucci 2013, Martins & Donatelli
2014); act as agship species (Sergio et al. 2008, Donázar
et al. 2016); and as indicators of environmental quality
(Jullien & iollay 1996, Blendinger et al. 2004, iollay
2007). Recent studies (e.g., Alexandrino et al. 2016) are
putting in check traditional classications of sensitivity
to disturbance, widely used for Neotropical avifauna,
such as the landmark database by Stotz et al. (1996). In
fact, despite some valuable eorts (e.g., Jullien & iollay
1996, iollay 2007), little is actually known about the
extent to which each species of Neotropical raptor ts in
the sensitivity gradient (Bierregaard-Jr. 1995, Touchton et
al. 2002, Roda & Pereira 2006).
As mentioned before, nest site choices of Accipitridae
demonstrate habitat use (Ferguson-Lees & Christie
2001), and so highlight their sensitivity to environmental
changes (Trejo 2007a). en again, recent studies
indicate a need to update classications of sensitivity to
habitat change of some Accipitridae. For example, Harpy
Eagles and Short-tailed Hawks have an alleged need for
nest sites in relatively pristine native forest (Albuquerque
1995). Yet, such allegation does not match a series of
recent breeding records that demonstrate a much greater
degree of tolerance, with successful nesting reported at
human-altered habitats (Silva 2007, Monsalvo 2012,
and references therein). ese recent reports also showed
that both prey delivery rates and edgling success in
such situations are similar or higher than those on more
pristine habitats. Nonetheless, nesting in such modied
conditions might lead to still undetected impacts, like
higher nest predation risks (Newton 2010). us, further
studies are necessary, to verify the occurrence of possible
negative eects.
Open-country raptors are generally considered
to be less threatened than forest species (e.g., Piana &
Marsden 2014), as mentioned by Bierregaard-Jr. (1995).
In fact, recent research shows that suitable habitats for
species such as the Roadside Hawk might increase with
anthropogenic changes (Carrete et al. 2009), and lead
to a substantial rise in nest productivity, in human-
modied habitats (Panasci & Whitacre 2002). On the
other hand, we also retrieved studies that claim that other
raptors of open habitats may be negatively impacted by
changes in land use. roughout the Americas, species
such as Cinereous Harriers (Circus cinereus) (Camilotti
et al. 2008), Chaco Eagles (Albuquerque et al. 2006),
and even White-tailed Hawks (Brown & Glinski 2009)
are apparently losing breeding areas. In any case, there
is a shortage of data about how environmental changes
aect the breeding of dierent species and populations.
So, for proper management of such potentially aected
populations, additional research on reproductive rates is
essential.
e relevance of studying generalist and abundant
species should not be disregarded, given the extremely
signicant participation of raptors in trophic webs
(Estes et al. 2011). Breeding range expansions have been
reported recently for some generalist species, such as some
Buteo hawks (Williams-III et al. 2007, Sandoval 2009).
ese expansions result in insertion of these raptors into
new food webs, interacting with populations of prey
species with which they had no previous contact. Some
Accipitriformes can prey upon introduced or invasive
species (Wheeler 2003, Pineda-López et al. 2012,
Martins & Donatelli 2014), and the eects of the latter
on breeding parameters of native predators still require
further research (Speziale & Lambertucci 2013). For
instance, in Snail Kite breeding areas the introduction
of an alien novel prey increased reproductive success
(Cattau et al. 2016), highlighting the ecological relevance
of raptor species.
is assessment of current knowledge of the
breeding biology of Neotropical Accipitriformes
indicated that, albeit 66% of the evaluated species had
some improvement on levels of knowledge, the scarcity
of breeding data on many South American Accipitridae
persists. Yet, we noted that records from both a citizen
science digital database and oological collections resulted
in a signicant increase in breeding information for a
total of 13 species, relative to recent literature. ere is a
persistent need for research to be conducted north of the
Southern Cone of South America, and we recommend
that breeding biology studies should focus on the 24
species selected as research priorities. Knowledge of the
breeding biology of Accipitridae not only plays a key role
in enabling proper management and conservation of their
populations. It also will point the way for more ecient
studies in the future, generating better data about the
biology of these predators and, in the nal analysis, on
the functioning of ecosystems as a whole (Bierregaard-Jr.
1995, Trejo 2007a).
ACKNOWLEDGEMENTS
J.A.B.M. and M.Â.M. thank Conselho Nacional de
Desenvolvimento Cientíco e Tecnológico (CNPq) for
their master's and researcher fellowships, respectively.
N.M.H. thanks Coordenação de Aperfeiçoamento de
Pessoal de Nível Superior (PNPD/Capes) for a post-doc
fellowship. We thank J. Holfeltz from the Peregrine
Fund for sending us pdf's, and Ryan Phillips and R.A.
Figueroa for sending us reprints and sharing information.
We also thank the latter, R.O. Bierregaard-Jr., and other
reviewers, for suggestions that greatly improved the
manuscript.
Review of breeding biology of Neotropical Accipitriformes
Monsalvo et al.
164
Revista Brasileira de Ornitologia 26(2): 2018 Revista Brasileira de Ornitologia 26(2): 2018
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Associate Editor: Gustavo S. Cabanne
APPENDIX I
Literature references with breeding data of 56 species of Neotropical Accipitriformes, produced between 1995–2016.
Species Located references
Elanus leucurus Erichsen et al. 1996; McMillian & Pranty 1997; Pranty & McMillian 1997; Sick 1997;
Arballo & Cravino 1999; Carvalho et al. 2001b; Maceda & Kin 2001; Wheeler 2003;
Antas 2004; Leveau et al. 2004; Chatellenaz 2005; de la Peña 2005; Di Giacomo 2005;
Joppert 2007; Niemela 2007; Pérez-León 2007; Scheibler 2007; Carvalho-Filho et al.
2008; Gussoni & Guaraldo 2008; González-Acuña et al. 2009; Chatellenaz et al. 2010;
Furman & Bastías 2012; Montalvo et al. 2014; Alvarado et al. 2015; Camacho-Varela
& Acosta-Chaves 2015; Romano et al. 2015; Marsden et al. 2016.
Gampsonyx swainsonii Martínez 1998; Reichle et al. 2003; Di Giacomo 2005; Jones 2005; Strewe et al. 2009;
Sandoval et al. 2010.
Chondrohierax uncinatus Ericson & Amarilla 1997; Di Giacomo 2000; orstrom et al. 2001; Clark 2002; 2003;
Krügel 2003; Reichle et al. 2003; Clark 2004; Rappole et al. 2007; Carvalho-Filho et
al. 2008; orstrom & McQueen 2008; Canuto 2009; Whitacre 2012; Sampaio et al.
2013; Phillips et al. 2015.
Leptodon cayanensis orstrom 1997; Bornschein & Reinert 2000; Carvalho-Filho et al. 2002; Cabanne
2005; Carvalho-Filho et al. 2005; Olmos et al. 2006; Carvalho-Filho et al. 2008;
Canuto 2009; Bodrati et al. 2010; Ghizoni-Jr. & Azevedo 2010; Whitacre 2012.
Leptodon forbesi Pereira et al. 2006; Dénes 2009; Dénes et al. 2011.
Elanoides forcatus Meyer & Collopy 1995; Brown et al. 1997; Gerhardt et al. 1997; Sykes-Jr. et al. 1999;
Naka & Rodrigues 2000; Coulson 2001; Blihovde 2002; Coulson 2002; Naka et al.
2002; Willis & Oniki 2002; Reichle et al. 2003; Gerhardt et al. 2004; Meyer et al.
2004; Soehren 2004; Zimmerman 2004; Azevedo & Di Bernardo 2005; Carvalho-
Filho et al. 2008; Coulson et al. 2008; Crease 2009; Gruber 2009; Lopes et al. 2009;
Whitehead & Jones 2009; Bodrati et al. 2010; Chiavacci et al. 2011; Whitacre 2012;
Carpenter & Allen 2013; Kjeldsen 2103; Enge et al. 2014.
Morphnus guianensis Whitacre et al. 2002; Mikich & Bérnils 2004; Vargas-González et al. 2006a; Raine
2007; Cintra & Naka 2012; Whitacre 2012; Crease & Tepedino 2013; Gomes 2014;
Gomes & Sanaiotti 2015; Sanaiotti et al. 2015.
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Species Located references
Harpia harpyja Chebez 1995; Alvarez et al. 1996; Alvarez-Cordero 1996; de Lucca 1996; Sick 1997;
Machado et al. 1998; Galetti & Carvalho 2000; Ibáñez et al. 2002; Piana 2002; Rettig
2002; Sanaiotti 2002; Hennessey et al. 2003; Peterson et al. 2003; Willis & Oniki
2003; Mikich & Bérnils 2004; Suárez et al. 2004; Luz 2005; Muñiz-López 2005;
Silveira et al. 2005; Olmos et al. 2006; Pereira & Salzo 2006; Vargas-González et al.
2006a; b; Giudice et al. 2007; Pacheco et al. 2007; Piana 2007; Silva 2007; Anfuso
et al. 2008; Trinca et al. 2008; Pinheiro & Dornas 2009; May 2010; Seymour et al.
2010; Sánchez-Lalinde et al. 2011; Ubaid et al. 2011; Vargas-González & Vargas 2011;
Aguiar-Silva et al. 2012; Cintra & Naka 2012; Muñiz-López et al. 2012; O'Shea &
Ramcharan 2012; Rotemberg et al. 2012; Aguiar-Silva et al. 2014; Vargas-González et
al. 2014; Aguiar-Silva et al. 2015; Kuniy et al. 2015; Sanaiotti et al. 2015; Sousa et al.
2015; Watson et al. 2016.
Spizaetus tyrannus Sick 1997; Olmos et al. 2006; Sigrist 2006; Lopes & Braz 2007; Canuto 2008;
Carvalho-Filho et al. 2008; Jones & Komar 2008a; Phillips 2009; Pimentel & Olmos
2011; Canuto et al. 2012; Cintra & Naka 2012; Whitacre 2012; Straube et al. 2014;
Meyer 2016.
Spizaetus melanoleucus Andrade et al. 1996; Sick 1997; Reichle et al. 2003; Anderson et al. 2004; Eisermann
2007; Canuto 2008; Carvalho-Filho et al. 2008; Canuto 2009; Phillips 2009; Phillips
& Seminario 2009; Bodrati et al. 2010; Canuto et al. 2012; Whitacre 2012; Kohler &
Rezini 2013.
Spizaetus ornatus Sick 1997; orstrom 1997; Andrade & Andrade 1998; Machado et al. 1998; Naveda-
Rodríguez 2002; Seipke & Cabanne 2002; Reichle et al. 2003; Greeney et al. 2004;
Mikich & Bérnils 2004; Naveda-Rodríguez 2004; Mendonça-Lima et al. 2006; Giudice
2007; Canuto 2008; Carvalho-Filho et al. 2008; Canuto 2009; Kirwan 2009; Phillips
2009; Joenck et al. 2011; Canuto et al. 2012; Cintra & Naka 2012; Whitacre 2012;
Joenck et al. 2013; Kjeldsen 2013; Phillips & Hatten 2013; Harvey et al. 2014.
Spizaetus isidori Valdez & Osborn 2002; Strewe & Navarro 2003; Valdez & Osborn 2004; Roesler et
al. 2008; Greeney et al. 2011; Castañeda 2012; Araóz & Alvedaño 2013; Zuluaga &
Echeverry-Galvis 2016.
Busarellus nigricollis Sick 1997; Di Giacomo 2000; Reichle et al. 2003; Willis & Oniki 2003; Antas
2004; Chatellenaz 2005; de la Peña 2005; Di Giacomo 2005; Márquez et al. 2005;
Chatellenaz et al. 2010; Knight 2010; Bertassoni et al. 2012; Evangelista et al. 2012.
Rostrhamus sociabilis Alvarez-López & Kattan 1995; Rodgers-Jr. 1996; Sick 1997; Valentine-Darby et al.
1997; Bennetts et al. 1998; Palmer 1998; Valentine-Darby et al. 1998; Angehr 1999;
Arballo & Cravino 1999; Bennetts & Kitchens 1999; Dreitz et al. 1999; Bennetts &
Kitchens 2000; Dreitz 2000; Dreitz & Duberstein 2001; Dreitz et al. 2001; Rodgers-Jr.
et al. 2001; Welch & Kitchens 2001; Beissinger & Snyder 2002; Bennetts et al. 2002;
Dreitz et al. 2002a; b; Petracci & Basanta 2002; Reichle et al. 2003; Rodgers-Jr. &
Schwikert 2003; Wheeler 2003; Antas 2004; Dreitz et al. 2004; Chatellenaz 2005; de
la Peña 2005; Angehr 2006; Jiménez & Zook 2007; Rodgers-Jr. 2007; Carvalho-Filho
et al. 2008; Jones & Komar 2008a; Reichert 2009; Chatellenaz et al. 2010; Palmer
2011; Bowling et al. 2012; Posso et al. 2012; Reichert et al. 2012; Román & Wiley
2012; Fortes & Denis 2013; Hernández-Vázquez et al. 2013; Bencke & Pereira 2014;
Machado et al. 2015; Cattau et al. 2016.
Helicolestes hamatus Greeney et al. 2004.
Harpagus bidentatus Schulze et al. 2000; Walther 2003; Greeney et al. 2004; Carvalho-Filho et al. 2008;
Greeney & Gelis 2008; Cintra & Naka 2012; Whitacre 2012.
Harpagus diodon Naka & Rodrigues 2000; Azevedo et al. 2003; Cabanne 2005; Azevedo et al. 2006;
Sigrist 2006; Cabanne & Roesler 2007; Carvalho-Filho et al. 2008; Canuto 2009;
Bodrati et al. 2010; Lees & Martin 2014.
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Species Located references
Ictinia plumbea Seavy et al. 1997; Sick 1997; Seavy et al. 1998; Reichle et al. 2003; Antas 2004;
Cabanne 2005; Chatellenaz 2005; de la Peña 2005; Di Giacomo 2005; Angehr 2006;
Carvalho & Bohórquez 2007; Pérez-León 2007; Carvalho-Filho et al. 2008; Gussoni
& Guaraldo 2008; Salvador-Jr. & Silva 2009; Bodrati et al. 2010; Chatellenaz et al.
2010; Jacomassa 2011; Whitacre 2012; Kjeldsen 2013; Pinto-Ledezma & Justiniano
2013; Chatellenaz 2015; Maciel et al. 2016.
Circus cinereus Saggese & de Lucca 1995; Donázar et al. 1996; Maurício & Dias 1996; Sick 1997;
Arballo & Cravino 1999; Bó et al. 2000; Jaksic et al. 2002; Bó et al. 2004; de la Peña
2005; Baladrón et al. 2007; Camilotti et al. 2008; Capllonch et al. 2011; Alvarado et
al. 2015.
Circus buoni Bó et al. 1996; Sick 1997; Arballo & Cravino 1999; Bó et al. 2004; Chatellenaz 2005;
Carvalho-Filho et al. 2008; Kirwan & Shirihai 2008; Chatellenaz et al. 2010; Alvarado
et al. 2015.
Accipiter poliogaster de Vries & Melo 2000; 2002; orstrom 2002a; Bodrati et al. 2010; Lima & Ribeiro
2011; Boesing et al. 2012.
Accipiter superciliosus Hennessey et al. 2003; iollay 2007; Carvalho-Filho et al. 2008; Bodrati et al. 2010.
Accipiter collaris Cuervo et al. 2008.
Accipiter gundlachi Rompré et al. 1999; Wallace et al. 1999; Peña et al. 2012; Ferrer-Sánchez & Rodríguez-
Estrella 2014; Ferrer-Sánchez 2015; Ferrer-Sánchez & Rodríguez-Estrella 2016.
Accipiter bicolor Pavez & González 1998; orstrom & Ki 1999; orstrom & Quixchán 2000; Reid et
al. 2002; Figueroa et al. 2004a; b; Mikich & Bérnils 2004; Ojeda et al. 2004; Carvalho-
Filho et al. 2005; Figueroa et al. 2007; Marini et al. 2007; Azpiroz & Menéndez 2008;
Carvalho-Filho et al. 2008; Canuto 2009; Bodrati et al. 2010; Zorzin 2011; Whitacre
2012; Hayes 2014; Alvarado et al. 2015; Medel-Hidalgo et al. 2015; Rivas-Fuenzalida
2015b; Rivas-Fuenzalida et al. 2015c.
Geranospiza caerulescens Sick 1997; Arballo & Cravino 1999; Sutter et al. 2001; del Ángel 2002; Reichle et
al. 2003; Chatellenaz 2005; Sigrist 2006; Carvalho-Filho et al. 2008; Canuto 2009;
Whitacre 2012.
Buteogallus anthracinus Barrantes 1998; Boal 2001; Barradas-García et al. 2004; Márquez et al. 2005; Barradas-
García & Morales-Mávil 2007; Clark 2007b; Pérez-León 2007; Flesch 2008; Sadoti
2008; Troy & Stahlecker 2008; Flesch 2009; Ruvalcaba-Ortega & González-Rojas
2009; Alava et al. 2011; Sadoti 2012; Uribe-Hernández et al. 2012; Etzel et al. 2014;
Smith & Finch 2014; Licence & McCarty 2015.
Buteogallus aequinoctialis Mikich & Bérnils 2004.
Buteogallus meridionalis Narozky & Martelli 1995; Best et al. 1996; Sick 1997; Andrade & Andrade 1998;
Arballo & Cravino 1999; Reichle et al. 2003; Antas 2004; Chatellenaz 2005; de la Peña
2005; Di Giacomo 2005; Navarro et al. 2007; Carvalho-Filho et al. 2008; Strewe et
al. 2009; Chatellenaz et al. 2010; Marini et al. 2012; Maurício et al. 2013; Camacho-
Varela et al. 2015; Silva & Machado 2015.
Buteogallus lacernulatus Carvalho-Filho et al. 2008; Canuto 2009.
Buteogallus urubitinga Best et al. 1996; Seavy & Gerhardt 1998; Arballo & Cravino 1999; Di Giacomo 2000;
Naveda-Rodríguez 2002; Reichle et al. 2003; Antas 2004; Naveda-Rodríguez 2004;
Chatellenaz 2005; de la Peña 2005; Di Giacomo 2005; Carvalho-Filho et al. 2006;
Carvalho-Filho et al. 2008; Canuto 2009; Chatellenaz et al. 2010; Whitacre 2012;
Kjeldsen 2013.
Buteogallus solitarius Mee et al. 2002; Strewe & Navarro 2003; Jones 2005; Clark 2007a; Seminario et al.
2011; Phillips 2012; Phillips & Martinez 2013; Phillips et al. 2014.
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Species Located references
Buteogallus coronatus Sick 1997; Bellocq et al. 1998; Machado et al. 1998; Carvalho et al. 2002; Maceda et
al. 2003; Mikich & Bérnils 2004; de la Peña 2005; Di Giacomo 2005; Albuquerque
et al. 2006; Barcellos & Accordi 2006; Granzinolli et al. 2006; Torres et al. 2006;
Bragagnolo et al. 2007; Lobos et al. 2007; Maceda 2007; Maceda et al. 2007; Carvalho-
Filho et al. 2008; Tizianel 2008; Chiaravalloti et al. 2009; Sarasola et al. 2010; Banhos
& Sanaiotti 2011; Lobos et al. 2011; Berkunsky et al. 2012; Fandiño & Pautasso 2013;
Urios et al. 2014; Kilpp 2015; Montalvo et al. 2015; Barbar et al. 2016.
Morphnarchus princeps Sánchez & Sánchez-M. 2002; Muela & Valdez 2003; Márquez et al. 2005; Greeney &
Nunnery 2006; Gelis & Greeney 2007; Greeney et al. 2008.
Rupornis magnirostris Best et al. 1996; Capllonch 1997; Maragliano & Montalti 1997; Arballo & Cravino
1999; Naka & Rodrigues 2000; Panasci & Whitacre 2000; Carvalho et al. 2001a;
Höing & Camargo 2002; Naka et al. 2002; Panasci & Whitacre 2002; Reichle et al.
2003; Antas 2004; Bó et al. 2004; Chatellenaz 2005; de la Peña 2005; Di Giacomo
2005; Marini et al. 2007; Navarro et al. 2007; Carvalho-Filho et al. 2008; Gussoni &
Guaraldo 2008; Salvador-Jr. & Silva 2009; Santos & Rosado 2009; Santos et al. 2009;
Verea et al. 2009; Bodrati et al. 2010; Chatellenaz et al. 2010; Cavicchia & García
2012; Cintra & Naka 2012; Mojica 2012; Panasci 2012; Panasci unpub. data apud
GRIN 2012; Uribe-Hernández et al. 2012; Romano et al. 2015.
Parabuteo unicinctus Blue 1996; Silva & Olmos 1997; Arballo & Cravino 1999; Gerstell & Bednarz 1999;
Patten & Erickson 2000; Maceda & Kin 2001; Willis & Oniki 2003; de la Peña 2005;
Márquez et al. 2005; Dwyer 2006; Figueroa & González-Acuña 2006; Jenner et al.
2007; Pérez-León 2007; Dwyer & Mannan 2009; Ellis et al. 2009; Cavicchia & García
2012; Furman & Bastías 2012; Alvarado et al. 2015.
Parabuteo leucorrhous Freile & Chaves 2000; Mikich & Bérnils 2004; Greeney & Nunnery 2006; Tobias &
Seddon 2007; Zilio & Mendonça-Lima 2012.
Geranoaetus albicaudatus Sick 1997; Bellatti 2000; Granzinolli 2003; Reichle et al. 2003; Di Giacomo 2005;
Granzinolli & Motta-Junior 2006; Granzinolli et al. 2006; Actkinson et al. 2007;
Granzinolli & Motta-Junior 2007; Rappole et al. 2007; Carvalho-Filho et al. 2008;
Haralson 2008; Actkinson et al. 2009; Brown & Glinski 2009; Salvador-Jr. & Silva
2009; Motta-Junior et al. 2010; Greeney et al. 2011; Maurício et al. 2013.
Geranoaetus polyosoma Jiménez 1995; Donázar et al. 1996; Jaksic & Lazo 1999; Bó et al. 2004; de la Peña
2005; Alvarado & Figueroa 2006a; Cabot & de Vries 2009; Cabot et al. 2010a; b;
Greeney et al. 2011; Hahn et al. 2011; Lüthi 2011; Alvarado et al. 2015; Shirihai et al.
2015.
Geranoaetus melanoleucus de Lucca & Saggese 1995; Hiraldo et al. 1995; Narozky & Martelli 1995; Best et
al. 1996; Donázar et al. 1996; Sick 1997; Arballo & Cravino 1999; Jaksic & Lazo
1999; Sousa 1999; Bellatti 2000; Pavez 2001; Saggese & de Lucca 2001; Bencke et al.
2003; de la Peña 2005; Trejo et al. 2006b; Zorzin et al. 2007; Salvador-Jr. et al. 2008;
Chatellenaz et al. 2010; Arriagada et al. 2011; Lüthi 2011; de Lucca & Saggese 2012;
Alvarado et al. 2015; Ignazi 2015; Pérez 2015; Raimilla et al. 2015; Lemos 2016.
Pseudastur polionotus Willis & Oniki 2002; Bencke et al. 2003; Corrêa et al. 2008; Canuto 2009.
Pseudastur albicollis Draheim 1995; Cisneros-Heredia 2006; Cintra & Naka 2012; Whitacre 2012.
Pseudastur occidentalis Vargas 1995; Best et al. 1996.
Leucopternis semiplumbeus Ferguson-Lees & Christie 2001.
Leucopternis melanops Ferguson-Lees & Christie 2001; Cintra & Naka 2012.
Leucopternis kuhli Kirwan 2009.
Buteo plagiatus Bibles & Mannan 2004; Werner 2004; Patrikeev 2007; Rappole et al. 2007; Flesch
2008; Flesch & Saavedra 2008; Flesch 2009; Ruvalcaba-Ortega & González-Rojas
2009; Sandoval 2009; Vargas-Masís & Ramírez 2012.
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Species Located references
Buteo nitidus Sick 1997; Reichle et al. 2003; Navarro et al. 2007; Sandoval 2009; Strewe et al. 2009;
Cintra & Naka 2012.
Buteo ridgwayi orstrom 2002b; orstrom et al. 2005; 2007; Woolaver 2011; Woolaver et al. 2013a,
b, c, Woolaver et al. 2015.
Buteo albigula Gelain et al. 2001; Trejo et al. 2001; Ojeda et al. 2003; Pavez et al. 2004; Trejo et al.
2004; Trejo et al. 2006a; Silva-Rodríguez et al. 2008; Henry & Aznar 2009; Rivas-
Fuenzalida et al. 2013; Alvarado et al. 2015; Rivas-Fuenzalida et al. 2015b.
Buteo brachyurus Naka & Rodrigues 2000; Carvalho et al. 2001a; Jones 2002; Wheeler 2003; Meyer
2004, 2005; Meyer & Zimmerman 2007; Rappole et al. 2007; Williams-III et al. 2007;
Brush 2008; Carvalho-Filho et al. 2008; Flesch 2008; Rizkalla et al. 2009; Salvador-
Jr. & Silva 2009; Howell 2010; Snyder et al. 2010; Monsalvo 2012; Enge et al. 2014;
Straube et al. 2014; Oliveira et al. 2015; FWC [s.d.].
Buteo galapagoensis Faaborg et al. 1995; DeLay et al. 1996; Bollmer et al. 2003; Whiteman & Parker 2004a;
b; Bollmer et al. 2005; Jaramillo & Vargas 2010; Rivera et al. 2011; Muñoz 2012.
Buteo albonotatus Kennedy et al. 1995; Sick 1997; Pérez-León 2007; Carvalho-Filho et al. 2008; Flesch
2008; Howell 2010; Olmos & Albano 2012.
Buteo ventralis Figueroa et al. 2000; Imberti 2003; Rivas-Fuenzalida et al. 2009, 2011; Norambuena
et al. 2012; Medel-Hidalgo et al. 2013; Norambuena et al. 2013; Raimilla et al. 2013;
Rivas-Fuenzalida & Asciones-Contreras 2013; Figueroa unpub. data apud GRIN
2015; Rivas-Fuenzalida 2015a; Rivas-Fuenzalida & Asciones-Contreras 2015; Rivas-
Fuenzalida et al. 2015a, 2016.
APPENDIX II
Complete list of references retrieved in this review and cited in Appendix I.
Actkinson M.A., Kuvlesky-Jr. W.P., Boal C.W., Brennan L.A. &
Hernandez F. 2007. Nesting habitat relationships of sympatric
Crested Caracaras, Red-tailed Hawks, and White-Tailed Hawks
in south Texas. Wilson Journal of Ornithology 119: 570–578.
Actkinson M.A., Kuvlesky-Jr. W.P., Boal C.W., Brennan L.A. &
Hernandez F. 2009. Comparison of the breeding biology of
sympatric Red-Tailed Hawks, White-Tailed Hawks, and Crested
Caracaras in south Texas. Journal of Raptor Research 43: 50–56.
Aguiar-Silva F.H., Junqueira T.G., Sanaiotti T.M., Guimarães V.Y.,
Mathias P.V.C. & Mendonça C.V. 2015. Resource availability
and diet in Harpy Eagle breeding territories on the Xingu River,
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APPENDIX III
Results of the search for literature breeding data of two species of Accipitriformes not presented on Bierregaard-Jr.'s (1995)
review.
Sharp-shinned Hawk Accipiter striatus – the vast majority of breeding records of the so-called “Central and South
American group” of subspecies (sensu Ferguson-Lees & Christie 2001) refer to A. s. erythronemius, whose breeding traits were
classied as entirely unknown by Bierregaard-Jr. (1995; but see comments by Di Giacomo 2005). Dierent populations
of this subspecies' range were studied in detail, but most other breeding reports are anecdotal. Central American A. s.
chionogaster (also labeled as having unknown breeding biology by 1995) now at least had one of its populations studied
in detail. Finally, the Andean form A. s. ventralis have no new breeding data; its nest remains undescribed, and knowledge
on breeding behavior is based solely on older scattered information (Bierregaard-Jr. 1995).
Located references:
Arballo E. & Cravino J.L. 1999. Aves del Uruguay: manual ornitológico,
v. 1. Montevideo: Editorial Hemisferio Sur.
Bodrati A., Cockle K., Segovia J.M., Roesler I., Areta J.I. & Jordan
E. 2010. La avifauna del Parque Provincial Cruce Caballero,
provincia de Misiones, Argentina. Cotinga 32: 41–64.
Carvalho C.E.A., Carvalho-Filho E.P.M. & Carvalho G.D.M. 2002.
Descripción de nidos, huevos, pichones y aspectos de la biologia
reproductiva del Gavilan Muslirrufo (Acciptier(sic) erythronemius)
en el Sureste de Brasil. Panamá: Conferencia Sobre Aves Rapaces
Neotropicales y Simposio del Águila Arpía.
Carvalho-Filho E.P.M., Carvalho G.D.M. & Carvalho C.E.A. 2005.
Observations of nesting Gray-Headed Kites (Leptodon cayanensis)
in southeastern Brazil. Journal of Raptor Research 39: 89–92.
Carvalho-Filho E.P.M., Zorzin G., Canuto M., Carvalho C.E.A.
& Carvalho G.D.M. 2008. Aves de rapina diurnas do Parque
Estadual do Rio Doce, Minas Gerais, Brasil. MG Biota 1: 4–43.
Chatellenaz M.L. 2005. Aves del Valle del Río Paraná en la provincia
del Chaco, Argentina: riqueza, historia natural y conservación.
INSUGEO 14: 527–550.
Di Giacomo A.G. 2005. Aves de la Reserva El Bagual, p. 201–465.
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y paisaje de la Reserva El Bagual, provincia de Formosa, Argentina:
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protegida del chaco húmedo. Buenos Aires: Asociación Ornitológica
del Plata.
Jenner T. 2008. Accipiter chionogaster. Aratinga 2: 4–5.
Jenner T. 2010. Life history of the White-Breasted Hawk (Accipiter
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Neotropical 13: 273–282.
Seipke S.H. & Cabanne G.S. 2008. Breeding of the Rufous-
ighed Hawk (Accipiter erythronemius) in Argentina and Brazil.
Ornitología Neotropical 19: 15–29.
Willis E.O. & Oniki Y. 2002. Birds of Santa Teresa, Espírito Santo,
Brazil: do humans add or subtract species? Papéis Avulsos de
Zoologia 42: 193–264.
Willis E.O. & Oniki Y. 2003. Aves do estado de São Paulo. Rio Claro:
Divisa Editora.
Cuban Black Hawk Buteogallus gundlachii – apparently there is still little breeding data, as we located very few reports,
and just two of these studies provide more detailed descriptions of breeding events.
Located references:
Ferrer-Sánchez Y. 2015. Variables que inuyen em la distribución y
abundancia de rapaces diurnas y em la ubicación de sus sitios de
anidación em Cuba. Ph.D. esis. La Paz: Centro de Investigaciones
Biológicas del Noroeste.
Ferrer-Sánchez Y. & Rodríguez-Estrella R. 2016. How rare species
conservation management can be strengthened with the use of
ecological niche modelling: the case for endangered endemic
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García-Quintas A. & Ávila D.D. 2012. Un ejemplo teórico de
modelación del hábitat y la distribución potencial por análisis
factorial del nicho ecológico. Mesoamericana 16: 12–21.
Wiley J.W. & Garrido O.H. 2005. Taxonomic status and biology of
the Cuban Black-Hawk, Buteogallus anthracinus gundlachii (Aves:
Accipitridae). Journal of Raptor Research 39: 351–364.
Review of breeding biology of Neotropical Accipitriformes
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APPENDIX IV
Literature references with breeding data of Neotropical Accipitriformes, produced between 1995‒2016, but not retrieved
in this review.
Altamirano T.A., Ibarra J.T., Hernandez-Rojas I., Laker J. & Bonacic
C. 2012. [Nesting habits of the birds of the Andean temperate forests
of Chile]. Santiago: Fondo de Proteccion Ambiental.
Alvarado S.A. & Figueroa R.A. 2006b. Function of reverse sexual
dimorphism in the reproductive behavior and parental care of
the Chilean Hawk (Accipiter chilensis), in the Nevados de Chillan
biological corridor, Chile. In: Valparaiso: VIII Jornadas de Etologia
de la Universidad de Playa Ancha.
Anderson D.L. 1999. Tawahka Project, Honduras: 1999 eld season
report. Boise: e Peregrine Fund.
Couve E. & Vidal C.F. 2004. Birds of Torres Paine National Park,
Patagonia, Chile. Puntarenas: Editorial Fantastico Sur.
Donaghy Cannon M. 2001. Breeding ecology of cooperatively
polyandrous Galapagos Hawks (Buteo galapagoensis) on Santiago
Island, Galapagos. MSc. Dissertation. Jonesboro: Arkansas State
University.
Figueroa R.A., Corales S.S. & Lopez R.R. 2001. Records of the
White-throated Hawk (Buteo albigula) and notes on its hunting
methods and movements in the Andes of central-southern Chile.
International Hawkwatcher 4: 3–9.
Giudice R. 2006. Tree architecture as a determinant factor in the nest
tree selection of Harpy Eagles (Harpia harpyja). In: Iguazu: II
Neotropical Raptor Conference.
Jones H.L. 2003. Central America. North American Birds 57: 414–
416.
Jones HL. 2004. Central America. North American Birds 58: 290–292
Jones H.L. & Komar O. 2008b. Central America. North American
Birds 61: 648–651.
Levenstein K. 2008. Reproductive ecology of the cooperatively polyandrous
Galapagos Hawk on Santiago Island, Galapagos. Ph.D. esis.
Jonesboro: Arkansas State University.
Lobos R.P. & Alvarado O.S. 2006. Mutualism between the Crowned
Solitary Eagle (Harpyhaliaetus coronatus) and the Monk Parakeet
(Myiopsitta monacha) during nest building in the Telteca Natural
Forest Reserve, Department of Lavalle, Mendoza province,
Argentina. Valparaiso: VIII Jornadas de Etologia de la Universidad
de Playa Ancha.
Meyer K.D. & Arnett J.E. 1996. Age-class distinctions and delayed
reproduction of American Swallow-tailed Kites in Florida. Boise:
Abstracts of the 114th Stated Meeting of the American Ornithologists'
Union and the 1996 Annual Meeting of the Raptor Research
Foundation.
Meyer K.D., Duvall D.J. & Arnett J.E. 1995. Depressed success of
American Swallow-tailed Kites (Elanoides forcatus) nesting in
introduced Australian Pines (Casuarina spp.). Duluth: Abstracts of
the Raptor Research Foundation 1995 Annual Meeting.
Olveira L. 2001. Rufous-thighed Hawk (Accipiter erythronemius) in
Mar del Plata, Buenos Aires, Argentina. Nuestras Aves 17: 34.
Pineda L., Navas E.M. & Fernandez R.A. 2016. New location for
and rst record of nesting Pearl Kite (Gampsonyx swainsoni) in El
Salvador. Spizaetus 22: 6–13.
Sánchez T. 2000. [Behavior of the Galapagos Hawk (Buteo
galapagoensis) during the incubation period and morphological
variation between populations at Espanola, Santa Fe, Isabela and
Antiago(sic) Islands, Galapagos, Ecuador]. Undergraduate esis.
Quito: Ponticia Universidad Católica del Ecuador.
Tuño P. 2007. Cunsi Pindo: conservation of the Harpy Eagle in Ecuador.
Quito: SIMBIOE.
Woods R.W. & Woods A. 1997. Atlas of breeding birds of the Falkland
Islands. Shropshire: Anthony Nelson.
Review of breeding biology of Neotropical Accipitriformes
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APPENDIX V
Results of the search for photographic breeding records of Neotropical Accipitriformes on the WikiAves database.
Species Records' reference numbers
Elanus leucurus WA1251178; WA1253853; WA1263279; WA1272398; WA1272409;
WA1279861; WA1281964; WA1288071; WA1290781; WA1293418;
WA1300395; WA1376684; WA1499798; WA1720428; WA1721934;
WA1770437; WA1835635; WA1835637; WA1837761; WA1841067;
WA1904171; WA2037143; WA2071064; WA2090188; WA21325; WA21537;
WA2271383; WA250965; WA466357; WA661980; WA698506; WA719423;
WA729366; WA732823; WA915133; WA915840; WA915852; WA936035.
Chondrohierax uncinatus WA1160532; WA1688095; WA1937776; WA1968066; WA1981003.
Leptodon cayanensis WA723947; WA723948.
Leptodon forbesi WA938449.
Spizaetus melanoleucus WA1140737; WA1140739; WA1370302; WA1378059; WA1438023;
WA195643; WA2206395; WA2242350; WA2249207; WA2322423.
Rostrhamus sociabilis WA1214147; WA1218422; WA1280372; WA147627; WA1493450;
WA1588325; WA1771055; WA2021254; WA2021256; WA2108507;
WA226747; WA24193; WA36753; WA474247; WA484024; WA64884;
WA696195; WA696196; WA81214; WA819399.
Helicolestes hamatus WA1589021; WA1966794; WA953944.
Harpagus bidentatus WA2198552; WA2240795; WA668871.
Harpagus diodon WA1156861; WA1200479; WA1228366; WA123732; WA1237599; WA14961;
WA14962; WA1966820; WA1966889; WA206624; WA209513; WA219297;
WA219978; WA222095; WA222762; WA244381; WA250110; WA251551;
WA252886; WA255778; WA255779; WA255914; WA257012; WA275906;
WA280598; WA507006; WA73820; WA76435; WA76436; WA76815;
WA785304; WA819506; WA82627; WA860802; WA861618; WA884512;
WA887671; WA887710; WA889684; WA897892; WA900167; WA98349.
Accipiter poliogaster WA1920902; WA1985763; WA1989199; WA1992309; WA1994808;
WA2005934; WA2034929; WA2047459; WA2132296; WA2319849;
WA779787.
Accipiter bicolor WA106136; WA1744297; WA89938.
Geranospiza caerulescens WA140630; WA141005; WA1444043; WA1565980; WA1649149.
Buteogallus anthracinus WA950092.
Buteogallus aequinoctialis WA1503515.
Pseudastur polionotus WA1570081; WA1570097; WA1581106.
Pseudastur albicollis WA215803; WA722126.
Buteo nitidus WA1184610; WA1392108; WA2187978; WA2187993; WA388429;
WA476978; WA506191.
Buteo brachyurus WA1116480; WA1356894; WA1356902; WA176090; WA176091;
WA2033914; WA225567; WA33877; WA513759; WA513770; WA513777;
WA513781; WA513790; WA513819; WA513828; WA819112; WA819113.
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Revista Brasileira de Ornitologia 26(2): 2018
APPENDIX VI
Proposed corrections to four misidentied museum egg sets of Neotropical Accipitriformes. Arguments referring to
geographical distribution are not presented since all species involved are sympatric at these collection localities (del Hoyo
et al. 2016).
Set WFVZ 15561 - formerly assigned to Lined Forest-Falcon Micrastur gilvicollis. Seemingly, no information exists on
Lined Forest-Falcon's eggs (Bierregaard-Jr. 1995, GRIN 2009, del Hoyo et al. 2016). is one-egg set was obtained by G.
D. Smooker, whose identications have already been questioned (orstrom & Ki 1999). More importantly, the egg is
much larger than those of another similar-sized, closely-related Micrastur falcon (Whitacre 2012). us, we doubt it could
be properly atributed to Lined Forest-Falcon.
Measurements, clutch-size and overall appearance are suitable with known clutches of the Gray-headed Kite measured by
us and to other data presented by Whitacre (2012). us, it almost certainly belongs to this species.
We recommend the treatment of this set as cfr. Leptodon cayanensis.
Set WFVZ 15951 - previously assigned to Black-collared Hawk Busarellus nigricollis. Also from Smooker's collection.
Measurements of theses two eggs are much smaller than Black-collared Hawk's eggs (GRIN 2010), but consistent with
those of Zone-tailed Hawk Buteo albonotatus (del Hoyo et al. 2016), as suggested by L. Ki on the data slip of this set. Yet,
contrary to the previous and next cases, these species overall appearances and “eld jizzes” are quite dierent (J.A.B.M.,
pers. obs.) to justify such a misidentication by the collector. Also, dimensions, clutch-size and general appearance of the
eggs did not allow a rigorous identication. We do not discard that the clutch refers to Zone-tailed Hawk, but evidence is
not conclusive as they may refer to other hawks as well.
We recommend that this set should not be treated as Busarellus nigricollis, and tentatively identify as cfr. Buteo albonotatus.
Sets WFVZ 16312 and 16313 - both formerly assigned to Hook-billed Kite Chondrohierax uncinatus.
ese three eggs are very distinct from, and much larger than, Hook-billed Kite's (J.A.B.M., pers. obs., Di Giácomo
2000, Whitacre 2012). Both dimensions, clutch-sizes and overall appearance ts with Gray-headed Kite's clutches. Albeit
measurements of the two-egg clutch (WFVZ 16312) are slightly smaller than most Gray-headed Kite's, they t with those
of another two egg-clutch of this species, provided by Carvalho-Filho et al. (2005).
We assign these sets to Leptodon cayanensis.
References:
Bierregaard-Jr. R.O. 1995. e biology and conservation status of
Central and South American Falconiformes: a survey of current
knowledge. Bird Conservation International 5: 325–340.
Carvalho-Filho E.P.M., Carvalho G.D.M. & Carvalho C.E.A. 2005.
Observations of nesting Gray-Headed Kites (Leptodon cayanensis)
in southeastern Brazil. Journal of Raptor Research 39: 89–92.
del Hoyo J., Elliott A., Sargatal J., Christie D.A. & de Juana E. 2016.
Handbook of the birds of the world alive. http://www.hbw.com
(Access on 24 October 2016).
Di Giacomo A.G. 2000. Nidicación de algunas rapaces poco
conocidas en el Chaco oriental argentino. Hornero 15: 135–139.
GRIN (Global Raptor Information Network). 2009. Species account:
Lined Forest Falcon Micrastur gilvicollis. http://www.globalraptors.
org/grin/SpeciesResults.asp?specID=8062 (Access on 24 October
2016).
GRIN (Global Raptor Information Network). 2010. Additional details
on Breeding. Busarellus nigricollis. http://www.globalraptors.org/
grin/SpeciesExtended.asp?specID=8010&catID=2006 (Access on
24 October 2016).
orstrom R. & Ki L. F. 1999. Notes on eggs of the Bicolored Hawk
Accipiter bicolor. Journal of Raptor Research 33: 244–247.
Whitacre D.F. 2012. Neotropical birds of prey: biology and ecology of a
forest raptor community. Ithaca: Cornell University Press.
