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Feeding Ecology of Military Macaws (Ara militaris) in a Semi-Arid Region of

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DOI: 10.1676/08-034.1

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 Feeding Ecology of Military Macaws (Ara militaris) in a Semi-Arid Region of
Central México
Author(s): A. M. Contreras-González , F. A. Rivera-Ortíz , C. Soberanes-González , A. Valiente-Banuet ,
and M. C. Arizmendi
Source: The Wilson Journal of Ornithology, 121(2):384-391. 2009.
Published By: The Wilson Ornithological Society
DOI: http://dx.doi.org/10.1676/08-034.1
URL: http://www.bioone.org/doi/full/10.1676/08-034.1

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The Wilson Journal of Ornithology 121(2):384–391, 2009

FEEDING ECOLOGY OF MILITARY MACAWS (ARA MILITARIS) IN

A SEMI-ARID REGION OF CENTRAL MÉXICO

A. M. CONTRERAS-GONZÁLEZ,1 F. A. RIVERA-ORTÍZ,1
C. SOBERANES-GONZÁLEZ,1 A. VALIENTE-BANUET,2 AND M. C. ARIZMENDI1,3

ABSTRACT.—The Military Macaw (Ara militaris) is an endangered species with a fragmented distribution
and declining populations. Their diet has not been described in detail but the species is considered to be a
specialized granivore, and individuals may make seasonal movements in search of food resources. This study
examined the diet of Military Macaws in tropical dry and oak (Quercus spp.) forests of Cuicatlán, México. We
followed macaws to identify diet composition, and quantified the nutritional content of all items which they
consumed. Military Macaws used 10 plant species during the year having a standardized Levins niche breath
index of B ⫽ 0.18. Their diet consisted principally of seeds but also included fruits, leaves, and latex. Foods
rich in protein and lipids composed the diet of macaws, especially during the breeding season. Macaws preferred
some plant species using them more than predicted by their abundances. Food resources must be considered
when managing and preserving habitat to protect this endangered species. Received 25 February 2008. Accepted
30 September 2008.

Military Macaws (Ara militaris) occur in
Mexico and northern South America (Forshaw
1989). In Mexico, the species is currently re-
stricted to a few isolated populations in semi-
deciduous and deciduous forests (Forshaw
1989, Carreón 1997, Collar 1997, Loza 1997),
making occasional seasonal altitudinal move-
ments to pine (Pinus spp.) and pine-oak
(Quercus spp.) forests (Forshaw 1989, Peter-
son and Chaliff 1989, Collar and Juniper
1992, Howell and Webb 1995). The species is
classified under Appendix I of the Convention
on International Trade in Endangered Species
(CITES 1998), and considered Vulnerable in
the Red List of the International Union for the
Conservation of Nature (SEMARNAT 2001,
Birdlife International 2003). The species is
considered in danger of extinction in Mexico,
principally due to habitat loss and trade pres-
sures (Iñigo-Elias 2000).
Macaws are considered to have narrow di-
ets using only between 10 and 23% of the
available resources throughout the year (Car-
reón 1997, Loza 1997, Iñigo-Elias 1999).
Adult Scarlet Macaws (Ara macao), for ex-
1 Laboratorio de Ecologı́a, UBIPRO, FES Iztacala
UNAM, Avenida de los Barrios 1, Los Reyes Iztacala,
Tlanepantla, Estado de México 05490.
2 Instituto de Ecologı́a, UNAM, Circuito Exterior s/n,
Ciudad Universitaria, Coyoacán, México D.F. 04510.
3 Corresponding author; e-mail:
coro@servidor.unam.mx
384
ample, used between 12 and 15 plant species
in Belize (Renton 2006). The same species
used 52 plant species in the Amazonian rain-
forest (Gilardi 1996), while in tropical dry for-
est in Peru they only used 28 species for food
(Marineros and Vaughan 1995). Military Ma-
caws in a semi-deciduous forest in western
Mexico used six species for food that repre-
sented 10% of the available resources (Loza
1997).
A narrow diet makes populations vulnera-
ble to changes in food resource availability
associated with fragmentation and other forms
of habitat loss, while a varied opportunistic
diet can allow populations to adapt to habitat
changes (Saunders 1977, Renton 2006). The
objectives of our study were to describe: (1)
the dietary composition of Military Macaws
during the year, and (2) the nutritional com-
position of foods eaten.
METHODS
Study Area.—Field work was conducted at
two sites in the Cuicatlan Basin in the State
of Oaxaca, Mexico. The first was in Sabino
Canyon (17⬚ 51⬘ 44.60⬙ N, 97⬚ 2⬘ 43.89⬙ W)
where a colony of 70–90 Military Macaws
roosts and nests in the vertical walls of the
canyon (Salazar 2001, Rivera-Ortı́z et al.
2008). The canyon occurs at 610–885 m asl
and is dominated by tropical deciduous forest
with columnar cacti (Valiente-Banuet et al.
 Contreras-González et al. • DIET OF MILITARY MACAWS IN MÉXICO
2000). The mean annual rainfall is between
304 and 432 mm, and the mean annual tem-
perature is 32⬚ C. The rainy season occurs
from June to September, and the dry season
lasts up to 8 months (Rivera-Ortı́z et al. 2008).
The second site was San Juan Coyula, be-
tween 17⬚ 54⬘ 47.45⬙ N and 96⬚ 55⬘ 42.64⬙ W
with an elevation of 1,400–1,610 m asl. The
main vegetation is oak forest (Valiente-Banuet
et al. 2000). The mean annual temperature is
between 16.9 and 18.7⬚ C, and mean annual
rainfall is 1,114 to 1,201 mm. The foothills of
this area are dominated by tropical dry forest,
which was also studied at el Aguacate, La
Chirimoya, and la Mulatera. These foothill
sites were between 17⬚ 55⬘ 43.50⬙ N and 96⬚
56⬘ 11.38⬙ W at 1,062 and 1,216 m asl. The
mean annual rainfall in this area is between
650 and 890 mm, and mean annual tempera-
ture is between 19 and 23⬚ C (Rivera-Ortı́z et
al. 2008).
Food Availability.—We conducted monthly
phenological observations between March
2005 and March 2006. We randomly estab-
lished four 50 ⫻ 20 m plots in Santa Marı́a
Tecomavaca, and three plots each in oak and
tropical dry forest vegetation types in San
Juan Coyula that were surveyed each month.
Area surveyed was selected after performing
a minimum area test. All trees, shrubs, and
cacti ⬎5 cm diameter at breast height (DBH)
were marked in each plot and numbers of
fruits were recorded for each month through-
out the year. A series (5–10) of samples (i.e.,
branches) were counted for trees with abun-
dant fruits, and the number of total branches
was counted to calculate the total number of
fruits (Chapman et al. 1992, Borchert 1994).
Relative abundance of trees in plots was cal-
culated for trees where macaws fed during the
year. Relative abundance of plants and fruits
were used to identify plant preferences.
Feeding Biology.—Observations of feeding
macaws were conducted by walking on and
off trails. We recorded macaws feeding by
date, location, habitat type, plant species,
plant part consumed, number of individuals
foraging, number of fruits or seeds consumed,
and time taken per feeding observation. We
also recorded other plant parts consumed. A
feeding observation ended when macaws
ceased foraging and moved; observers then
continued walking and searching for addition-
385
al foraging activities in the area. Observations
were made by two or three observers that fol-
lowed individuals and counted ingested items.
Trees were identified by collection of speci-
mens for comparison at the Herbarium of FES
Iztacala, UNAM.
Nutritional Quality.—Samples of food
items consumed were collected and placed on
ice for transportation to the laboratory. We
quantified total protein using micro-Kjeldahl
procedures (Izaki 1993, González and Peña-
losa 2000, Levey 2000), total carbohydrates
by Cleggs Antrone procedure (Osborne 1986),
total lipids using Soxhlet apparatus (González
and Peñalosa 2000), and water content by sub-
traction between fresh and dry weight.
Data Analysis.—All data were tested for
normality (Kolmogorov-Smirnov test) (Sokal
and Rohlf 1979, Zar 1999). Variation in fruit
abundance during the year was analyzed using
a non-parametric repeated measures Friedman
test. We used a log-linear multiple contingen-
cy analysis to test for preferences for one or
more of the consumed species by vegetation
type, assuming that birds consume fruits ac-
cording to their relative abundance at the site
(Everitt 1977, Sokal and Rohlf 1979, Zar
1999).
Niche breadth for Military Macaw diet was
calculated using the standardized Levins niche
breadth index (Krebs 1989), where a value
close to zero indicates dietary specialization
and a value close to one indicates a broad diet
(Colwell and Futuyma 1971). Differences in
foraging time on different species of plants
consumed by Military Macaws and seed con-
sumption rates were analyzed using one-way
ANOVA (Sokal and Rohlf 1979, Zar 1999).
RESULTS
Temporal Variation in Food Resources.—
There was significant variation in abundance
of food items among months during the year
(Fig. 1). The abundance of food resources for
macaws in the tropical dry forest of Santa Ma-
rı́a Tecomavaca was high during the dry sea-
son in February and March (Fig. 1). Signifi-
cant differences were found between months
in the number of fruits throughout the year
(Friedman test, Xr2 ⫽ 11.00, df ⫽ 1, P ⫽
0.01). Peak fruiting in the oak forest occurred
in September (Fig. 1). Significant differences
were found in number of fruits throughout the
386 THE WILSON JOURNAL OF ORNITHOLOGY • Vol. 121, No. 2, June 2009
FIG. 1. Temporal patterns in availability of fruits for Military Macaws in two locations in Mexico with
different vegetation cover.
year at this site (X2r ⫽10.00, df ⫽ 1, P ⫽
0.02).
Military Macaw Diet.—We observed Mili-
tary Macaws feeding on 10 plant species from
eight families (Table 1). Seeds were the most
consumed item in 39.02% of foraging obser-
vations. Macaws were also observed taking
water among the leaves of Tillandsia grandis
and consuming latex from the stems of Plu-
meria rubra.
Macaws used some species more than pre-
dicted by plant abundance, as was the case for
Bursera schechtendalii (X2 ⫽ 98020.67; df ⫽
12; P ⫽ 0.0001) in the tropical dry forest of
Santa Marı́a Tecomavaca, and Celtis caudata
(X2 ⫽ 991.75; df ⫽ 4; P ⫽ 0.0001) in the
tropical dry forest of San Juan Coyula. The
rate of seed consumption was highest in com-
parison with other food items (Table 1; F5,63
⫽ 5.93; P ⫽ 0.0004). A one-way ANOVA
showed significant differences in the rate of
consumption among seeds from different
plants (F3,45 ⫽ 3.939; P ⫽ 0.015).
The Military Macaw had a standardized
Levins niche breadth index of B ⫽ 0.18. The
standardized Levins niche breadth index in the
breeding season was B ⫽ 0.033, while it was
higher (B ⫽ 0.158) in the non-breeding sea-
son.
Nutritional Value of Food Items.—Leaves
and flowers of Cyrtocarpa procera and Til-
landsia makoyana, and the latex of Plumeria
rubra had the highest water content. Fruits of
Neobuxbaumia tetetzo had the highest protein
content while fruit of Bunchosia montana was
highest in sugar content. Lipids occurred in
 Contreras-González et al. • DIET OF MILITARY MACAWS IN MÉXICO 387

TABLE 1. Diet of Military Macaws in the Cuicatlán Region, México. (a Site: SMT ⫽ Santa Marı́a Teco-
mavaca, SJC ⫽ San Juan Coyula; b RA: relative abundance in site; c Part eaten: rp ⫽ ripe, unrp ⫽ unripe, dr
⫽ dry, lf ⫽ leaf, fl ⫽ flower, lx ⫽ latex, ft ⫽ fruit, se ⫽ seed; d FO ⫽ number of foraging observations; e NM
⫽ number of macaws; f CR ⫽ consumption rate).
Family/species Sitea RA in sitesb Part eatenc FOd NMe CRf/min ⫾ SD (n) Month

Breeding season
Anacardiaceae SMT 0.036 rp lf 3 6 2.66 ⫾ 0.19 (3) Aug 2005
Cyrtocarpa procera rp fl 2 7 3.4 ⫾ 0.1 (5) Mar 2006
Apocyanaceae SMT 0.043 Stem lx 3 3 Aug 2005
Plumeria rubra
Bromeliaceae SMT Water 5 6 Jun 2005
Tillandsia grandis from lf
T. makoyana SJC rp lf 1 3 2 (1) Mar 2006
Burseraceae SMT 0.026 unrp se 1 3 4 (1) Mar 2005
Bursera aptera
B. schlechtendalii SMT 0.046 unrp se 2 2 3 (1) Mar 2005, Dec 2005
Cactaceae SMT 0.36 rp ft ⫹ se 1 3 4.4 (1) Jun 2005
Neobuxbaumia
tetetzo
Leguminosae SMT dr se 1 4 2.5 ⫾ 0.7 (2) Mar 2006
Lysiloma divaricata
Non-breeding season
Burseraceae SMT 0.046 unrp se 2 2 3.25 ⫾ 1.62 (2) Dec 2005
B. schlechtendalii
Malpighiaceae SJC 0.18 rp fr 5 31 2.9 ⫾ 0.11 (9) Jan 2006, Feb 2006
Bunchosia montana
Ulmaceae SJC 0.12 dr se 5 87 5.65 ⫾ 0.04 (39) Mar 2006
Celtis caudata

high concentrations (10%) in fruits of Neo-
buxbaumia tetetzo, leaves and flowers of Cyr-
tocarpa procera, seeds of Celtis caudata and
Bursera aptera, and latex of Plumeria rubra
(Table 2).
DISCUSSION
Resource availability changed seasonally in
our study site. Tropical dry forest in Santa
Marı́a Tecomavaca provided food resources
for Military Macaws during the prolonged dry
season and when breeding occurs from March
until September (Rivera-Ortiz et al. 2008).
Macaws may move during winter to other
sites with higher food resource availability
(Gaucı́n et al. 1999). Seasonal migrations are
common among macaws; other psittacines are
known to track food resource availability
(Pizo et al. 1995; Bonadie and Bacon 2000;
Renton 2001, 2002; Symes and Perrin 2003;
Karubian et al. 2005).
The diet of Military Macaws was highly
variable seasonally. The diet was mostly
seeds, as in other Psittacidae, but also con-
tained fruits of Bunchosia montana and Neo-
buxbaumia tetetzo, and leaves of Cyrtocarpa
procera. Military Macaws had a narrow diet
based on the Levins niche breadth index.
Sosa-Asanza (2000) reported values for the
Levins index for Maroon-tailed Parakeet (Pyr-
rhura melanura) of B ⫽ 0.5, Cobalt-winged
Parakeet (Brotogeris cyanoptera) of B ⫽ 0.55,
Black-headed Parrot (Pionites melanocephal-
us) of B ⫽ 0.75, Blue-headed Parrot (Pionus
menstruus) of B ⫽ 0.55, and Orange-winged
Amazon (Amazona amazona) of B ⫽ 0.55.
Renton (2001) reported the Lilac-crowned
Parrot (Amazona finschi) had an index of B ⫽
0.55 in the rainy season, while Scarlet Ma-
caws had an index of B ⫽ 0.39 (Renton 2006).
Other studies also indicate high diet speciali-
zation for macaws, even though niche breadth
was not calculated (Loza 1997, Iñigo-Elias
1999).
Niche breadth was low, but varied between
the two sites and between seasons, being
broader in Santa Marı́a Tecomavaca where
macaws breed. Macaws, during breeding,
388 THE WILSON JOURNAL OF ORNITHOLOGY • Vol. 121, No. 2, June 2009

spent more time in the canyon (Rivera-Ortiz

et al. 2008) consuming available food resourc-

63.4 2 ⫾ 1.65 (2)

67.95 ⫾ 0.29 (6)

29.72 ⫾ 1.78 (2)

31.61 ⫾ 1.93 (3)

47.64 ⫾ 1.17 (3)

5.27 ⫾ 0.08 (4)
es. These variations have been reported for

75.49 ⫾ 1.3 (6)

other parrots that had a correlation between
Water
availability of food resources and diet breadth

80.59 (1)

1.2 (1)
(Wermundsen 1997, Renton 2001) or seasonal
variation that could not be attributed to food
abundance (Matuzak et al. 2008). Macaws
consumed the latex of Plumeria rubra in our
study site when this plant was flowering. This
is the first record of a parrot consuming plant
18.38 ⫾ 0.16 (3)

41.13 ⫾ 0.83 (3)

9.66 ⫾ 0.56 (3)
25.54 ⫾ 0.36 (3)
7.39 ⫾ 0.35 (3)
36.12 ⫾ 0.14 (3)
9.63 ⫾ 0.13 (3)
latex. Latex had high water content, but also
16.4 ⫾ 0.56 (3)

15.58 ⫾ 0.1 (3)

was rich in lipids which constitute a good en-
Lipid

ergetic source for large birds such as macaws.

The use of leaves as a food resource is also
uncommon for parrots (Pizo et al. 1995, Gau-
Nutritional content of food items in the diet of Military Macaws. Sample sizes are in parentheses.

cı́n et al. 1999, Kristosch and Marcondes-Ma-

chado 2001, Matuzak et al. 2008).
Parrots are usually considered seed preda-
tors (Janzen 1972, 1981; Higgins 1979; Howe
1.07 (3)
0.09 (3)
0.15 (3)
0.62 (3)

0.94 (3)
0.11 (3)
0.89 (3)
0.07 (3)

1980; Galetti 1993; Kristosch and Marcondes-

0.1 (3)
Carbohydrate

Machado 2001; Renton 2001, 2006; Francisco

et al. 2002; Bollen and Elsacker 2004) with
⫾
⫾
⫾
⫾
⫾
⫾
⫾
⫾
⫾

potential to reduce seedling recruitment (Gal-

5.77
6.57
8.19
15.16
2.72
7.93
5.50
31.16
2.51

etti and Rodrigues 1992, Galetti 1993, Fran-

cisco et al. 2002). Renton (2001) proposed
that parrots can contribute to maintenance of
tree diversity in tropical forests. Military Ma-
caws in our study, consumed only the pulp of
Bunchosia montana dropping the seeds where
0.10 (3)
0.07 (3)

0.09 (3)
0.12 (3)
0.12 (3)
1.58 (3)
0.13 (3)
0.92 (3)
0.1 (3)

secondary dispersers could act. Macaws also

Protein

consumed fruits of Neobuxbaumia tetetzo, a

⫾
⫾
⫾
⫾
⫾
⫾
⫾
⫾
⫾

columnar cacti that has a high number of

6.31
9.04
0.78
1.44
1.05
6.21
28.63
3.47
7.42

small seeds, and probably serves as a primary

disperser as described for other parrots eating
fruits with small seeds (Eitniear et al. 1994,
Norconk et al. 1998). Further studies are need-
ed to evaluate the role of Military Macaws as
seed dispersers.
The nutritional content of food resources
Neobuxbaumia tetetzo (seeds and fruits)

eaten by Military Macaws is comparable to

that in foods used by other parrots (Abramson
1995, Pepper 1996, Pepper et al. 2000). The
lipid content of fruits was similar to that re-
Tillandsia makoyana (leaves)
Cyrtocarpa procera (leaves)

Bunchosia montana (fruits)

ported for fruit eaten by Lilac-crowned Parrots

B. schlechtendalii (seeds)

(Gutı́errez-Pérez 2005), and by parrots in Peru

Bursera aptera (seeds)
Plumeria rubra (latex)

Celtis caudata (seeds)

Species

C. procera (flowers)

(Gilardi 1996), except for fruits of Neobux-

baumia tetetzo which had high protein content
(28.6%). Military Macaws used food resourc-
TABLE 2.

es rich in protein and lipids during the breed-

ing season when they require energetically
rich foods (Vriends 1991); nestlings also have
high protein requirements for growth (Sakai
 Contreras-González et al. • DIET OF MILITARY MACAWS IN MÉXICO
et al. 1986, Vriends 1991, Koutsos et al.
2001).
Military Macaws had a narrow but variable
diet during the year. This variation and their
high mobility may represent adaptability to
changing food resources. However, the high
energetic content of their preferred foods sug-
gests that Military Macaws can be considered
as seasonally specialized birds which track
food resources in a landscape context. Con-
servation of Military Macaws depends on
conservation of food resources; management
plans for the region should focus on preserv-
ing plant species used throughout the year.
ACKNOWLEDGMENTS
We appreciate revision of an earlier draft of this pa-
per by Katherine Renton. Financial assistance was pro-
vided by projects PAPIIT-DGAPA IN 207305,
UNAM, and CONABIO DT006 to M. C. Arizmendi,
and by a fellowship to A. M. Contreras (CONACyT:
189305). We thank the local authorities of Santa Marı́a
Tecomavaca and San Juan Coyula, México for use of
facilities.
LITERATURE CITED
ABRAMSON, J. 1995. Nutritional requirements. Pages
111–146 in The large macaws: their care, breed-
ing and conservation (J. Abramson, B. L. Speer,
and J. B. Thomsen, Editors). Raintree Publica-
tions, Fort Bragg, California, USA.
BIRDLIFE INTERNATIONAL. 2003. BirdLife’s online
world bird data base: the site for bird conserva-
tion. Version 2.0. Cambridge, United Kingdom.
http://www.birdlife.org (accessed 28 January
2004).
BOLLEN, A. AND L. V. ELSACKER. 2004. The feeding
ecology of the Lesser Vasa Parrot, Coracopsis ni-
gra in south-eastern Madagascar. Ostrich 75:141–
146.
BONADIE, W. A. AND P. R. BACON. 2000. Year-round
utilization of fragmented palm swamp forest by
Red-bellied Macaw (Ara manilata) and Orange-
winged Parrots (Amazona amazonica) in the Na-
riva Swamp (Trinidad). Biological Conservation
95:1–5.
BORCHERT, R. 1994. Soil and stem water storage de-
termine phenology and distribution of tropical dry
forest trees. Ecology 75:1437–1449.
CARREÓN, G. G. 1997. Estimación poblacional, biolo-
gı́a reproductiva y ecologı́a de la nidificación de
la guacamaya verde (Ara militaris) en una selva
estacional del oeste del estado de Jalisco. Tesis de
licenciatura. Facultad de Ciencias, Universidad
Nacional Autónoma de México, México, D.F.
CHAPMAN, C. A., L. J. CHAPMAN, R. K. H. WANGHAM,
D. GEBO, AND L. GARDNER. 1992. Estimators of
389
abundance of tropical trees. Biotropica 24:527–
531.
CITES. 1998. Apendices I, II and III to the Convention
on International Trade in Endangered Species of
wild fauna and flora. USDI, Fish and Wildlife Ser-
vice, Washington, D.C., USA.
COLLAR, N. J. 1997. Order Psittaciformes family Psit-
tacidae (parrots). Pages 280–339 in Handbook of
the birds of the world. Volume 4 (J. del Hoyo, A.
Elliott, and J. Sargatal, Editors). Lynx Edicions,
Barcelona, Spain.
COLLAR, N. J. AND A. T. JUNIPER. 1992. Dimensions
and causes of the parrot conservation crisis. Pages
1–24 in New World parrots in crisis: solutions
from conservation biology (S. R. Bessinger and
N. F. R. Snyder, Editors). Smithsonian Institution
Press, Washington, D.C., USA and London, Unit-
ed Kingdom.
COLWELL, R. K. AND D. J. FUTUYMA. 1971. On the
measurement of niche breadth and ovelap. Ecol-
ogy 52:567–576.
EITNIEAR, J. C., S. MCGHEE, AND W. WADDELL. 1994.
Observations on the feeding upon Psittacanthus
calyculatus by Brown-hooded Parrots (Pionopsit-
ta haematotis). Ornitolologia Neotropical 5:119–
120.
EVERITT, B. S. 1977. The analysis of contingency ta-
bles. Chapman and Hall, London, United King-
dom.
FORSHAW, J. M. 1989. Parrots of the world. Third Edi-
tion. Lansdowne Press. Melbourne, Australia.
FRANCISCO, M. R., V. D. LUNARDI, AND M. GALETTI.
2002. Massive seed predation of Pseudobombax
grandiflorum (Bombacaceae) by parakeets Broto-
geris versicolurus (Psittacidae) in a forest frag-
ment in Brazil. Biotropica 34:613–615.
GALETTI, M. 1993. Diet of the Scaly-headed Parrot
(Pionus maximiliani) in a semideciduous forest in
southeastern Brazil. Biotropica 25:419–425.
GALETTI, M. AND M. RODRIGUES. 1992. Comparative
seed predation on pods by parrots in Brazil. Bio-
tropica 24:222–224.
GAUCÍN, R. N., E. E. IÑIGO, C. J. TREVIÑO, T. B.
ZÚÑIGA, H. G. SIFUENTES, G. L. CARMONA, AND R.
M. A. SÁNCHEZ. 1999. Biologı́a de conservación
de la Guacamaya Verde (Ara militaris) en el Só-
tano del Barro, Querétaro. Informe final del proy-
ecto. Facultad de Ciencias Naturales, Universidad
Autónoma de Querétaro, Querétaro, México.
GILARDI, J. D. 1996. Ecology of parrots in the Peruvian
Amazon: habitat use, nutrition and geophagy. Dis-
sertation. University of California, Davis, USA.
GONZÁLEZ, M. AND I. PEÑALOZA. 2000. Biomoléculas.
Métodos de Análisis. Universidad Nacional Au-
tónoma de México, Mexico, D.F.
GUTIÉRREZ-PÉREZ, A. 2005. Disponibilidad y calidad
nutritiva de recursos y su relación con la dieta de
las crı́as del Loro Corona Lila (Amazona finschi)
en un bosque tropical caducifolio. Thesis. Instituto
de Biologı́a, Universidad Nacional Autónoma de
México, Mexico, D.F.
390 THE WILSON JOURNAL OF ORNITHOLOGY • Vol. 121, No. 2, June 2009
HIGGINS, M. L. 1979. Intensity of seed predation on
Brosimum utile by Mealy Parrots (Amazona fari-
nosa). Biotropica 11:80.
HOWE, H. F. 1980. Monkey dispersal and waste of a
neotopical fruit. Ecology 61:944–959.
HOWELL, S. N. G. AND S. WEBB. 1995. A guide to the
birds of Mexico and northern Central America.
Oxford University Press, New York, USA.
ÍÑIGO-ELIAS, E. 1999. Las guacamayas verde y escar-
lata en México. Biodiversitas Año. 5:1–25.
IÑIGO-ELIAS, E. 2000. Estado de conservación de las
Guacamayas verde (Ara militaris) y escarlata (Ara
macao) en México. Audubon Latin Americana 3:
1–3.
IZAKI, I. 1993. Influence of no protein nitrogen on es-
timation of protein from total nitrogen in fleshy
fruit. Journal of Chemical Ecology 19:2605–2615.
JANZEN, D. H. 1972. Escape in space by Sterculia apa-
tela seeds from the bug Dysdercus fasciatus in a
Costa Rican deciduous forest. Ecology 53:350–
361.
JANZEN, D. H. 1981. Ficus ovalis seed predation by an
Orange-chinned Parakeet (Brotogeris jugularis) in
Costa Rica. Auk 98:841–844.
KARUBIAN, J., J. FABARRA, D. YUNES, J. P. JORGENSON,
D. ROMO, AND T. B. SMITH. 2005. Seasonal and
spatial variation in macaw abundance in the Ec-
uadorian Amazon. Condor 107:617–626.
KOUTSOS, E. A., K. D. MATSON, AND K. C. KLASING.
2001. Nutrition of the birds in the Order Psitta-
ciformes: a review. Journal of Avian Medicine
and Surgery 15:257–275.
KREBS, C. J. 1989. Ecological methodogy. Harper Col-
lins Publisher, New York, USA.
KRISTOSCH, G. C. AND L. O. MARCONDES-MACHADO.
2001. Diet and feeding behavior of the Reddish-
bellied Parakeet (Pyrrhura frontalis) in an Arau-
caria forest in southeastern Brazil. Ornitologia
Neotropical 12:215–223.
LEVEY, D. J., H. A. BISSELL, AND S. F. O’KEEFE. 2000.
Conversion of nitrogen to protein and amino acids
in wild fruits. Journal of Chemical Ecology 26:
1749–1763.
LOZA, S. C. A. 1997. Patrones de abundancia, uso de
hábitat y alimentación de la guacamaya verde
(Ara militaris) en la Presa Cajón de Peña, Jalisco,
México. Thesis. Facultad de Ciencias, Universi-
dad Nacional Autónoma de México, México, D.F.
MARINEROS, L. AND C. VAUGHAN. 1995. Scarlet Ma-
caws of Carara: perspectives for management.
Pages 445–467 in The large macaws: their care,
breeding and conservation (J. Abramson, B. L.
Speer and J. B. Thomsen, Editors). Raintree Pub-
lications, Fort Bragg, California, USA.
MATUZAK, G. D., M. B. BEZY, AND D. BRIGHTSMITH.
2008. Foraging ecology of parrots in a modified
landscape: seasonal trends and introduced species.
Wilson Journal of Ornithology 120:353–365.
NORCONK, M. A., B. W. GRAFTON, AND N. L. CONKLIN-
BRITTAIN. 1998. Seed dispersal by neotropical
seed predators. American Journal of Primatology
45:103–126.
OSBORNE, D. R. 1986. Análisis de los nutrientes de los
alimentos. Editoria Acribia, S. A. España.
PEPPER, J. W. 1996. The behavior and ecology of the
Glossy Black-cockatoo Calyptorhynchus lathami
halmaturinus. Dissertation. University of Michi-
gan, Ann Arbor, USA.
PEPPER, J. W., T. D. MALE, AND G. E. ROBERTS. 2000.
Foraging ecology of the South Australian Glossy
Black-cockatoo (Calyptorhynchus lathami hal-
maturinus). Australian Ecology 25:6–24.
PETERSON, R. T. AND E. L. CHALIFF. 1989. Guı́a de
Aves de México. Editorial Diana, México, D. F.
PIZO, M. A., I. SIMAO, AND M. GALETTI. 1995. Diet and
flock size of sympatric parrots in the Atlantic For-
est of Brazil. Ornitologia Neotropical 6:87–95.
RENTON, K. 2001. Lilac-crowned Parrot diet and food
resource availability: resource tracking by a parrot
seed predator. Condor 103:62–69.
RENTON, K. 2002. Seasonal variation in occurrence of
macaws along a rainforest river. Journal of Field
Ornithology 73:15–19.
RENTON, K. 2006. Diet of adult and nestling Scarlet
Macaws in southwest Belize, Central America.
Biotropica 38:280–283.
RIVERA-ORTÍZ, F. A., A. M. CONTRERAS-GONZÁLEZ, C.
A. SOBERANES-GONZÁLEZ, A. VALIENTE-BANUET,
AND M. DEL C. ARIZMENDI. 2008. Seasonal abun-
dance and breeding chronology of the Military
Macaw (Ara militaris) in a semi-arid region of
central Mexico. Ornitologia Neotropical 19:255–
263.
SAKAI, H. F., C. J. RALPH, AND C. D. JENKINS. 1986.
Foraging ecology of the Hawaiian Crow, an en-
dangered generalist. Condor 88:211–219.
SALAZAR, T. J. M. 2001. Registro de la guacamaya ver-
de (Ara militaris) en los cañones del Rı́o Sabino
y Rı́o Seco, Santa Marı́a Tecomavaca, Oaxaca,
México. Huitzil 2:18–20.
SAUNDERS, D. A. 1977. The effect of agricultural clear-
ing on the breeding success of the White-tailed
Black Cockatoo. Emu 77:180–184.
SEMARNAT. 2001. Norma Oficial Mexicana NOM-
059-ECOL-2001, Protección ambiental-Especies
nativas de México de flora y fauna silvestres-Ca-
tegorı́as de riesgo y especificaciones para su in-
clusión, exclusión o cambio-Lista de especies en
riesgo. Diario Oficial de la Federación. Miércoles
6 de marzo del 2002. [En lı́nea] http://
www.ine.gob.mx/ueajei/norma59a.html
SOKAL, R. R. AND F. J. ROHLF. 1979. Biometry. H.
Blume Editores. Madrid, España.
SOSA-ASANZA, A. C. 2000. Food choice, seasonal,
movements, and population trends of parrots in
the Ecuadorian Amazon. Dissertation. University
of Georgia, Athens, USA.
SYMES, C. T. AND M. R. PERRIN. 2003. Seasonal oc-
currence and local movements of the Grey-headed
(Brown-necked) Parrot Poicephalus fuscicollis
 Contreras-González et al. • DIET OF MILITARY MACAWS IN MÉXICO
auahelicus in South Africa. African Journal of
Ecology 41:299–305.
VALIENTE-BANUET, A., A. CASAS, A. ALCÁNTARA, P.
DÁVILA, H. N. FLORES, M. DEL C. ARIZMENDI, J.
L. VILLASEÑOR, AND R. J. ORTEGA. 2000. La ve-
getación del Valle de Tehuacan-Cuicatlán. Bol-
etı́n de la Sociedad Botánica de México 67:24–
74.
View publication stats
391
VRIENDS, M. 1991. Feeding parrots the right way. AFA
Watchbird 18:47–51.
WERMUNDSEN, T. 1997. Seasonal change in the diet of
the Pacific Parakeet Aratinga strenua in Nicara-
gua. Ibis 139:566–568.
ZAR, H. J. 1999. Biostatistical analysis. Fourth Edition.
Prentice-Hall Inc., Englewood Cliffs, New Jersey,
USA.