Ara glaucogularisblue-throated macaw

Ge­o­graphic Range

Blue-throated macaws (Ara glaucogu­laris) are found only in the Beni de­part­ment of Bo­livia (200 to 300 m above sea-level). In total, blue-throated macaws in­habit an area of 2508 square kilo­me­tres. (Hesse and Duffield, 2000; "Blue-throated Macaw", 2009)

There are two areas in­hab­ited by two sub-pop­u­la­tions of Ara glaucogu­laris: one is to the north­west of Trinidad (the cap­i­tal city of Beni), and the other is to the south of Trinidad. This sep­a­ra­tion may have oc­curred be­cause of the in­dige­nous peo­ples that his­tor­i­cally in­hab­ited this area and hunted blue-throated macaws to use the feath­ers in or­na­men­tal cos­tume. This sep­a­ra­tion could also have been caused more re­cently by the wild-bird trade. With the high pop­u­la­tion of hu­mans, any blue-throated macaws in the vicin­ity would have a higher chance of being caught. The for­ma­tion of large human set­tle­ments in this area also re­sulted in a loss of suit­able habi­tat and habi­tat frag­men­ta­tion for this species. Hence, there are no blue-throated macaws in the vicin­ity of Trinidad. (Hesse and Duffield, 2000; Strem, 2008)

Habi­tat

Blue-throated macaws in­habit gallery forests and is­lands of trees sur­rounded by a trop­i­cal sa­vanna. Sea­sonal rains cause flood­ing from Oc­to­ber to May and trans­form the sa­vanna into a grassy swamp sur­round­ing per­ma­nently dry el­e­vated for­est is­lands. The pres­ence of Motacú palms (At­talea phaler­ata) is re­quired for the sur­vival of Ara glaucogu­laris as this species of macaw feeds and nests in these palms more than any other species of plant. They occur most often be­tween the el­e­va­tions of 200 and 300 m. (Ben­stead, et al., 2009; Hesse and Duffield, 2000; Jor­dan and Munn, 1993)

Most of blue-throated macaws' habi­tat is used for cat­tle ranch­ing. How­ever, the land is un­suit­able for cul­ti­va­tion, so habi­tat al­ter­ation for agri­cul­tural use does not occur. Al­though cat­tle will tram­ple saplings, ma­ture Motacú palms are very hardy and re­sist dam­age. This palm is some­what fire re­sis­tant as well. As a re­sult, Motacú palms often dom­i­nate the for­est frag­ments in the Beni Sa­vanna. (Hesse and Duffield, 2000)

  • Range elevation
    200 to 300 m
    656.17 to 984.25 ft

Phys­i­cal De­scrip­tion

Blue-throated macaws have very vivid col­oration. They have bright turquoise-blue feath­ers cov­er­ing their throat, crown, back and the dor­sal side of their wings and tail. Golden-yel­low feath­ers grow in a stripe be­tween the blue crown and throat on the side of the face and on the ven­tral side of their body, wings and tail. On the face there is a sparsely feath­ered patch of skin near the base of the large dark-col­ored bill that has 5 or 6 hor­i­zon­tal stripes of blue feath­ers which are unique for every blue-throated macaw and can be used to in­di­vid­u­ally iden­tify adults. The skin on this patch is pre­dom­i­nantly white with a pink tint close to the bill. (Ben­stead, et al., 2009; Kyle, 2007a)

Adult blue-throated macaws' mass ranges from 600 to 1000 g with a length of about 85 cm (mea­sured from the top of the head to tip of the long, ta­pered tail) and a wingspan of ap­prox­i­mately three feet or 0.9 m. Ara glaucogu­laris shows lit­tle eas­ily ob­serv­able sex­ual di­mor­phism; how­ever, males tend to be a lit­tle big­ger than fe­males with ap­prox­i­mate masses of 600 g and 800 g re­spec­tively. (Ben­stead, et al., 2009; Clubb, 1994; Clubb, 2009; Kyle, 2007a)

Newly hatched blue-throated macaws are com­pletely pink and have no feath­ers. Gray down grows in as they age, and is later re­placed by col­ored, fully-de­vel­oped feath­ers. The iris also changes color with age. The eye color of a nestling is ini­tially black and changes to brown soon after the eyes open. When the macaaw is one to three years old, its eyes will turn grey, then white. As the macaw ma­tures, the iris turns yel­low and will be more golden at 10 years and will be­come a richer gold with age. El­derly macaws show a ring of dark grey sur­round­ing the pupil where the iris has be­come thin­ner and the back of the retina shows through. This con­tin­uum of the iris’ colour can be used to es­ti­mate the age of a macaw. (Clubb, 1994; Clubb, 2009; Kyle, 2007b)

Blue-throated macaws look very sim­i­lar to the more com­mon blue and gold macaws (Ara ara­rauna). How­ever, they can be dis­tin­guished most clearly by the col­ors of the feath­ers on the throat and crown. Blue-throated macaws have 5 or 6 hor­i­zon­tal lines of feath­ers across the oth­er­wise bare fa­cial patch while blue and gold macaws have 3. Blue-throated macaws are smaller and also have a more nasal, higher-pitched and softer voice in com­par­i­son to blue and gold macaws. (Ben­stead, et al., 2009; Kyle, 2007a; Riv­iere, et al., 1986; "Blue and Gold Macaw", 2009)

Poly­mor­phisms, sea­sonal and ge­o­graph­i­cal vari­a­tions have not been ob­served in blue-throated macaws.

  • Sexual Dimorphism
  • sexes alike
  • male larger
  • Range mass
    600 to 1000 g
    21.15 to 35.24 oz
  • Average length
    85 cm
    33.46 in
  • Average wingspan
    0.9 m
    2.95 ft

Re­pro­duc­tion

Blue-throated macaws are monog­a­mous and mate for life. It is not known if these macaws will pair with an­other mate if their orig­i­nal mate dies. Noth­ing else is known about mat­ing sys­tems for Ara glaucogu­laris in the wild. (Kyle, 2007b)

Blue-throated macaws breed once a year if the en­vi­ron­men­tal con­di­tions per­mit them; how­ever, if the eggs or nestlings are lost, the breed­ing pair may pro­duce a sec­ond clutch in the same breed­ing sea­son. It has been spec­u­lated that the two sub-pop­u­la­tions breed at slightly dif­fer­ent times: the north­ern pop­u­la­tion breed from Au­gust to No­vem­ber and the south­ern pop­u­la­tion breed from No­vem­ber to March. The fe­male will lay one to three eggs per clutch and in­cu­bates for 26 days. Blue-throated macaws have a mass of ap­prox­i­mately 18 g at hatch­ing. The nestlings fledge at 13 to 14 weeks. Young blue-throated macaws will not be fully in­de­pen­dent of their par­ents for a full year. Blue-throated macaws will be sex­u­ally ma­ture at about 5 years. (Ben­stead, et al., 2009; Clubb, 1994; Kyle, 2007b; Kyle, 2007a; Strem, 2008; "Blue-throated Macaw", 2009)

Ara glaucogu­laris usu­ally nests in cav­i­ties of palm trees, most often At­talea phaler­ata, al­though it will nest in other palm species as well. Dead palms are ideal to nest in as they are hol­lowed out by large grubs after the tree has died. Some macaw species, in­clud­ing Ara glaucogu­laris, will eat the palm fronds to an ex­tent that will kill the tree. The trunk of the palm is hol­lowed out by grubs re­sult­ing in the cre­ation of a po­ten­tial nest. It is not yet known whether this is co­in­ci­dence or if these birds do this in­ten­tion­ally to cre­ate nest sites. (Gi­lardi, et al., 2005; Hesse and Duffield, 2000; Jor­dan and Munn, 1993; Kyle, 2007b; Strem, 2008)

Nest­ing pairs of Ara glaucogu­laris don’t con­sis­tently stay at one nest for con­sec­u­tive breed­ing sea­sons and will usu­ally search for dif­fer­ent nest­ing sites every year. (Kyle, 2007b)

  • Breeding interval
    Blue-throated macaws breed once or twice yearly.
  • Breeding season
    Blue-throated macaws breed from November to March.
  • Range eggs per season
    1 to 3
  • Average time to hatching
    26 days
  • Range fledging age
    13 to 14 weeks
  • Average time to independence
    1 years
  • Average age at sexual or reproductive maturity (female)
    5 years
  • Average age at sexual or reproductive maturity (male)
    5 years

A mat­ing pair of Blue-throated macaws must find a suit­able nest be­fore fer­til­iza­tion oc­curs. If a nest site can­not be found, the pair may ex­ca­vate their own nest. While de­cid­u­ous trees can be used, dead palms are eas­ier to hol­low out and are used quite often. Prospec­tive par­ents must also with­stand com­pe­ti­tion for nest sites from other macaw species such as Ara ara­rauna, Ara chloroptera and Ara macao as well as toco tou­cans, large wood­peck­ers, barn owls, bats, and bees. (Ben­stead, et al., 2009; Kyle, 2007b; Kyle, 2007a)

Fe­male blue-throated macaws lay the eggs and in­cu­bate them until they hatch. Males feed the fe­males dur­ing in­cu­ba­tion. Both par­ents feed the nestlings once the chicks have hatched and the pre­co­cial chicks must be kept warm by their par­ents be­fore their feath­ers grow in. After the nestlings’ feath­ers grow in they are often left alone while both par­ents gather food. The young macaws are still de­pen­dent upon their par­ents for food after they fledge until they are fully weaned and ca­pa­ble of for­ag­ing by them­selves. Even after the ju­ve­nile macaws are able to pro­vide for them­selves, it has been ob­served that young blue-throated macaws will stay with their par­ents up to a year. Dur­ing this time, the par­ents will skip an en­tire breed­ing sea­son. (Kyle, 2007b; Kyle, 2007a)

  • Parental Investment
  • altricial
  • male parental care
  • female parental care
  • pre-fertilization
    • provisioning
    • protecting
      • female
  • pre-hatching/birth
    • provisioning
      • male
    • protecting
      • male
      • female
  • pre-weaning/fledging
    • provisioning
      • male
      • female
    • protecting
      • male
      • female
  • pre-independence
    • provisioning
      • male
      • female
    • protecting
      • male
      • female
  • post-independence association with parents
  • extended period of juvenile learning

Lifes­pan/Longevity

In cap­tiv­ity, blue-throated macaws are able to breed up to about 30 to 35 years, after which aging and age re­lated dis­eases begin to show. Very few macaws in cap­tiv­ity live to 50 years of age. Blue-throated macaws have not been stud­ied very long in the wild, and in­for­ma­tion on lifes­pan is yet to be dis­cov­ered. (Clubb and Karpin­ski, 1992; Clubb, 1994)

  • Range lifespan
    Status: captivity
    50 (high) years

Be­hav­ior

Blue-throated macaws are so­cial birds as they form monog­a­mous mat­ing pairs and will also live in small groups. One group of 70 in­di­vid­u­als was known at one point, but this species is not often seen in large flocks as other species of macaws have. This is most likely be­cause blue-throated macaws are very rare and have such a low pop­u­la­tion that the for­ma­tion of such large groups is very un­likely. A mat­ing pair of blue-throated macaws can be very af­fec­tion­ate to­wards one an­other and are often seen preen­ing each other's feath­ers and perch­ing close to­gether. Blue-throated macaws are some­times seen in­ter­act­ing with blue and gold macaws (Ara ara­rauna) as well. (Ben­stead, et al., 2009; Kyle, 2007b; Ben­stead, et al., 1992)

Blue-throated macaws live, feed and nest in trees. Their main mode of lo­co­mo­tion is fly­ing, but they are also able to climb trees, ma­neu­ver along branches and walk on the ground. These birds are ac­tive dur­ing the day and usu­ally stay in one gen­eral area. There have been sight­ings of blue-throated macaws in Paraguay, which in­di­cates that this species will some­times travel long dis­tances. (Kyle, 2007b; Ben­stead, et al., 1992)

Home Range

There is not very much in­for­ma­tion avail­able per­tain­ing to the home ranges of in­di­vid­ual blue-throated macaws or pairs.

Com­mu­ni­ca­tion and Per­cep­tion

Blue-throated macaws com­mu­ni­cate mostly by sound. When they sus­pect dan­ger, they emit a very loud alarm­ing call and promptly fly off. Blue-throated macaws are known to com­mu­ni­cate with each other with quiet caws as well. Toa Kyle (2007b) de­scribes his ob­ser­va­tions of the “al­most fledg­ing” of a blue-throated macaw chick dur­ing which the chick re­ceived “light caws of en­cour­age­ment from its par­ents perched nearby.” Bird trap­pers have been known to use “caller” bait birds to at­tract blue-throated macaws of the op­po­site sex, so acoustic com­mu­ni­ca­tion is known to have a role for at­tract­ing mates. Tac­tile com­mu­ni­ca­tion is used as well. Mates show af­fec­tion for each other and their chicks quite often in the form of preen­ing. Like all birds, blue-throated macaws per­ceive their en­vi­ron­ment through audio, vi­sual, tac­tile, and chem­i­cal stim­uli. (Kyle, 2007b; Kyle, 2007a; Ben­stead, et al., 1992)

Food Habits

Blue-throated macaws do not eat seeds and nuts to the same ex­tent as many other macaw species do. In­stead, they eat pri­mar­ily fruit from large palms. The palm species At­talea phaler­ata is the most pre­dom­i­nant source, but they will also eat from Acro­co­mia ac­uleata and Mau­ri­tia fleux­osa. The macaws eat the meso­carp from ripe and nearly ripe fruit and have also been ob­served drink­ing the liq­uid from very im­ma­ture fruit. (Ben­stead, et al., 2009; Jor­dan and Munn, 1993; Kyle, 2007b; Strem, 2008)

  • Plant Foods
  • leaves
  • seeds, grains, and nuts
  • fruit

Pre­da­tion

Blue-throated macaws’ cau­tious­ness is one of their be­hav­ioral adap­ta­tions to pre­vent nest pre­da­tion. Par­ents will often perch just out­side their nest and ob­serve their sur­round­ings for preda­tors and other dan­gers. When re­turn­ing from for­ag­ing, blue-throated macaws will ap­proach the nest tree grad­u­ally only after ob­serv­ing the area first. While one par­ent en­ters the nest to feed the chicks, the other par­ent has been ob­served to stand guard near the nest tree. (Kyle, 2007a; Ped­er­son and Ped­er­son, 2002)

Few preda­tors of Ara glaucogu­laris exist. Adults can be preyed upon by great horned owls, south­ern caracaras, coati, tayras, and brown ca­puchin mon­keys. Nestlings are preyed upon by crane hawks, while both eggs and nestlings are eaten by toco tou­cans. (Gi­lardi, et al., 2005; Kyle, 2007b)

Ecosys­tem Roles

Blue-throated macaws play a role as prey in their ecosys­tem. They are also sub­ject to par­a­sites such as mites and bot­fly lar­vae. As fru­gi­vores, they are likely an im­por­tant seed dis­perser for na­tive, fruit­ing plants. (Kyle, 2007b)

  • Ecosystem Impact
  • disperses seeds

Eco­nomic Im­por­tance for Hu­mans: Pos­i­tive

Blue-throated macaws are used as pets. Their feath­ers have also been used for dec­o­ra­tion of or­na­men­tal cos­tume for in­dige­nous groups. (Hesse and Duffield, 2000; Strem, 2008)

  • Positive Impacts
  • pet trade
  • body parts are source of valuable material

Eco­nomic Im­por­tance for Hu­mans: Neg­a­tive

There are no known ad­verse ef­fects of Ara glaucogu­laris on hu­mans.

Con­ser­va­tion Sta­tus

Ara glaucogu­laris is cur­rently rated as crit­i­cally en­dan­gered on the IUCN Red List and is listed in Ap­pen­dix I by CITES. Trap­ping is il­le­gal as blue-throated macaws have been pro­tected by the na­tional leg­is­la­tion of Bo­livia and the Con­ven­tion on In­ter­na­tional Trade in En­dan­gered Species of Wild Fauna and Flora (CITES) since 1986. (Strem, 2008; "Blue-throated Macaw", 2009)

Trap­ping for the pet trade is the main rea­son that blue-fronted macaws are so crit­i­cally en­dan­gered. The rar­ity of this species drove the sell­ing price up re­sult­ing in in­creased pres­sure from trap­ping. As more birds were caught, blue-throated macaws be­came rarer. This be­came a vi­cious cycle that greatly re­duced the wild pop­u­la­tion of blue-throated macaws to the num­bers presently ob­served. There are cur­rently an es­ti­mated 50 to 250 in­di­vid­u­als in the wild. (Hesse and Duffield, 2000; Jor­dan and Munn, 1993; "Blue-throated Macaw", 2009)

Since spix's macaws (Cyanop­sitta spixii) went ex­tinct in 2000, Ara glaucogu­laris is now the most rare species of macaw in the world. With a wild pop­u­la­tion es­ti­mated to be be­tween 50 and 250 in­di­vid­u­als, ex­treme con­ser­va­tion ac­tions are nec­es­sary. The World Par­rot Trust has many vol­un­teers and em­ploy­ees work­ing to­wards the con­ser­va­tion of blue-throated macaws. These peo­ple mon­i­tor the nests to pro­tect the chicks from pre­da­tion. Chicks are also ex­am­ined pe­ri­od­i­cally to en­sure that they are healthy and re­ceiv­ing ad­e­quate food from their par­ents. If the chick is not doing as well as is ex­pected, then it is sup­ple­mented with for­mula. New nest boxes have been built and cur­rent nest sites im­proved. Sup­port from the local landown­ers has also been es­tab­lished. (Gi­lardi, et al., 2005; Kyle, 2006; Kyle, 2007b; Kyle, 2007a)

Con­trib­u­tors

Shelby Wyatt (au­thor), Uni­ver­sity of Al­berta, Au­gus­tana Cam­pus, Doris Audet (ed­i­tor), Uni­ver­sity of Al­berta, Au­gus­tana Cam­pus, Rachelle Ster­ling (ed­i­tor), Spe­cial Pro­jects.

Glossary

Neotropical

living in the southern part of the New World. In other words, Central and South America.

World Map

acoustic

uses sound to communicate

agricultural

living in landscapes dominated by human agriculture.

altricial

young are born in a relatively underdeveloped state; they are unable to feed or care for themselves or locomote independently for a period of time after birth/hatching. In birds, naked and helpless after hatching.

arboreal

Referring to an animal that lives in trees; tree-climbing.

bilateral symmetry

having body symmetry such that the animal can be divided in one plane into two mirror-image halves. Animals with bilateral symmetry have dorsal and ventral sides, as well as anterior and posterior ends. Synapomorphy of the Bilateria.

chemical

uses smells or other chemicals to communicate

diurnal
  1. active during the day, 2. lasting for one day.
endothermic

animals that use metabolically generated heat to regulate body temperature independently of ambient temperature. Endothermy is a synapomorphy of the Mammalia, although it may have arisen in a (now extinct) synapsid ancestor; the fossil record does not distinguish these possibilities. Convergent in birds.

female parental care

parental care is carried out by females

frugivore

an animal that mainly eats fruit

herbivore

An animal that eats mainly plants or parts of plants.

iteroparous

offspring are produced in more than one group (litters, clutches, etc.) and across multiple seasons (or other periods hospitable to reproduction). Iteroparous animals must, by definition, survive over multiple seasons (or periodic condition changes).

male parental care

parental care is carried out by males

monogamous

Having one mate at a time.

motile

having the capacity to move from one place to another.

native range

the area in which the animal is naturally found, the region in which it is endemic.

oviparous

reproduction in which eggs are released by the female; development of offspring occurs outside the mother's body.

pet trade

the business of buying and selling animals for people to keep in their homes as pets.

seasonal breeding

breeding is confined to a particular season

sedentary

remains in the same area

sexual

reproduction that includes combining the genetic contribution of two individuals, a male and a female

social

associates with others of its species; forms social groups.

swamp

a wetland area that may be permanently or intermittently covered in water, often dominated by woody vegetation.

tactile

uses touch to communicate

terrestrial

Living on the ground.

tropical

the region of the earth that surrounds the equator, from 23.5 degrees north to 23.5 degrees south.

tropical savanna and grassland

A terrestrial biome. Savannas are grasslands with scattered individual trees that do not form a closed canopy. Extensive savannas are found in parts of subtropical and tropical Africa and South America, and in Australia.

savanna

A grassland with scattered trees or scattered clumps of trees, a type of community intermediate between grassland and forest. See also Tropical savanna and grassland biome.

temperate grassland

A terrestrial biome found in temperate latitudes (>23.5° N or S latitude). Vegetation is made up mostly of grasses, the height and species diversity of which depend largely on the amount of moisture available. Fire and grazing are important in the long-term maintenance of grasslands.

visual

uses sight to communicate

Ref­er­ences

In­ter­na­tional Union for Con­ser­va­tion of Na­ture and Nat­ural Re­sources. 2009. "Ara glaucogu­laris" (On-line). The IUCN Red List of Threat­ened Species. Ac­cessed Sep­tem­ber 13, 2009 at http://​www.​iucnredlist.​org/​details/​142580/​0.

World Par­rot Trust. 2009. "Blue and Gold Macaw" (On-line). World Par­rot Trust. Ac­cessed No­vem­ber 18, 2009 at http://​www.​parrots.​org/​index.​php/​encyclopedia/​profile/​blue_​and_​gold_​macaw/​.

World Par­rot Trust. 2009. "Blue-throated Macaw" (On-line). World Par­rot Trust. Ac­cessed No­vem­ber 10, 2009 at http://​www.​parrots.​org/​index.​php/​encyclopedia/​profile/​blue_​throated_​macaw/​.

Wild­screen. 2009. "Blue-throated macaw" (On-line). ARKive. Ac­cessed Sep­tem­ber 13, 2009 at http://​www.​arkive.​org/​blue-throated-macaw/​ara-glaucogularis/​.

Ben­stead, P., J. Bird, S. Butchart, D. Cap­per, T. Stu­art, A. Symes. 1992. "BLUE-THROATED MACAW Ara glaucogu­laris" (On-line pdf). Ac­cessed Oc­to­ber 10, 2009 at http://​www.​birdlife.​info/​docs/​AmRDBPDFs/​Ara_​glaucogularis_​eng.​pdf.

Ben­stead, P., J. Bird, S. Butchart, D. Cap­per, T. Stu­art, A. Symes. 2009. "Species fact­sheet: Ara glaucogu­laris" (On-line). Ac­cessed Sep­tem­ber 13, 2009 at http://​www.​birdlife.​org/​datazone/​species/​index.​html?​action=SpcHTMDetails.​asp&​sid=1548#​FurtherInfo.

Clubb, S. 1994. "Life His­tory And Med­ical Man­age­ment of Macaws" (On-line pdf). Ac­cessed Oc­to­ber 09, 2009 at http://​www.​susanclubb.​com/​research.​html.

Clubb, S., L. Karpin­ski. 1992. "Aging in Macaws" (On-line pdf). Ac­cessed Oc­to­ber 09, 2009 at http://​www.​susanclubb.​com/​research.​html.

Clubb, S. 2009. "Blue-Throated Macaw" (On-line). SusanClubb.​com. Ac­cessed No­vem­ber 18, 2009 at http://​www.​susanclubb.​com/​education.​html.

Gi­lardi, J., T. Kyle, J. Eck­les. 2005. Bo­li­vian "Blue­beards". Psit­taScene, Vol. 17/No. 1: 2-10.

Hesse, A., G. Duffield. 2000. The sta­tus and con­ser­va­tion of the Blue-Throated Macaw Ara glaucogu­laris. Bird Con­ser­va­tion In­ter­na­tional, vol 10/iss 3: 255-275.

Jor­dan, O., C. Munn. 1993. First ob­ser­va­tions of the Blue-throated Macaw in Bo­livia. The Wil­son Bul­letin, vol 105/iss 4: 694-695.

Kyle, T. 2007. "Par­rot Blog­ger - Toa Kyle" (On-line). World Par­rot Trust. Ac­cessed No­vem­ber 10, 2009 at http://​www.​parrots.​org/​index.​php/​blog_​kyle/​P0/​.

Kyle, T. 2007. Rais­ing the Bar. Psit­taScene, Vol 19 / No 2: 14-17.

Kyle, T. 2006. Sav­ing Bo­livia's Blue-throated Macaw. Psit­taScene, vol 18 / no 1: 4-8.

Ped­er­son, B., B. Ped­er­son. 2002. Ob­ser­va­tions at an ac­tive nest of Blue-throated Macaws in the Beni De­part­ment of Bo­livia. Psit­taScene, Vol 14 / No 50: 2-3.

Riv­iere, S., S. Clubb, K. Clubb. 1986. The Elu­sive Caninde. AFA Watch­bird, vol 13: 6-9.

Strem, R. 2008. "Pop­u­la­tion Vi­a­bil­ity Analy­sis of the Blue-throated Macaw (Ara glaucogu­laris)" (On-line pdf). Ac­cessed Oc­to­ber 10, 2009 at http://​etd.​ohiolink.​edu/​send-pdf.​cgi/​Strem%20Cuellar%20Rosa%20Ines.​pdf?​acc_​num=bgsu1219175814.