Society for Range Management

Diets of Plains Vizcacha, Greater Rhea and Cattle in Argentina

Author(s): Javier A. Pereira, Rubn D. Quintana and Susana Monge
Source: Journal of Range Management, Vol. 56, No. 1 (Jan., 2003), pp. 13-20
Published by: Allen Press and Society for Range Management
Stable URL: http://www.jstor.org/stable/4003875 .
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J. Range Manage.
56: 13-20 January2003

Diets

of

plains

vizcacha,

greater

rhea

and

cattle

in

Argentina

JAVIER A. PEREIRA, RUBEN D. QUINTANA, AND SUSANA MONGE

Authors are Dissertation Student and Researcher and Laboratory Chief, Laboratorio de Ecologia Ambiental y Regional, Departamento de Ciencias
Biol6gicas, FCEyN, Universidad de Buenos Aires. Ciudad Universitaria, Pabell6n II, 1428 Buenos Aires, Argentina; and CONICETTechnician, Instituto
Argentino de Investigaciones de Zonas Aridas (IADIZA),CRICyT,CONICET.Calle Bajada del Cerro S/N, 5500 Mendoza, Argentina. At the time of the
research, the senior author was Thallmann Visiting Scholar (UBA) and Research Associate, Dept. of Organismic and Evolutionary Biology, Harvard
University, Cambridge,Mass. USA.

Abstract

Resumen

Food habits of plains vizcacha(Lagostomusmaximus),greater

Se estudiaron los haibitos alimenticios de la vizcacha

(Lagostomus maximus), el iiandu (Rhea americana) y el ganado
River Delta, Argentina,were studied over 2 years using micro- vacuno (Bos taurus) durante 2 auios en el Delta del Rio Parana,
histologicalanalysisof faeces.This was the first study of feeding Argentina, utilizando analisis microhistologico de heces. El prehabits of these herbivoresgrazing in common in a wetland of sente es el primer estudio que focaliza en la comparacion de los
Argentina.Poaceaewas the main diet componentthroughoutthe haibitos alimenticios de estos herbivoros en coexistencia en un
fueron el componente principal en
year for all 3 herbivores,with the exceptionof spring and sum- area de humedal. Las poa&ceas
mer, when greater rhea consumed a higher proportion of la dieta de los 3 herbivoros a lo largo del aiio, excepto en primavProsopisnigra (Griseb.)pods. Botanicalcompositionof plains era y verano, cuando los iiandutesconsumieron una mayor provizcachaand cattle diets was generallysimilarfor the same sea- porci6n de vainas de Prosopis nigra (Griseb.). La composici6n
son but different from that of greater rhea. Panicummilioides botanica de la dieta de la vizcacha y el ganado fue en general simNees., Dichondra microcalyx (Hallier) Fabris. and P. nigra were ilar para una misma estaci6n pero diferente de la del fiandu.
the most consumedspeciesfor vizcacha,whileP. nigra,Plantago Panicum milioides Nees., Dichondra microcalyx (Hallier) Fabris. y
myosuros Lam., Solanum sp. L., Spilanthes stolonifera (H. et A.) P. nigra fueron las especies mas consumidas por la vizcacha,
Baker and D. microcalyxdominatedthe greater rhea diet. The mientras que P. nigra, Plantago myosuros Lam., Solanum sp. L.,
species most consumedby cattle were LuziolaperuvianaGmel. Spilanthes stolonifera (H. et A.) Baker y D. microcalyx dominaron
and P. milioides.Similaritiesbetweenthe diets of plainsvizcacha en la dieta del fiandu. Las especies mas consumidas por el ganado
and cattle seem to support the ranchers' view that vizcachas fueron Luziola peruviana Gmel. y P. milioides. La similitud entre
compete with domestic herbivores for forage. However, high las dietas de la vizcacha y el ganado parecen apoyar la idea de los
overlapin food habits wouldresult in competitiononly if forage ganaderos sobre la competencia por el forraje entre estos heris scarce. Greater rhea and cattle have different foraging pat- bivoros. Sin embargo, un elevado solapamiento dietario no se traterns and hunting of greater rhea is not justifled solely on the duce en competencia a menos que el forraje resulte escaso. Por
basisof foragecompetitionwith cattle.
otra parte, el fiandu y el ganado presentan diferentes patrones de
forrajeo y la caceria que sufren estas aves no esta justificada utnicamente sobre la base de la competencia por el forraje.
rhea (Rhea americana) and cattle (Bos taurus) in the Parana

Key Words: diet composition, herbivory, Lagostomus maximus, Parana River Delta, Rhea americana

In farmingecosystems,cattle and otherdomesticspecies often

co-exist with wildlife. Roughly80%of Argentinais dedicatedto
extensive cattle ranching(CONAPA 1991) where cattle interact
with a large numberof wild species thatuse the same resources.
However,few studiesin Argentinahave quantifiedthese interacResearch was funded by the TX-16 and PICT 98 N? 04503 grants of the
UBACyTandthe FONCYTprogrammes,respectively.The authorswish to thank
Mr.I. Schojettfor allowingthe studyto be conductedon his ranch,D. Villarealfor
his bibliographycontributionandN. Madanes,M. Cagnoni,S. Arias,G. Aprile,N.
Fracassi,D. VarelaandF. Gagliardifor theirfield support.A specialthanksto L.
Azcoagafor herinvaluablehelp withthe Englishtranslation,Dr. 0. Solbrigfor his
supportat HarvardUniversity,andthe anonymousreviewersfor constructivecriticism.

Manuscriptaccepted14 May02.

tions (e.g., Bonino et al. 1986, Kufner and Pelliza 1987, Martella
et al. 1996, Quintana et al. 1998a, 1998b).
Plains vizcacha (Lagostomus maximus) and greater rhea (Rhea
americana) are 2 native species found in these ecosystems. Plains
vizcacha are large nocturnal rodents of the Chinchillidae family
that dwell in communal burrows ("vizcacheras")of grasslands and
semi-arid scrublandsfrom southernParaguayand Bolivia to central
Argentina (Llanos and Crespo 1952, Branch 1993). Greater rhea
also live in grasslands and bush country from Brazil and Bolivia to
central Argentina. These birds live in polygamous social clusters
and are generally associated with farming and cleared fields where
native vegetation has been replaced by improved pastures (Martella
et al. 1996, Reboreda and Fernandez 1997).
The plains vizcacha is often considered to be an agricultural
pest, damaging soil and vegetation (Weir 1974) due to its burrow-

JOURNAL OF RANGE MANAGEMENT56(1) January 2003

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13

ing and grazinghabits and causing losses

in yield of crops such as corn, soybean,
pastures,andhorticulturalspecies (Rendel
1990, Navarroet al. 1997). It is also hunted for its pelt (Rendel 1990, Bruggersand
Zaccagnini 1994) and used as a food
resource by humans (Mares and Ojeda
1984). Huntinghas severely reducedthe
numbersof plain vizcachas,and it is now
extinct in large tracts of the wet Pampa
(Redford and Eisenberg 1992). Greater
rheaalso have been huntedintensivelyfor
meat, feathersand skin, and was declared
a harmful species due to its negative
impact on crops (Bertonatti 1997) and
potentialcompetitionwith cattlefor forage
(Martellaet al. 1996).
There have been few scientific studies
on the feeding habitsof these wild herbivores despitetheir widespreadcontinental
distribution.Plains vizcacha are selective
grazers (Llanos and Crespo 1952) while
greaterrhea feed on vegetable matteras
well as arthropodsand small vertebrates
(Bruning1974).
This studyfocuses, for the first time, on
a wetland area of Argentina(the Parana
River Delta Region) and surveysthe food
habits of plains vizcachas, greaterrheas
andcattlesharingthe samegrazingarea.

Materialsand Methods
The studyareawas locatedon the "Don
Jose" Ranch (1,500 ha, 33?27'S,
58?48'W), 6 km north of Ceibas,
Departmentof Gualeguaychd,EntreRios
province,Argentina.Mean annualrainfall
is 978 mm, and temperatureaverages17.4
C' (Servicio Metereologico Nacional
1972). The studyareais situatedin one of
the 11 wetlandlandscapeunits identified
for the Parana'River Delta Region. The
landscape pattern correspondsto plains
with a savannaphysiognomy of grasses,
patches of xerophitic Prosopis nigra
(Griseb.) and Acacia caven (Mol.) Mol.
forest, and baldspotswith a large percentage of bare soil and sparse cover of
Portulaca sp. L. The area is also crisscrossed by small lentic streams covered
with floating and rooted aquatic plants
(Malvairez1997).
Unlike its neighboringareas, the study
area is free from the periodicalfloods of
the Parana'River. Large areas are waterlogged by rainfall due to the minimum
slope of the land (Malvairez1997) and the
type of soils (Pratolongo 2000). During
1998, coincidentwith an "El Ninio"event,
both the waterloggedareas and the duration of the waterloggingwere longerthan
for 1996/97.
14

Soils are composed of a clayey, sodic

horizon close to the surface.These sodic
clays hamperwater penetrationbecause
they swell when wet, creatinga layer that
is effectively impermeable.Tramplingby
cattle compounds the problem by compressingand hardeningthe soil, and eroding thin layers of topsoil (Arias 2000).
Extensive cattle ranching (0.7 cows per
hectare per year), hunting of wildlife
species for food and trade,and exploitation of P. nigra trees for lumberare the
mainhumanactivitiesin this region.
Vizcachasdig theirburrowsboth in the
grasslandsand the forestpatchesand their
foraging is restricted to the areas surroundingthe burrows(Branch and Sosa
1994, Arias2000). Greaterrheaandcattle,
on the other hand, graze over extensive
areas,including,in the case of cattle,vegetation from the streams. Cattle were
always presenton the study area and the
overall level of forage utilization was
moderate.
Fresh faeces of plains vizcachas and
cows were collected seasonallyfor 2 consecutive years, between November 1996
(spring) and August 1998 (winter).
Greaterrhea fresh faeces were collected
only in spring/summer1997 and fall/winter 1998, whenthesebirdswere presenton
the study area. Collection dates were in
the middle of the respective season. All
faeces were collected within the single
grazingunitthatcomprisedthe studyarea.
Faeces of plains vizcachas were taken
from 8 activeburrowslocatedon the edge
between grassland and forest patches,
while faeces of greater rhea and cattle
were collectedfromdroppingsfoundboth
in grasslandand forest patches.We verified the vizcachas' burrowswere active
throughdirectobservationof the animals,
or signs of recent activity such as fresh
faeces, footprintsor diggings (Branchet
al. 1994b). Fifty pellets were collected at
each vizcachaburrow(a total of 400 pellets per season)to form 8 compositesamples (1 sampleper burrowwith 50 pellets
each).This is an adequatenumberof samples to estimatethis rodent'sdiet (Bontti
et al. 1997).Forgreaterrheaandcattle,32
and 24 samplesof faeces of 4 g were collected, respectively, and formed into 8
composite samples for both herbivores
(eachcomposedof 4 and3 faeces).
The botanicalcompositionof the diets
was determinedby means of microhistological analysisusing the William's technique (1969). Four slides were prepared
from each of the composite samples and
100 randomlychosen microscopic fields
were observed at 400x for each slide

(Holechek and Vavra 1981, Holechek et

al. 1982). Frequenciesof each consumed
item were convertedto percentagesof the
total sample weight (Holechekand Gross
1982,Martellaet al. 1996).
Compositesamplesof greaterrheaunderwent a special procedure;whole or fragmented pods and seeds of P. nigra and
arthropodswere separatedfrom the herbaceouscomponentsandeachof these3 componentswas driedandweighed.Herbaceous
componentswere then analyzedfollowing
thetechniquedescribedabove.
The percentof each food item was calculatedfor each season. Diet correlations
and dietarysimilarityamong seasons and
among herbivore species were analyzed
using two-tailedSpearman'srankcorrelation coefficients (Zar 1996) and
Kulczynski'ssimilarityindex (Smith and
Shandruk 1979, Henley et al. 2001),
respectively.Datawere averagedfor the 2
years for plains vizcacha and cattle to
compare botanical composition of diets
amongseasonsandherbivores.

Results
While vizcachas and cattle fed exclusively on plant leaves, greaterrhea also
fed on P. nigra pods and consumedsome
arthropods(Tables1, 2, and3). Plainsvizcachafed on a wide varietyof plantsduring all seasons (Table 1). A total of 45
plant species were identified in vizcacha
diets over the lengthof the study.Grasses
were the staplediet of this rodent,both in
amount consumed and in number of
species, 42.7% of the diet in winter with
20 species to 65.1% of the diet in spring
with 17 species. Panicum milioides Nees.

was the most consumedgrass, contributing 8.8% and 16.4%of the diet in winter
and spring,respectively.The item "Other
Grasses"was at times similar to P. milioides (14.8% vs. 15.1% in summer)or
even higher (13.4% vs. 8.8% in winter).
Other major items were Dichondra micro-

calyx (Hallier)Fabris.(20.4%and 19.3%,

fall and winter)and P. nigra (18.7% and
13.2%in fall andwinter,respectively).
Legumes were the most common component of the greater rhea diet during
spring and summer (25.3% and 38.9%;
Table 2) while grasses were more common in fall andwinter(35.8%and45.3%).
The high content of legumes was due
mainly to the consumption of P. nigra
pods (21.2% and 37.8%, for spring and
summer,respectively).Plantago myosuros
Lam.(20.5%)andSolanaceae,particularly
Solanumsp. L. (13.9%)were also important in the spring diet while Spilanthes

JOURNAL OF RANGE MANAGEMENT56(1) January 2003

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Table 1. Botanical composition (%) of plains vizcacha diets in different seasons in the Delta of the Parana River.

Food item
Perennialgrasses
Aristida sp.
Briza sp.
Bromussp.
Cynodondactylon (L.)
Chloris berroi Arech.
Eleusine tristachya(Lam.) Lam.
Festuca sp.
Luziolaperuviana Gmel.
Panicum milioides Nees.
Pappophorumsp.
Paspalum spp.
Piptochaetiumnapostaense Lam.
Poa sp.
Setaria sp.
Sporobolusindicus (L.) R. Br.
Stipa brachychaetaGodr.
Stipa neesiana Trin. et Rupr.
Stipa sp.
Zizaniopsisbonariensis
(Balansaet Poitr.) Speg.
Total perennialgrasses
Annual grasses
Hordeumeuclaston Steud.
Loliumsp.
Phalaris sp.
Polypogon monspeliensis(L.) Desf.
Total annualgrasses
OtherGrasses
Grass-likeplants (Cyperaceae)
Carex bonariensis Desf.
Eleocharis sp.
OtherCyperaceae
Total grass-likeplants
Perennialforbs
Alternantheraphiloxeroides (Mart.)
Griseb.
Chenopodiaceae
Dichondra microcalyx(Hallier)Fabris.
Eichhorniaazurea (Sw.) Kunth.
Eryngiumsp.
Gomphrenapulchella Mart.
Holocheilus hieracioides (Don) Cabr.
Modiolastrumsp.
Oxalis sp.
Pamphalea bupleurifoliaLess.
Phyla canescens (HBK) Greene
Physalis viscosa L.
Solanumsp.
Spergularialevis Camb.
Spilanthesstolonifera (H. et A.) Baker
Trifoliumsp.
Total perennialforbs
Annual forbs
Gamochaetasp.
Medicago sp.
Plantago myosurosLam.
Total annualforbs
Shrubsand trees
Acacia caven (Mol.) Mol.
Prosopis nigra (Griseb.)
Total shrubsand trees

stolonifera

1997

Winter
1998

3.3
1.0
0.0
1.2
0.7
0.5
0.0
8.0
12.1
0.9
1.4
0.1
0.7
1.6
0.0
0.2
1.0
0.0
2.3

3.8
2.7
0.0
0.3
0.7
0.3
0.0
5.2
9.7
0.5
0.3
0.0
0.3
0.0
0.0
0.0
0.0
0.0
3.2

0.5
0.0
0.5
1.5
0.0
1.5
0.0
4.0
8.0
0.0
0.0
0.0
0.0
0.4
0.0
0.7
0.1
0.1
0.4

2.2
1.3
0.2
0.9
0.3
0.9
0.0
4.6
8.8
0.2
0.2
0.0
0.2
0.2
0.0
0.4
0.1
0.1
1.8

23.7

35.0

27.0

17.7

22.4

0.0
0.7
0.5
0.2
1.4
0.0

0.2
6.8
0.4
0.0
7.4
9.9

0.1
3.8
0.4
0.1
4.4
4.9

0.0
0.0
1.3
3.3
4.6
17.5

0.1
7.5
0.5
0.9
9.0
9.4

0.1
3.7
0.9
2.1
6.8
13.4

2.9
0.2
4.4
7.5

0.7
0.0
6.8
7.5

3.4
0.0
10.1
13.5

2.0
0.0
8.5
10.5

1.5
0.0
9.0
10.5

4.6
0.0
7.1
11.7

3.0
0.0
8.0
11.0

0.0

0.0

0.0

0.4

0.2

0.0

1.6

0.8

0.0
11.2
2.7
0.0
3.2
0.0
0.0
0.0
0.7
4.0
1.3
0.7
0.0
0.7
0.3
24.8

0.0
14.2
0.0
0.0
2.2
0.0
0.1
0.0
0.0
1.5
0.0
1.7
0.0
1.4
0.1
21.2

0.0
12.7
1.3
0.0
2.7
0.0
0.1
0.0
0.3
2.7
0.7
1.2
0.0
1.0
0.2
22.9

0.3
21.0
0.0
0.0
0.7
0.0
0.2
0.0
0.0
2.0
0.0
0.3
0.0
0.0
2.7
27.2

0.4
19.7
0.0
0.4
0.0
0.1
0.0
0.0
0.0
0.4
0.5
0.1
0.0
0.6
0.9
23.5

0.3
20.4
0.0
0.2
0.3
0.1
0.1
0.0
0.0
1.2
0.2
0.2
0.0
0.3
1.8
25.3

0.0
15.7
0.0
0.0
2.2
0.0
0.2
0.2
0.0
0.2
0.0
0.2
0.0
0.3
0.2
19.2

0.5
22.9
0.0
0.0
2.5
0.5
0.6
0.0
0.0
0.0
0.0
1.5
0.1
2.2
2.9
35.3

0.2
19.3
0.0
0.0
2.3
0.2
0.4
0.1
0.0
0.1
0.0
0.8
0.1
1.3
1.5
27.1

1.1
0.0
4.2
5.3

0.0
0.0
0.0
0.0

0.7
0.0
0.6
1.3

0.4
0.0
0.3
0.7

0.5
0.0
0.0
0.5

0.0
0.9
0.1
1.0

0.2
0.4
0.1
0.7

1.2
0.0
0.8
2.0

2.0
0.0
7.0
9.0

1.6
0.0
3.9
5.5

0.0
4.4
4.4

0.2
1.5
1.7

0.0
7.7
7.7

0.1
4.6
4.7

0.8
16.7
17.5

0.0
20.7
20.7

0.4
18.7
19.1

0.5
18.8
19.3

0.1
7.6
7.7

0.3
13.2
13.5

1997

Fall
1998

0.9
0.0
0.0
2.6
0.6
2.6
0.0
7.8
18.6
0.7
0.7
0.0
0.0
0.1
0.0
2.1
1.1
0.0
5.6

0.4
0.0
0.0
5.5
0.8
1.9
0.0
7.0
15.1
1.0
1.5
0.0
3.8
0.3
0.3
4.8
1.1
0.0
3.4

6.2
2.0
0.0
1.2
1.0
0.3
0.0
10.7
12.7
1.8
2.8
0.0
1.3
2.5
0.0
0.2
0.2
0.0
3.2

0.4
0.0
0.0
1.2
0.4
0.6
0.0
5.4
11.6
0.0
0.0
0.1
0.0
0.6
0.0
0.2
1.7
0.0
1.5

50.5

43.4

46.9

46.1

1.0
0.0
0.2
6.6
7.8
8.2

0.5
0.0
0.0
0.7
1.2
21.5

0.0
2.7
0.4
0.3
3.4
8.1

0.2
1.4
0.2
0.5
2.3
14.8

3.6
0.0
6.5
10.1

1.8
0.0
4.9
6.7

0.3
0.3
0.0
0.6

5.5
0.0
8.7
14.2

0.0

0.0

0.0

0.0

0.0
14.3
0.0
0.0
0.5
0.0
0.0
0.0
0.0
2.0
0.0
1.7
0.0
0.0
0.0
18.5

0.0
15.0
0.0
0.0
1.2
0.0
0.0
0.0
1.2
0.7
0.0
0.1
0.0
0.0
0.1
18.3

0.0
14.7
0.0
0.0
0.9
0.0
0.0
0.0
0.6
1.4
0.0
0.9
0.0
0.0
0.1
18.6

0.0
0.0
3.3
3.3

2.2
0.0
5.0
7.2

0.0
6.0
6.0

0.0
2.7
2.7

Summer
1997 1998

1996

Spring
1997

0.0
0.0
0.0
5.5
15.2
5.5
3.3
0.0
16.5
0.0
0.0
0.0
0.0
0.0
0.0
0.0
12.7
0.0
0.0

9.2
3.6
0.0
1.0
0.4
0.4
0.0
4.5
16.4
0.5
0.5
0.0
0.0
0.6
0.0
0.0
0.4
0.0
1.9

4.6
1.8
0.0
3.2
7.8
2.9
1.7
2.2
16.4
0.2
0.2
0.0
0.0
0.3
0.0
0.0
6.5
0.0
0.9

0.0
0.0
0.0
8.3
1.2
1.2
0.0
6.3
11.5
1.2
2.2
0.0
7.7
0.5
0.7
7.5
1.0
0.0
1.2

58.7

39.4

48.7

2.0
0.0
0.0
5.3
7.3
2.8

0.0
0.0
0.5
7.9
8.4
13.6

0.0
0.0
3.3
3.3

(H. et A.) Baker was a major

item in summer(14.9%)and D. microcalyx was the dominant food item in fall

(34.7%). In winter, "OtherDicots" sup-

plied 18.2%of the diet. Therewere traces

Grasseswere also the dominantgroupin
of arthropodconsumptionthroughoutthe the cattlediet throughoutthe year,ranging
year, with the highest values in spring between78.3%in fall and 87.3%in sum(2.9%)andwinter(3.0%).
mer (Table 3). Luziola peruviana Gmel.

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15

Table 2. Botanicalcomposition(%) of greaterrhea diets in differentseasonsin the Delta of the withinseasonandthe similarityindex valParanaRiver.
ues oscillated between 50.5 and 57.9
Summer
1997

Spring
1997

Food item

- ----------------(
Perennialgrasses
Aristida sp.
Bromus sp.
Chloris berroi Arech.
Luziolaperuviana Gmel.
Panicum milioides Nees.
Pappophorumsp.
Setaria sp.
Zizaniopsisbonariensis
(Balansaet Poitr.) Speg.
Total perennialgrasses
Annual grasses
Lolium sp.
Phalaris sp.
Total annualgrasses
OtherGrasses
Grass-likePlants (Cyperaceae)
Carex bonariensis Desf.
OtherCyperaceae
Total grass-likeplants
Perennialforbs
Chenopodiaceae
Dichondra microcalyx
(Hallier) Fabris.
Gomphrenapulchella Mart.
Holocheilus hieracioides (Don) Cabr.
Oxalis sp.
Pamphalea bupleurifoliaLess.
Phyla canescens (HBK) Greene
Physalis viscosa L.
Solanumsp.
Spilanthesstolonifera (H. et A.) Baker
Trifoliumsp.
Total perennialforbs
Annual forbs
Medicago sp.
Plantago myosurosLam.
Total annualforbs
Shrubsand trees
Acacia caven (Mol.) Mol.
Prosopis nigra (Griseb.)leaves
Prosopis nigra (Griseb.)pods
Lyciumsp.
Total shrubsand trees
OtherDicots
Arthropods

%)0.0
0.9
0.5
0.5
0.0
0.2
0.0
0.0

0.5
0.0
0.2
8.1
1.9
0.0
0.5
1.2

0.0
0.0
0.2
10.7
2.9
0.0
0.0
6.1

0.8

2.1

12.4

19.9

0.0
0.0
0.0
7.2

0.0
0.0
0.0
6.0

2.1
0.0
2.1
21.5

2.9
0.7
3.6
21.8

0.0
0.0
0.0

0.0
1.0
1.0

0.7
0.0
0.7

3.4
4.6
8.0

1.0
4.8

0.9
1.6

0.2
34.7

0.0
6.3

10.8
0.0
1.1
0.0
9.1
0.0
13.9
1.9
1.0
43.6

13.0
0.0
1.0
0.6
13.1
1.1
1.7
14.9
0.0
47.9

0.0
0.5
0.0
0.0
4.2
0.0
0.2
1.9
0.0
41.7

1.0
0.5
0.0
0.0
1.0
0.0
0.0
2.2
0.0
11.0

0.2
20.5
20.7

0.0
0.3
0.3

5.8
6.5
12.3

3.6
8.2
11.8

0.0
3.0
21.2
0.8
25.0
0.0
2.9

0.3
0.8
37.8
0.0
38.9
1.0
0.8

0.0
0.5
6.6
0.2
7.3
1.4
1.0

0.0
2.7
0.0
0.0
2.7
18.2
3.0

with Zizaniopsis bonariensis (Balansa et

Poitr.) Speg. somewhat lower (5.8% in

springand 9.8% both in winterand summer). The item "OtherGrasses"averaged
about 15%over all seasons.Luziolaperutogether with

"OtherGrasses",constituted60%, 63%,
and58%of the spring,summerandwinter
diets,respectively.
Withinseasons,vizcachaandcattlediets
differedin botanicalcompositionbetween
years (Table4). Summerand winterdiets

16

Winter
1998

0.0
0.0
0.0
0.2
0.6
0.0
0.0
0.0

and P. milioides were commonly eaten

(the formerbetween 13.8%in winterand
25.0% in summer,and the latter ranging
from 9.3% in winterand 20.0%in spring)

viana and P. milioides

Fall
1998

of the vizcachawere significantlycorrelated between years but the correlationvalues were low. Similarity index values
showed the same trend, except for fall,
when similaritywas highest(Table 4).
Plains vizcacha and cattle diets were
similaramongthe differentseasons,something that matchesthe ratherhigh values
of the similarityindex (Table5). The diet
of greaterrhea differed among most seasons, being similar only between spring
and summerand betweenfall and winter.
Both comparisonsalso showedthe highest
similarityvaluesbetweendiets (Table5).
Botanical composition of the diets of
plains vizcacha and cattle was similar

(Table 6). Inversely,plains vizcacha and

greater rhea segregated their grazing
resources,withoutsignificantassociations
between diets, except in winter, which
again had the highest similarity index
value (Table 6). Cattle and greaterrhea
diets differedmost with significantnegative correlations;also, the observedsimilarity index's values showed low values,
especiallyin springandsummer(Table6).

Discussion
Grasseswere the main foragefor plains
vizcacha and cattle in all seasons. For
greaterrhea,grasseswere the most important diet componentwhen P. nigra pods
were lacking.The importanceof grassesin
the plains vizcacha diet has also been
reportedby other authors (Giulietti and
Jackson 1986, Kufner et al. 1992, Jofre
1994, Branchet al. 1994a, Navarroet al.
1997).Althoughplainsvizcachagrazedon
a largevarietyof plantspecies, only a few
food itemscomposedthe bulkof its diet in
each season.This last fact was observedin
other grassland habitats (Giulietti and
Jackson 1986), although we found a
greaterconsumptionof dicotscomparedto
that study (23 species versus 3 species).
Ourresultsare more similarto those from
the semiaridscrub of CentralArgentina,
where53%of the species in the diet were
dicots (Branchet al. 1994a). The number
of items consumed by vizcachas in the
Delta Region was 45, while Giuliettiand
Jackson(1986) and Branchet al. (1994a)
record 20 and 62 items in grasslandand
scrub,respectively.The lower numberof
items found in the grassland might be
explained by higher availabilityof more
palatablespecies, a fact that might allow
the rodentsto fulfill theirnutritionalneeds
with fewer species, in agreement with
classic foraging theory (Stephens and
Krebs1986).
In agreement with other studies
(Martella et al. 1996, Comparatoreand
Martinez 1997), greaterrhea had a high
intake of greens throughoutthe year. P.
nigra pods were an importantfood item
during certain parts of the year. These
highly nutritiouspods appearin spring,
reach their peak during summer, their
availabilitydecreasesin fall, andthey cannot be found in winter(Pratolongo2000),
which correlateswith their abundancein
the rheadiet.
Variationsin digestibilityof the different food itemscould be takingplace in the

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Table 3. Botanical composition (%) of cattle diets in different seasons in the Delta of the Parana River.
Summer
Fall
Winter
Spring
1997
1998
x
1997
1998
x
1997 1998
x
1996
1997
-------(-%-)

Food item
Perennialgrasses
Aristida sp.
Briza sp.
Cynodondactylon (L.)
Chloris berroi Arech.
Eleusine tristachya(Lam.) Lam.
Festuca sp.
Luziolaperuviana Gmel.
Panicum milioides Nees.
Pappophorumsp.
Paspalum sp.
Poa sp.
Setaria sp.
Stipa brachychaetaGodr.
Stipa neesiana Trin. et Rupr.
Zizaniopsis bonariensis .
(Balansaet Poitr.) Speg
Total perennialgrasses
Annual grasses
Hordeumeuclaston Steud.
Loliumsp.
Phalaris sp.
Polypogon monspeliensis(L.) Desf.
Total annualgrasses
OtherGrasses
Grass-likePlants (Cyperaceae)
Carex bonariensis Desf.
OtherCyperaceae
Total grass-likeplant
Perennialforbs
Alternantheraphiloxeroides (Mart.)
Griseb.
Chenopodiaceae
Dichondra microcalyx(Hallier)Fabris.
Gomphrenasp.
Phyla canescens (HBK) Greene
Solanumsp.
Spergularialevis Camb.
Spilanthesstolonifera (H. et A.) Baker
Trifoliumsp.
Total perennialforbs
Annual forbs
Gamochaetasp.
Plantago myosurosLam.
Total annualforbs
Shrubsand trees
Prosopis nigra (Griseb.)
OtherDicots

0.0
0.0
3.5
4.0
0.0
5.0
21.5
31.0
0.0
0.0
0.0
0.0
0.0
4.0
0.0

10.5
5.5
0.0
0.0
0.0
0.0
26.5
9.0
0.0
0.0
0.0
0.0
0.0
0.0
11.5

5.3
2.7
1.8
2.0
0.0
2.5
24.0
20.0
0.0
0.0
0.0
0.0
0.0
2.0
5.8

5.0
0.0
0.5
5.5
4.0
0.0
22.0
14.0
0.0
0.0
1.5
0.0
1.0
0.0
3.0

0.0
0.0
0.0
1.0
2.5
0.0
28.0
11.5
0.0
0.0
0.0
2.5
0.0
3.5
16.5

2.5
0.0
0.3
3.3
3.3
0.0
25.0
12.8
0.0
0.0
0.8
1.3
0.5
1.8
9.8

3.5
2.0
3.0
4.0
0.0
0.0
17.5
14.0
0.0
0.5
0.0
5.5
0.0
2.5
7.5

0.0
0.0
4.0
0.0
1.5
0.0
11.0
8.5
3.5
0.0
0.0
0.0
0.0
9.0
8.5

1.8
1.0
3.5
2.0
0.8
0.0
14.3
11.3
1.8
0.3
0.0
2.8
0.0
5.8
8.0

8.0
12.0
0.0
0.0
0.0
0.0
13.0
4.0
0.0
0.0
3.0
0.0
0.0
0.0
11.5

0.0
0.0
0.0
0.0
0.0
0.0
14.5
14.5
0.0
0.0
0.0
0.0
0.0
0.0
8.0

4.0
6.0
0.0
0.0
0.0
0.0
13.8
9.3
0.0
0.0
1.5
0.0
0.0
0.0
9.8

69.0

63.0

66.1

56.5

65.5

61.4

60.0

46.0

53.4

51.5

37.0

44.4

1.0
0.0
0.0
3.5
4.5
6.5

0.0
0.0
0.0
0.0
0.0
25.0

0.5
0.0
0.0
1.8
2.3
15.8

0.0
0.0
0.0
0.0
0.0
35.5

0.0
2.5
0.0
0.0
2.5
14.5

0.0
1.3
0.0
0.0
1.3
25.0

0.0
0.0
3.0
0.0
3.0
16.0

0.0
9.0
1.5
0.0
10.5
21.0

0.0
4.5
2.3
0.0
6.8
18.5

0.0
0.0
0.0
0.0
0.0
37.5

0.0
9.0
1.0
0.0
10.0
31.5

0.0
4.5
0.5
0.0
5.0
34.5

0.0
0.5
0.5

6.0
2.5
8.5

3.0
1.5
4.5

0.0
0.0
0.0

12.5
0.0
12.5

6.3
0.0
6.3

0.0
3.0
3.0

7.5
8.5
16.0

3.8
5.8
9.6

1.0
0.0
1.0

5.5
4.0
9.5

3.3
2.0
5.3

0.5

0.0

0.3

0.0

0.0

0.0

0.0

0.0

0.0

0.0

0.0

0.0

0.0
7.0
0.0
0.5
0.0
0.0
0.5
0.0
8.5

0.0
1.5
0.0
0.0
0.0
0.0
0.0
0.0
1.5

0.0
4.3
0.0
0.3
0.0
0.0
0.3
0.0
5.2

0.0
1.0
1.0
2.0
1.0
0.0
0.0
0.5
5.5

0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0

0.0
0.5
0.5
1.0
0.5
0.0
0.0
0.3
2.8

1.0
4.0
0.0
0.0
0.0
1.0
0.0
5.5
11.5

0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0

0.5
2.0
0.0
0.0
0.0
0.5
0.0
2.8
5.8

0.0
2.5
0.0
0.0
0.0
0.0
0.0
1.0
3.5

1.0
5.0
0.0
0.0
0.0
0.0
2.0
0.0
8.0

0.5
3.8
0.0
0.0
0.0
0.0
1.0
0.5
5.8

0.0
0.5
0.5

0.5
0.0
0.5

0.3
0.3
0.6

0.0
2.5
2.5

0.0
0.0
0.0

0.0
1.3
1.3

0.0
0.0
0.0

0.0
0.0
0.0

0.0
0.0
0.0

0.0
0.5
0.5

0.0
0.0
0.0

0.0
0.3
0.3

10.5
0.0

1.5
0.0

6.0
0.0

0.0
0.0

5.0
0.0

2.5
0.0

6.5
0.0

6.5
0.0

6.5
0.0

2.5
3.5

4.0
0.0

3.3
1.8

digestive tract.Plantpartsin the rheafaeces were hardly degraded. There were

only tracesof arthropodsin the faeces, in
spite of their high abundancein the field
throughoutthe year. As with Martellaet
al. (1996), these traceswere highly digested, unlikethe plantmaterial.This seems to
indicatethatthis methodfor diet quantification is unsuitable to analyze the food
habitsof the greaterrhea, because differences in digestibility produce a bias in
establishing the true proportionof each
item in its diet (Moreby 1988, Rosenberg
andCooper1990, Martellaet al. 1996).
Although the results obtainedby com-

paringthe diet compositionsby means of

correlationsand similarityindices showed
on the whole a similar trend, the former
were clearer when comparing between
years,seasonsandspecies.
Differencesobservedin botanicalcompositionof the diets of plainsvizcachaand
cattlebetweenthe 2 yearscould be due to
environmentalvariability.Observeddifferencesin rainfallmightaccountfor variability in both vegetationabundanceand
grazingareabecauseof the waterlogging,
resultingin changesin forageavailability.
Plains vizcacha avoid waterloggedareas.
Waterloggingmay also provoke changes

in the vegetationwhich may in turninfluence changesin the grazingpatternsfrom

one year to the next. While some grasses
were eaten less in the wettest year (e.g.
Chloris berroi, Stipa neesiana, Polypogon

monspeliensis),intakeof Cyperaceaegrew
(See Tables 1 and 3). Despite the yearly
changesin the foragingpatterns,however,
the staple components of the diet kept
their high values between the 2 years of
the study (e.g. Dichondra microcalyx and
Panicum milioides for both vizcachas and
cattle and Luziola peruviana for cattle).

Thus, these environmentalchangescorrespondedto changesin the grazingpatterns

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17

Table 4. Comparison of the composition of plains vizcacha and cattle diets (Spearman's rank correlation coefficient, rS and Kulczynski's similarity index, K) between the same season of different years in the Parana River Delta.

Comparisona

nb

Plains vizcacha
P
rs

SP 96 - SP 97
SU 97 - SU 98
F 97 - F 98
W97-W98

28
36
39
39

0.10
0.35
0.30
0.42

0.61
0.04
0.07
<0.01

Cattle
K

rs

51.7
54.7
68.5
61.4

21
21
22
18

0.01
0.17
0.23
0.21

0.96
0.47
0.30
0.41

40.5
54.5
59.5
62.5

aSP = Spring;SU = Summer;W = Winter;F = Fall.

n = Numberof food items.

of bothherbivores.Somethingsimilarwas
observedfor wild anddomesticherbivores
in central Entre Rios (Quintana et al.
1998b).This underscoresthe need to conduct surveys for at least 2 years, as proposedby HansenandLucich(1978).
Variationsobservedin the greaterrhea
diet throughoutthe year may be explained
by changes in basic nutritionalrequirementsassociatedwith reproductiveactivities (Bruning 1974, Robbins 1981,
Lombardi 1994, Martella et al. 1995,

of greater rhea were collected in 1997,

while the fall and winter samples were
from 1998, one might thinkthat variation
of intakeof Prosopis nigra pods between
spring/summerand fall/wintercould also
be due to environmentalchangesthattook
place from one year to the next. Yet,
intake of these pods correspondedto the
availability

of Prosopis

nigra fruits

throughthe year in this region (Burkart

1976, Pratolongo 2000), supportingour
previousargument.

europaeus)(Boninoet al. 1986).Increased

intake of P. nigra leaves by plains vizcachain fall and summerdid not coincide
with the results for cattle (except for a
slightintakein fall) andthis maybe due to
the wider range of movementof cattle as
comparedto thatof the rodents.Cattlehad
easy access to areas with more tender,
palatablegrass,a preferredforage(Hansen
and Gold 1977, Vavraet al. 1977, Samuel
andHoward1982).This foragewas not as
availablefor vizcachasbecause their forof
aging was restrictedto the surroundings
their burrows (Branch and Sosa 1994,
Arias 2000). Greaterrhea diet was more
similarto those of the other 2 species in
seasons where the intake of Prosopis
seeds decreased.
Accordingto Kufneret al. (1992), the
largernumberof itemsin the diet of plains
vizcachasseems to indicatebetteradapted
grazing habits than those of greaterrhea
and cattle, when faced with the resources
available in their habitat. However, the

Table 5. Comparison of the composition of plains vizcacha, greater rhea and cattle diets (Spearman's rank correlation coefficient, rS and Kulczynski's
similarity index, K) among seasons in the Parana River Delta.

Comparisona

nb

SP- SU
SP-F
SP-W
SU - F
SU-W
F-W

38
41
41
43
44
43

Plains vizcacha
r5
P
0.38
0.47
0.51
0.61
0.50
0.60

0.02
<0.01
<0.01
<0.01
<0.01
<0.01

Cattle
K

r5

63.1
56.9
61.6
62.0
64.6
76.6

29
30
27
28
27
26

0.48
0.47
0.48
0.50
0.31
0.48

<0.01
<0.01
0.01
<0.01
0.11
0.01

71.2
67.9
63.3
68.0
69.0
68.6

24
25
25
30
28
25

Greaterrhea
r5
P
0.58
0.17
-0.17
0.07
-0.17
0.56

<0.01
0.42
0.43
0.72
0.36
<0.01

K
56.6
34.1
30.6
23.3
14.9
58.2

aSP = Spring;SU = Summer;W = Winter;F = Fall.

n = Numberof food items.

Reboredaand Fernandez1997). Intakeof

insects, small vertebrates,and seeds satisfy the needof minerals,vitamins,proteins,
or specific nutrients in larger or lesser
demandaccordingto the season (Robbins
1981, Martellaet al. 1996). This intake
was higherduringspringand summer,the
mating season for this species (Reboreda
and Fernandez1997). Consequently,the
similardiets observedin these seasonsand
theirdifferencewith the fall/winterintake
is logical. Since springand summerfaeces

relevanceof grasses as a resourceshared

by this rodentand cattle implies a significant dietaryoverlapthatmightbe a negative factor for the remainingpopulations
of vizcacha in this area. Plains vizcacha
and cattle had similar diets, which supports the ranchers' view that vizcachas
competewith domesticherbivoresfor foraging resources and leads to their being
huntedas pests. However,high overlapin
the use of resourcesimplies competition
(Do lichotis patagonum) (Kufner and
Pelliza 1987) and Europeanhares (Lepus only if the resources are scarce (Wiens
1989). Huntingof greaterrheaon ranches
Grazing on common grounds explains
the similarityobservedin botanicalcomposition of diet observed throughoutthe
year between plains vizcacha and cattle,
includinga few commondominantspecies
(mainlygrasses)and a wide rangeof less
relevant species (<3%). Diet similarities
between cattle and medium-sized wild
herbivores such as the plains vizcacha
have also been reported for mara

Table 6. Comparison of the composition of plains vizcacha (PV), greater rhea (GR) and cattle (CA) diets (Spearman's rank correlation coefficient, r'
and Kulczynski's similarity index, K) in different seasons in the Parana River Delta.
PVvsCA
Spring
Summer
Fall
Winter

na

rs

30
36
40
40

0.70
0.57
0.82
0.64

P
<0.01
<0.01
<0.01
<0.01

56.7
53.6
57.9
50.5

35
41
43
42

PVvsGR
P
rs
-0.19
0.01
0.22
0.41

0.29
0.98
0.16
<0.01

GRvsCA
P

rs

23.3
19.0
43.3
49.6

30
32
32
24

-0.38
-0.38
-0.04
0.34

0.04
0.04
0.83
0.11

K
16.1
10.8
36.2
59.6

an = Number
of fooditems.

18

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sis. J. RangeManage.34:337-338.
cacha (Lagostomus maximus, family
and farmshas been justified for the same
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