Interindividual interactions of Molina’s hog-nosed skunks Conepatus chinga in the Pampas grassland of Argentina, --home range overlap, dynamic interactions, static interactions,

8/14/2019 Interindividual interactions of Molinas hog-nosed skunks Conepatus chinga in the Pampas grassland of Argentina, --home range overlap, dynamic interaction

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Introduction

In general, the social organization of solitaryspecies is based on territoriality: both males andfemales defend their territories from other individuals of the same sex (Rogers 1977, Powell

1979). In this context, males usually range overlarger areas than females, and home range over-lap is wider intersexually than intrasexually(Gittleman and Harvey 1982). The purpose of this is to enable males to access to more than onefemale during mating (Kruuk and Macdonald

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Acta Theriologica 54 (1): 8794, 2009.PL ISSN 00017051

Interindividual interactions of Molinas hog-nosed skunksConepatus chinga in the Pampas grassland of Argentina

Juan I. REPPUCCI, Diego F. CASTILLO*, Mauro LUCHERINI, Estela M.

LUENGOS VIDAL and Emma B. CASANAVE

Reppucci J. I., Castillo D. F., Lucherini M., Luengos Vidal E. M. and CasanaveE. B. 2009. Interindividual interactions of Molinas hog-nosed skunks Cone - patus chinga in the Pampas grassland of Argentina. Acta Theriologica 54:8794.

During one year, we radiotracked two female and two male Molinashog-nosed skunks Conepatus chinga (Molina, 1782), a little studied mephitid,in the Pampas of central Argentina, to analyze the static and dynamic inter-actions between individuals. Mean home range overlap was large (44.5%), butmales shared a smaller proportion of their home ranges than females. Theaverage percentage of localizations in the overlap area (39.3%) indicates thatthese areas were not marginal sections of individual home ranges, but this value was greater for individuals of different genders than intrasexual dyads.The Coefficient of Spatial Association confirmed that the distances betweenindividuals of different sexes were smaller than between animals of the samesex. During simultaneous locations, females were closer than males, butreciprocal distances between individuals varied over the time. These patternsare congruent with those described as typical for mustelids and mephitids.Our results confirm that an analysis of home range overlap should not beconsidered complete without the study of dynamic interactions among individuals and their temporal variations, which are necessary to overcome thelimitations of spatial overlap analysis.Grupo de Ecologa Comportamental de Mamferos (GECM), Ctedra Fisiologa Animal, De-partamento Biologa, Bioqumica y Farmacia, Universidad Nacional del Sur, San Juan 670, (8000)Baha Blanca, Argentina (JIR, DFC, ML, EMLV, EBC); CONICET Researcher (ML, EBC)

Key words : home range overlap, dynamic interactions, static interactions,

Mephitidae, radiotelemetry

* Corresponding author e-mail: [email protected]


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1985, Eisenberg 1986). This social organizationsystem is common among the carnivore Families(Kruuk and Macdonald 1985), but has never beenstudied in the South American skunks of genusConepatus.

Until some years ago, skunks were consideredto be members of the Mustelidae family. However,genetic evidence suggested that skunks arephylogenetically distinct from other mustelids,and they were placed in a new family, called Mephitidae (Rosatte and Lariviere 2003). TheMolinas hog-nosed skunk Conepatus chinga(Molina, 1782) is one of the most widespreadSouth American species and is found from

Bolivia south through Uruguay and WesternParaguay into Argentina at least as far south asNeuqun province and central Chile (Redfordand Eisenberg 1992).

The analysis of home range overlap and theinteractions among the individuals that sharethese areas is used commonly in the analysis of the social organization of carnivores, whereterritoriality is frequently difficult to study(Powell 2000). Interindividual interactions canbe classified in two groups: static and dynamic.Static interactions are defined as the spatialoverlap between home ranges of two or more individuals. The study of static interactions repre-sents a first approach to the understanding of thespatial organization of species. However, if weaim to understand the importance of shared ar-eas, it is advisable to analyze dynamic interac-tions, which imply the addition of time factor andthe study of reciprocal positions and independentmovements of the individuals whose home rangesoverlap (Powell 2000). The comprehension of dy-namic interactions allows determining if two in-dividuals are using an area in contemporary form(Doncaster and Macdonald 1997, Powell 2000).

The present work aims to contribute to theunderstanding of interindividual interactionsand the social organization in C. chinga .

Study areaWe conducted our study at Parque Provincial Ernesto

Tornquist (PPET), located in the central part of the Ventaniamountain range (38 00S 62 00O), southern Buenos Airesprovince, Argentina. This protected area has a surface of approximately 6700 ha and a maximum altitude of 1240 m

a.s.l. The climate is temperate with mean annual precipita-tion of 500800 mm (Frangi and Bottino 1995). This area ispart of the Pampas ecoregion (USAID 1995) and vegetation

is characterized by native grassland (genuses Stipa , Pipto -chaetium , Festuca and Briza ; Cabrera 1976). Introducedplant patches ( Pinus , Cupressus, Acacia , Eucalyptus andspecimens of the Rosacea family) are also frequent (Zalbaand Villamil 2002). The vegetation in the study area wasextensively affected by the presence of a dense population of feral horses that feed on natural grasses, but there was alittle fenced area (24.5 ha), where horses were excluded andgrasses were denser and taller.

Material and methods

Capture and handling

Trapping was carried out from October 2002 to March2005. During the first two years, we used box traps, but, inspite of a relatively large trapping effort, we had little suc-cess (we only trapped 2 skunks). For this reason, we de-cided to use a different method, and spotlighted skunksfrom a vehicle to restrain them manually, using a blanketto avoid being sprayed. With this new method we trapped 6skunks. Restrained individuals were chemically immobi-lized for handling with a combination of ketamine hydro-chloride (24.64 mg/kg) and xylazine (1.96 mg/kg). Captureand handling procedures were conducted in accordance witha protocol elaborated by our team and tested in more than200 carnivore captures (Luengos Vidal et al . 2003), andwere authorized by Ministerio de Asuntos Agrarios, Buenos

Aires province (the governmental agency in charge of wild-life research and management).

Radiotelemetry

We fitted adult skunks with Telemetry Solutions (Con-cord, California) and AVM Instrument (Colfax, California)radiocollars, equipped with activity and mortality sensors.Radiotelemetry sampling was carried out on foot and homo-geneously distributed throughout the 24 hours. We usedstandard telemetry techniques to track skunks (White andGarrot 1990). Most (87%) locations were obtained using theHoming technique, which provides great precision (Whiteand Garrot 1990). The remaining locations were obtained bytriangulation (through the LOCATE II software; Nams 1990)of 3 bearings obtained with a 3-element hand-held yagiantenna and compass from points located through a GPSreceiver. To minimize telemetry error, locations with aminimum angle of intersection < 60 or a maximum angle> 120 were discarded.

We used Ranges V (Kenward and Hodder 1996) soft-ware to estimate the sizes of skunk home ranges and coreareas, as well as their percent overlap utilizing the Kernelestimator (Worton 1989). We calculated home ranges at95% isopleths (henceforward 95% K) to avoid including ex-cursions outside normally used area, while 45% isoplethswere used to define core areas. We utilized fixed Kernel es -timator because this method is superior to others to de-scribe spatial use (Worton 1995, Seaman and Powell 1996).

88 J. I. Reppucci et al .


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Study of interactions

We first calculated the sizes of overlap areas betweenindividual home ranges and core areas. Then we estimatedthe time spent by each individual in the portion of its homerange overlapping with another animal using the percent-age of locations in this area.

For the study of dynamic interactions, it is necessary todefine the distance at which an individual is aware of thepresence of a co-specific. While previous authors studying dynamic interactions adopted an arbitrary distance (Salvatori et al . 1999, Harrington and Macdonald 2008), we analyzedthe frequency distribution of distances between individualsand observed a marked decrease at distances of less than300 m. Over this base, we assumed that two skunks at lessthan 300 m are detecting one another, and we consideredthat two animals were together when the distance be-tween them was < 300 m.

We adopted the period of two hours as a threshold be-tween locations of two animals to assume them as contem-poraneous, based on the analysis of the distances traveledduring different time interval (D. Castillo, unpubl.). How-ever, the mean time between locations was 1h 03, the timelapse between locations was less than one hour in 64.6%and less than 1h 30 in 80.4% of the cases.

We calculated the coefficient of spatial association, asthe proportion of contemporaneous locations at distances J > 0) when dis -tances are greater than expected.

Finally, we analyzed distances between individuals inrelation to their activity stage (active-inactive), and the variation of these distances over the study period.

Results

We radiotagged 8 adult skunks, but for theanalyses of this paper we used only data fromthe four individuals (two males and two females)which were tracked simultaneously.

From October 2004 to October 2005, we col-lected an average ( SD) of 74.2 22.8 locationsper animal. Mean home range size was 109.1 51.8 ha. On average, male home ranges were 2.3times larger than female home ranges (Fig. 1).

Skunk interactions in the Pampas 89

0 500 m

M1

M2F1

F2

Fig. 1. Spatial distribution of home ranges (95% Kernel estimator) of 4 C. chinga individuals in the Pampas grassland of Ar-gentina. The shaded polygon indicates the fenced area from which horses were excluded. F female, M male.


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Mean home range overlap was 44.5% (Table1). The home range of F1 was almost completelyincluded in M2s. The two males shared a slightlymore reduced proportion of their home ranges incomparison to females (Table 1). The averagepercentage of locations in the overlap area was39.3%, but the time spent within the area of overlap ranged widely and was the greatest be-tween F1 and M2, and minimum for the M2-F2dyad (Table 2). Although this difference was notsignificant (Mann-Whitney U -test: Z = 1.622, p= 0.106), on average, in the overlap areas be-tween individuals of different gender, we re-corded 45.3% of the total locations while this

proportion was only 27.5% for intrasexual pairs(Fig. 2, Table 2).Core area overlap was extensive between F1

and M2, while it was more reduced for the otherpairs. No overlap was recorded between females,as well as between F2 and M2 (Table 2).

To study dynamic interactions, we collected213 pairs of simultaneous locations, 54 of themof 2 animals, 33 of 3, and 10 including 4 individuals. For simultaneous locations, the averagedistance ( SD) between males and females was732.9 428.9 m. Females were closer (mean dis-tance between the two females: 541.1 143.7 m)than males (1265.9 579.3 m; Student t -test: t =8.2, df = 62, p = 0.001). The distance did not vary with the activity stage for any type of dyad(female-female: t = 0.64, df = 2, p = 0.29; male--male: t = 0.44, df = 6, p = 0.33; male-female t =0.37, df = 18, p = 0.36).

The average Coefficient of Spatial Associa-tion had a greater value in intersexual dyads


Table 1. Home range percent overlap matrix as obtained us-ing the 95% Kernel estimator for 4 C. chinga individuals inthe Pampas grassland of Argentina. F female, M male.

ID M1 M2 F1 F2

M1 31.8 34.1 42M2 26.7 46.8 18.9F1 55.8 91.2 41.9F2 67.7 36.2 41.2

Table 2. Matrix showing the percentages of locations in the overlap area between individual home ranges(HR), calculated with 95% K, and core areas (CA), calculated with 45% K, for 4 C. chinga individuals inthe Pampas grassland of Argentina. F female, M male.

IDM1 M2 F1 F2

HR CA HR CA HR CA HR CA

M1 31 8.8 32 1.2 47 3.9M2 29.9 10.3 53.7 38.6 7.5 0F1 43.9 3.1 86.6 84.4 17.1 0F2 55.3 8.9 36.2 0 31.9 0

10

20

30

40

50

60

70

L o c a t i o n s

i n t h e o v e r l a p a r e a s

( % )

M-M F-F M-F

Fig. 2. Average percentages ( SD) of locations in the over-lap areas of home ranges (95% Kernel estimator) for intraand intersexual dyads of C. chinga in the Pampas grasslandof Argentina. F female, M male.


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(0.14) than for dyads of the same gender (0.02between males and 0 between females, Table 3).The values of this coefficient were large for F1with both males, whereas all other possibledyads spent less than 4% of their locations atless than 300 m of distance.

Mostly, Jacobs index values were close to 0,indicating neutral spatial association. The mostnegative value was between males (0.23), whilethe index value of 0.57 for the M1-F1 dyad issuggestive of spatial association (Table 3).

Reciprocal distances between individuals var-ied over the time. All animals were relativelycloser in March and April, and then their recipro-cal distances increased, until reaching a peak inJune, when the highest average distance be-tween dyads was recorded. The distance betweenfemales seemed to be more constant than for theother dyads. It is interesting to observe that

while the distance in the M1-F1 dyad decreasedalong the study period, the distance between F1and M2 was gradually increasing (Fig. 3).

Discussion

In general, overlap between individual homeranges in the Molinas hog-nosed skunk in thePampas grassland was wide. This was true evenif we excluded the most external locations,which can possibly due to occasional excursionsthat increase the home range size of an animal,and, as a consequence, its overlap with other in-dividuals. Furthermore, these areas of overlapdid not represent marginal sections of individ-ual home ranges, since they were used in a rela-tively intense form (collared animals spent3848% of their time there). The most solitary


Table 3. Matrix showing the Coefficient of Spatial Association (CSA) and Jacobs index (Ji) values for thedynamic interactions between 4 C. chinga individuals in the Pampas grassland of Argentina. F: female;M: male.

IDF2 M1 M2

CSA Ji CSA Ji CSA Ji

M1 0.028 0.085 M2 0.036 0.033 0.019 0.227 F1 0 0.149 0.295 0.574 0.235 0.010

0

500

1000

1500

2000

March April May June July August

Month

D i s t a n c e ( m

)

F1-F2

M1-M2

F1-M1

F1-M2

F2-M1

F2-M2

Fig. 3. Monthly variations of the distances between 4 individuals of C. chinga , in the Pampas grassland of Argentina. F fe-male, M male.


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carnivore species have reduced intrasexual homerange overlap, in particular between females,whose reproductive success is mainly deter-mined by the access to trophic resources (eg,Macdonald 1983, Sandell 1989). This spatial or-ganization has been described as typical in theMustelidae family (Powell 1979). However, it isexpected that the home range overlap in a popu-lation of carnivores be also strongly affected bythe dispersion of the main resources that theyuse (Kruuk and Macdonald 1985, Fournier et al .2008). Molinas hog-nosed skunks feed mostly oninsects (Travaini et al . 1998, Donadio et al .2001), and this resource, being abundant and of

quick renovation, can favor a greater overlapamong individual home ranges (Johnson et al .2000).

Home range overlap in our sample of individ-uals was larger intersexually than intrasexually.This result, together with a bigger home rangesize in males, is in accordance with expectations,since mustelid males usually try to expand theirterritories to increase their chances of accessing to a greater number of females (Powell 1979).This explanation is also supported by the factthat the average overlap of male home rangeswith females was 3239%, a proportion thatwould enable each male to overlap its actionarea with three females.

Also the study of the percentage of locationsin the overlap areas among home ranges showedthat their importance was greater for thefemale-male dyads than for intrasexual dyads.The last aspect studied by static interactionswas core area overlap. As it can be expected forsolitary species, we found a null or very reducedoverlap in core areas for most radiotrackedskunk dyads. Nevertheless, we also observed acomparatively wide overlap between M2 and F1,and also a certain degree of overlap between thetwo males. The explanations of these two casesare probably different. While the intersexualoverlap of M2 and F1 could be due to theformation of a reproductive pair, for the otherdyad a possible explanation is that the overlaparea is inside or near the horse exclusion area, apatch where food was probably very abundantbecause grassland is in better conditions andsupported a greater richness and abundance of

arthropods (D. Castillo, unpubl.). A similarbehaviour was reported by Bixler and Gittleman(2000) for striped skunks Mephitis mephitis in acampground/picnic area where people fed them.However, we are aware that other possibleexplanations cannot be ruled out. In particular,our study did not include genetic analysis of theindividuals and therefore does not allow us tomake conclusions on the specific role of kinshipin the interactions we observed.

In any case, our results confirm that an an-alysis of home range overlap cannot be con-sidered complete if the dynamic interactionsamong individuals have not been studied,

because this is the information that we need inorder to understand whether the animals whoshare parts of their home ranges try to reducecontacts, or not. The Coefficient of Spatial As-sociation confirmed that, in general, Molinashog-nosed skunks of different sexes tended tospend more time at small distances than individuals of the same sex. On the other hand, itallowed detecting differences between F1, whospent a lot of time in proximity of both males,and F2, who was rarely together with themales. Jacobs Index is a further refinement of this analysis, and in our case, showed that F1was only spatially associated to M1 and not toM2, in contrast with what suggested by thestatic interaction analysis. In agreement withthis result, the mean distance between F1 andM1 was smaller than that recorded for the dyadF1-M2. This index also indicates that the twomales, in spite of their extensive overlap in homeranges, are unfrequently spatially associated.

Finally, it is interesting to notice that the av-erage distance between radiotagged skunkscould vary markedly over time. This kind of ob-servations should discourage researchers fromanalyzing and interpreting interindividual in-teractions as a static phenomenon.

Our sample size (in term of number of loca-tions, and, especially, of individuals tracked)prevents us from drawing definitive conclusionsabout the social organization of C. chinga . Thepresence of other individuals in the area used bythe four radiotracked animals, proved by cap -tures and direct observations, could also possi-bly change the picture obtained by this study.



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Even if these limitations cannot be overlooked,we can conclude that our data for C. chinga , sug-gesting stronger intrasexual than intersexualterritoriality and a greater tolerance between fe-males than males, are in agreement with thepattern of social organization described in theliterature as typical for mustelids and mephitids,particularly if the availability of their typicalfood sources is accounted for. However, they alsoconfirmed that it is difficult to reduce the com-plexity and flexibility in the strategies for re-source use to fixed schemes (Johnson et al .2000), especially when individuals are used assampling units. Finally, we suggest that the

analysis of dynamic interactions is clearly a fun-damental tool to improve our understanding of social behavior since (as in our case) they can po-tentially give a different view of this complex as-pect of vertebrate ecology than that provided bystatic interaction studies and that more effortshould be devoted at developing standardize pro-tocols of study of dynamic interactions enabling direct comparisons among studies and species.

Acknowledgements: We thank C. Manfredi, S. Savini, H.Bindotti, J. Baglioni, R .Temperoni, H. Marrero, P. Pereyraand M. Rodriguez for their help in captures and radiotelem-

etry data collection. Thanks are extended to the rest of GECM members for their advices. The staff of TornquistProvincial Park provided logistic support. JIR, ML and ELV have been supported by a postgraduate scholarship fromConsejo Nacional de Investigaciones Cientficas y Tcnicas(CONICET), and DFC by postgraduate scholarships fromComisin de Investigaciones Cientficas of Buenos AiresProvince (CIC) and CONICET. The paper was supported byEarthwatch Institute, IM40 Grant by ANPCyT, and Sec-retara General de Ciencia y Tecnologa, UNS (PGI 24 B/123).

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Received 26 February 2008, accepted 29 September 2008.

Associate editor was Joseph F. Merritt.


Documents

Interindividual interactions of Molina’s hog-nosed skunks Conepatus chinga in the Pampas grassland of Argentina, --home range overlap, dynamic interactions, static interactions,