Habitat Use by Red Deer and Fallow Deer in Doñana National Park

HABITAT USE BY RED DEER AND FALLOW DEER IN DOÑANA NATIONAL PARK

Braza, F. & Álvarez, F., 1987. Habitat use by Red Deer and Fallow Deer in Doñana National Park. Misc. Zool., 11: 363-367.

Habitar use by Red Deer and Fallow Deer in Doñana National Park.- On the Doñana Reserve (Southern Spain) the Fallow Deer tends to occupy the marsh and its periphery, whereas the Red Deer is found in al1 different habitat types, showing a slight preference for the shrub and dunes areas. The Fallow Deer does not vary to a great extent its habitat preferences across seasons, whereas Red Deer moves towards the more humid areas during the summer and beginding o€autumn, a time at which habitat overlap between both species reaches its highest level, and towards the drier areas during the rest of the year. The observed patterns of habitat use are probably re- lated to the grazing habits of the Fallow Deer and the browsing tendencies of Red Deer in the study area.

Key words: Cervus elaphus, Dama dama, Red Deer, Fallow Deer, Habitat use.

(Rebur: 13-1-87)

F. Braza & F. Álvarez, Estación Biológica de ~oñana,Ap. 1056, Sevilla, España.

INTRODUCTION STUDY AREA AND METHODS

Knowledge relating habitat use and in- Habitat units terspecific competition in southern Iberia ungulates is sparse (ROGERS & MYERS, Within the National Park of Doñana least 1980). disturbed area, that is the inner Reserve, The matter is of special importance in an covering a surface of approximately 7500 ha area such as the National Park of Doñana, was chosen. For the purposes of the study, where high ungulate densities and the ab- the following environmental units (ALLIER sence of predators urgently demands popula- et al., 1974) were considered (fig. 1): tion control as a measure of habitat conser- Fixed dunes: These are the driest part of vation. the stabilized sand formations, covering an Indirect techniques for the estimation inactive old dune system where the crests are of ungulates habitat use and population occupied by sparse wood of Juniperus densities have been succesful used in a vari- phoenicea or scarce shrub with maximum ety of arcas (see NEFF, 1968 for review; cover of Cistus libanotis, and the interdune DZIECIOLOWSKI, 1973), thcir use being valleys by shrub with maximum cover of especially important for nocturna1 or secre- Halimium halimifolium and abundance of tive species. Cistus libanotis, Lavandula stoeehas, Ar- In thc present study the occupancy of dif- meria pungens, Helichrysum angustifolium ferent vegetation communities on the Park and Stauracanthus genistoides, as well as by Red Deer (Cervus elaphus L.) and Fallow planted Pinus pinea. Deer (Dama dama L.) is compared. Moving dunes: This constitutes a fraction f I xed dunes

shrub

Fig. 1. Environmental units in the Doñana Reserve and location of the transects. Unidades ambientales en la Reserva de Doñana y localización de los transectos.

of the extensive active dunes system with era ang groups of Pinus pinea and Quercus fronts parallel to the coastline. The most fre- suber. quent elements of the very sparse vegetation Marsh: Very flat area flooded in winter. are Corema alba, Armeria pungens and Am- Cover of Scirpus maritimus and Scirpus mophila arenaria. lacustris. Fine alluvial soil. Shrub: An extensive area, slightly lower, Marsh periphery: This narrow border be- and located to the east of the previous tween the marsh and the shrub areas consists habitat types. Over the sandy substrate, the of dry pastures of Tuberaria guttata, predominant vegetation are the shrubs Trifolium campestre, Plantago coronopus, Halimium halimifolium, Stauracanthus Cynodon dactylon, etc., in the nearest zone genistoides, Calluna vulgaris, etc. in high to the shrub and series of complex pasture areas, and of spots of Erica ciliaris, Ulex mosaics, with Trifolium fragijerum, Cyno- minor and Erianthus ravanae in the depres- don dactylon, Paspalum vaginatum and sions, as well as pasturelands of Agrostis Gaudinia fragilis in the transition to the stolonifera, Illecebrum verticillatum, Anagal- marsh, as well as extensions of Pteridium lis crassifolia, Senecio prealtus, etc., derived aquilinum and Juncus spp., stands of Quer- from the previous shrub. cus suber and isolated specimens of Populus Ponds: Located along the moving dunes alba. system parallel to the coastline, including more or less permanent ponds, scrub of Ulex minor and Erica ciliaris, extensions of Data collection and analysis Echinodorus ranunculoides and Eleocharis multicaulis, pasturelands of Agrostis stolonif- A total of 22 tra'nsects 500 m long by 2 m Table 1. Total percentage of presence (proportion of segments per transect containing either pellet groups or . footprints) in the different vegetation units. Po~etirajeioial de presencia (proporción de segmentos por iransecio que conleniun grupos de excrementos o lruellus de I>~.SLI~US)etr las dijerenies unidades umbieniules.

Environmental Units Number of transects Red Deer Fallow Deer Fixed dunes Moving dunes Ponds Shrub Marsh periphery Marsh

wide were cstablished. The number of trans- pancy appear quite different for the two deer ccts in the vegetation units were proportional species: comparisons of the index of pre- to their extcntion (see table 1). Each transect sence between the two resulted in statisti- was marked by a straight line of numbered cally significant differences in each of the wooden pegs in its middle line, so the whole plant communities considered (P < 0.001, transect was divided into 25 segments of 20 m Chi squared test). long each. Some degree of seasonal variability is ap- The transects were established on 30th parent in the habitat preferences of the Red June 1982, and animal signs (faeces, foot- Deer (fig. 2): the animals tend to move to the prints) wcre recorded at one monthly inter- more humid areas (the marsh and its vals. Signs were removed from the transects periphery) during the summer and beginning in each visit. of autumn (comparison of the periods June- An index of presence was calculated as the October vs. November-May: marsh, x2 = proportion of the 20 m long transect seg- 81.71, df = 1, p < 0.001; ponds, x2 = 26.94, ments per habitat unit per day containing df = 1, p < 0.001), and towards the drier signs left by the deer. areas (mainly the fixed dunes) during the rest Niche overlap for each month of the study of the year (comparisons of the periods June- was calculated applying PIANKA'SIndex October vs. November-May, x2 = 86.54, df (1973) to the indices of presence on the dif- = 1, p < 0,001). Although to a lesser degree, ferent environmental units. the Red Deer also shows a greater tendency towards using the shrub area during the more humid periods (x2 = 6.05, df = 1, p < 0.02) RESULTS as well as to occupy the marsh periphery to a higher degree during the drier of the two The overall values of the index of presence periods (x2 = 9.28, df = 1, p < 0.01). (table 1) show that the Red Deer extends The Fallow Deer, on the other hand, rather uniformly over al1 the areas of the re- maintains its presence without much vari- serve, intensity of use being slightly higher ation in the marsh and its periphery al1 year for the shrub and dunes areas. On the other round. hand, the Fallow Deer is found more often in As to the monthly variation in PIANKA's the marsh and its periphery than over the (1973) overlap index between the two rest of the environmental units; its presence species, figure 3 shows that it reaches its in the shrub and moving dunes is low and is higher values during the summer months, never found at al1 at the fixed dunes area. probably as a result of the Red Deer moving As a whole, the patterns of habitat occu- to the more humid areas as that time. Misc. Zool. 11, 1987

red deer

C fallow deer

" utilization of the different vegetation units. Porcentaje de utiliza- ción mensual de las unidades de vegetación.

habits: the varied diet of the Red Deer in other areas.(DZIECIOLOWSKI, 1967, 1970a, 1970 b, 1970c; KAY& STAINES,1981) and its more intense browsing tendencies over the shrub vegetation and more varied diet, as compared to the Fallow Deer in the study area (PALACIOSet al., 1980; VENERO, 20 L JASONDJFMAMJ 1982), allows the Red Deer to extend to more varied habitats as well as to approach or withdraw seasonally from more different Fig. 3. Monthly values of Pianka's niche overlap in- areas, whereas the less eclectic Fallow Deer dex. concentrates al1 year round on grazing on the Valores medios mensuales del índice de Pianka de meadows of the marsh periphery and on the marsh itself, the meadow areas bordering the ponds being the next more prefered habitat. According to the patterns of movements DISCUSSION detected for both species in the ecotone area of the Doñana Reserve (BRAZA et al., The differences detected in habitat occu- 1984), the occurrence of the Red Deer dur- pancy and in degree of seasonality seem to ing the summer and beginning of autumn in correspond to specific differences in feeding the marsh and its periphery would presumably be the cause of the higher habitat overlap ACKNOWLEDGEMENTS found between both species at that time, when diet overlap is also greatest (VENERO, We are grateful to al1 the companions and friends 1982). who helped us during data collection at Doñana as In fact, the pattern of seasonal habitat pre- well as to Prof. R. Dzieciolowski for reviewing an ference in the Red Deer, presumably con- earlier draft of the manuscript. trolled by feeding needs, is probably a direct response to the seasonally fluctuating depth of the water table, itself affecting the occur- REFERENCES rence of various plant communities (ALLIER et al., 1974). As a responde to the deeper water table during the summer, the Red ALLIER, C., GONZÁLEZBERKÁLDEZ, F. & Deer would be able to move to lower places, RAM~REZD~Az, L., 1974. Mapa ecológicol feeding to a large extent on the grass of the Ecological map. Reserva Biológica de Doñana. marsh and its periphery at that time and re- Estación Biológica de Doñana, C.S.I.C. Sevilla. BATCHELER,C.L., 1960. A study of the relations be- maining also in those traditional rutting areas tween roe, red and fallow deer, with special refer- for the first part of the autumn. ente to Drummond Hill Forest, Scotland. J. Besides the highly probable effect of food Anim. Eco!. ,29: 375-384. preferences we cannot discard microclimatic BRAZA,F., ALVAREZ,F., GELDOF,R. & BYLOO, H., 1984. Desplazamientos de ungulados silvestres conditions having an effect on thermal com- a través de una zona de ecotono en Doñana. fort of deer, in this way affecting the in- Doñana, Acta Vert., 11 : 275-287. terspecific and seasonal differences detected D~IECIOLOWSKI,R. M., 1967. Food of the red deer in habitat occupancy. in an annual cycle. Acta Theriol., 12: 503-520. Whereas differential habitat selection was - 1970a. Foods of the red deer as determined by rumen contents analysis. Acta Theriol., 15: 89-110. algo suggested by BATCHELER(1960) for - 1970b. Food selectivity in the red deer towards Red Deer and Fallow Deer in forests in Scot- twigs of trees, shrubs and dwarf shrubs. Acta land, the results of this study for the 1982-83 Theriol., 15: 361-365. period (just after a period of severe draught) - 1970c. Variation in red deer food selection in re- lation to environment. Ecologia Polska, 18: 635- agree to a great extent with those of ROGERS 645. & MYERS(1980) of 1973 (when precipitation - 1973. The use of pellet-group counts to census red was normal), although during the 1982-83 deer (Cervus elaphus). In: Proc. XI Intern. Congr. period both Red and Fallow Deer apparently Game Biologists: 559-563. Stockholm. extended to more varied environmental KAY,R.N.B. & STAINES,B. W., 1981. The nutriiion of the red deer (Cervus elaphus). Nutrition units, both towards more dry and more wet Abstracts and Reviews, Ser. B, 51: 601-622. areas than their favoured ones. NEFF, D.J., 1968. The pellet-group count technique for big game trend, census, and distribution: A review. J. Wildl. Manage., 32: 597-614. RESUMEN PALACIOS,F., MARTINEZ,T. & GARZ~N,P., 1980. Ecología alimenticia comparada del Ciervo (Cer- Uso del habitatpor el ciervo y el gamo en el Parque Na- vus elaphus hispanicus. Hilzheimer, 1909) y el cional de Doñana.- En la Reserva de Doñana, el Gamo (Dama dama, Linne, 1758) en el Parque gamo tiende a ocupar la marisma y su periferia, mien- Nacional de Doñana. In: Actas II Reun. tras que el ciervo se encuentra en todos los habitats Iberoamer. Zool. y Cons. Vert.:444-454. Cáceres. mostrando una ligera preferencia por las áreas de ma- PIANKA,E. R., 1973. The structure of lizard com- torral y de dunas. Las preferencias del gamo no varian munities. Ann. Rev. Ecol. and System., 4: 53-74. demasiado a lo largo de las estaciones, mientras que el ROGERS,P. M. & MYERS,K., 1980. Animal distri- ciervo, va hacia áreas más húmedas en verano y a butions, landscape classification and wildlife man- principios de otoño Cpoca durante la coincidencia en agement, Coto Doñana, Spain. J. Appl. Ecol., 17: el habitat de ambas especies es máxima y hacia las 545-565. áreas más secas durante el resto del año. El uso del ha- VENERO,J. L., 1982. Dieta de los grandes fitófagos bitat está relacionado con el modo de alimentación de silvestres del Parque Nacional de Doñana, Es- ambas especies. paña. Ph.D. Thesis, University of Seville.