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FEEDING ECOLOGY of VIPERA LA TASTEI in INTRODUCTJON Foraging Ecology Plays a Direct Or Indirect Role in Vir Tually All Aspect

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FEEDING ECOLOGY of VIPERA LA TASTEI in INTRODUCTJON Foraging Ecology Plays a Direct Or Indirect Role in Vir Tually All Aspect HERPETOLOGICAL JOURNAL, Vol. 14, pp. 13-19 (2004) FEEDING ECOLOGY OF VI PERA LA TA STEI IN NORTHERN PORTUGAL: ONTOGENETIC SHIFTS, PREY SIZE AND SEASONAL VARIATIONS JOSE C. BRITO Centro de Biologia Ambiental and Dep. de Biologia Animal, Faculdade de Ciencias da Universidade de Lisboa, Lisbon, Portugal The diet of Vipera latastei was investigated in northern Portugal from 1998 to 2002. Palpation of stomach contents and forced defaecation from 190 snake specimens resulted in the recovery of 83 identifiableprey items. V. latastei preys on fo ur species of small mammal (76%), two lizard species (14%), three amphibian species (5%) and arthropods s.l. (5%). Estimates of prey availability demonstrated that the most common prey were also the most frequent prey consumed. No differences between the sexes were detected in terms of the proportion of snakes with prey or diet composition. However, there was an ontogenetic shift in diet composition. Juveniles fed mostly on ecthotermic prey (60%), the maj ority of subadults fed on insectivorous mammals and lizards (60%), and adults fed mainly on rodents (88%). This ontogenetic shift is mostly due to the morphological constraints imposed on the juveniles, which cannot swallow large prey items. There is a positive correlation between snake size and prey size. V. latastei is selective in terms of both the species and size of prey ingested, with larger snakes being more selective than smaller snakes. Larger snakes have a narrower food niche breadth than smaller snakes, but their diet composition overlaps 1i1oderately. There is seasonal variation in the diet composition, with snakes taking amphibians mainly in spring and autu mn, lizards in spring, and mammals in summer and autumn. Feeding fr equencies indicate that both males and females - and subadults and adults - consume prey more frequently during summer. Key words: dietary habits, food selection, prey availability, snakes, Viperidae INTRODUCTJON Despite its former wide distribution in P01iugal, it pres­ ently only occurs in isolated and small populations, Foraging ecology plays a direct or indirect role in vir­ especially in the south (Godinho et al., 1999). This spe­ tually all aspects of snake life-history (Mushinsky, cies is threatened mainly by habitat loss, as a 1987). For instance, obtaining sufficientenergy is a cru­ consequence of forest fires and intensification of agri­ cial factor in reproduction and it may influence the culture, and direct human persecution (Brito al., frequency of reproduction (Naulleau & Bonnet, 1996), et 200 l ) . Additionally, the Portuguese preliminary con­ reproductive output (Gregory & Skebo, 1998) or the ma­ servation status of "Indete1minate" for V. latastei ternal condition after reproduction (Shine & Madsen, (S.N.P.R.C.N., 1991), demonstrates insufficient I 997). Feeding also affects demographic traits of knowledge concerning tltis species. Thus, a conserva­ populations, especially rates of growth and survival tion programme has been established in n01ihern (Madsen & Shine, 2000). Portugal, which aims to identifypriority areas for con­ Although the diet of western European vipers(V ipera servation and to develop management strategies for spp.) is relatively well documented (Bea et al., 1992), viper populations and their habitats. data on Lataste's viper (Vip era latastei) are scarce and Data on foraging ecology is important because mostly anecdotal (Valverde, 1967; Yericad & Escarre ( l) species with very specialized diets usually are unable to 1976; Lopez Jurado & Caballero, 1981 ), or come from studies carried out in captivity outside their natural dis­ use alternative prey types if their ha bitats are modified or destroyed; and (2) ontogenetic shifts in diet compo­ tribution area (Saint-Girons, 1980). Although general sition may contribute to the endange1ment of species, accounts (Bea & Brana, 1988) state that this species for instance due to declines in a specific prey type feedson lizards and small mammals, no study has spe­ by a particular age/size class of snakes (Webb cifically addressed its foraging ecology. needed & Shine, 1998). Within the framework of the conserva­ Vipera latastei is a small-sized viper, with snout-vent tion programme for latastei, the following questions length usually not larger than 60.0 cm. It is distributed V. were asked: (I) is it a size-selective predator and how fromnorth-west ern Africa (Morocco, Tunisia and Alge­ does prey availability influence the diet composition? ria) in the south, to the Iberian peninsula in the north, (2) with the exception of Cantabrian and Pyrenean regions. Are there ontogenetic shiftsin the diet composition? (3) Are there intersexual differences in the diet compo­ sition? ( 4) What is the frequency of feeding and are Correspondence: J. Brito, Centro de Investigai;:ao em there seasonal changes in diet composition? These Biodiversidade e Recursos Geneticos da Universidade do Porto (CIBIO/UP), Instituto de Ciencias Agrarias de Vairao, questions should help in understanding the foraging R. Padre Armando Quintas, 4485-661 Vairao, Portugal. E­ ecology of V. latastei and allow the identificationof fa­ mail: [email protected] vourable ecological niches for the species. 14 J. C. BRITO METHODS The availability of potential prey was estimated in a smaller area (2.24 ha), and compared with prey data STUDY AREA from thevipers capturedin this area (64% of the total ex­ The study was conducted in the extreme north-west­ amined snakes). Due to logistical reasons, prey ern part of the distribution area of V latastei (Brito & availability fo cuses only on small mammals and lizards. Crespo, 2002), in a National Woodland and Biosphere Small manu11al abundance was estimated using l 00 Reserve, "Mata de Albergaria ", Portugal ( 4 l 049'N, Shen11a1m traps, placed in line at 10 111 intervals, baited 8°07'W; elevation c. 750 m a.s.1.), included in a pro­ and surveyed for six nights, in June 1999, 2000 and tected area, Parque Nacional da Peneda-Geres. The area 2001 . This period should conespond to ilie annual abun­ is a series of granite mountains, witl1 altitudes ranging dance peak of small manmials in this region (Mathias, from 50 to 1500 m a.s.I. The climate is Atlantic Medi­ 1999). Traps were examined in early morning (at about terranean (Goday, 1953), characterized by wet winters 07:00 to 08:00 hr) and late afternoon (at about 19:00 to and hot summers. Mean monthly rainfall ranges from55 20:00 hr). Lizard abundance was estimated through mmin July to 457 nun in January (average annual rain­ visual encounter surveys of 1 OOO m, in the same area at fall above 3200 mm/year), and mean air temperature the same time period. Tluee lacertid lizard species were ranges from 7. 9 °C in January to 2 0 .4 °C in July (aver­ foW1d in this smaller study area (Podarcis bocagei, age annual air temperatme 14 °C) (INAG, 2002). The Lacerta schreiberi and L. lepida), but - due to the large vegetation consists mostly of deciduous oak forests size of adult L. schreiberi and L. lepida - only juveniles (Quercus robur) or mixed deciduous and coniferous were considered as potential prey for V latastei. forests (Pinus sp.) with arbutus trees (Arbutus unedo). The extent of sexual divergence in viper body mor­ Major shrubs include heath ( Calluna vulgaris and Erica phology was quantified according to the methods sp.), gorse (Ulex sp.) and brooms (Cytisus sp.). outlined by Shine (1991 a). For SVL, the quotient be­ tween fe male SVL and male SVL was found. For HL, FIELD AND ANALYTICAL METHODS HW and HH, a regression equation linking female meas­ Data were collected between 1998 and 2002 in an ures to the SVL was calculated. An estimate of the area of around 2.300 ha. 135 snakes were caught by female headmeasure was obtainedusing the mean ma le band, sexed by analysing external tail morphology, SVL. The ratio of female to male head length, at a mean measured for snout-vent length (SVL), head length male SVL, provided an index oftJ1e extent of sexual size (HL), bead width (HD), and head height (HH), and per­ dimorph ism in relative head measmes (Shine, 1991 a). manently marked for future identificationby clipping For analysis purposes, snakes were divided into tluee unique combinations of ventral scales. Snakes were pal­ size categories according to tl1eir sexual maturation sta­ pated to detect prey in the stomach and intestines, but tus: (1) Juveniles and neonates (SVL <30.0 cm); (2) only forced to defecate intestine contents and never subadults (SVL 30.0-40.0 cm); (3) adults (SVL >4 0.0 forcedto disgorge prey; however, prey presence was cm) (Brito & Rebelo, 2003). Diffe rences in diet compo­ recorded in such cases. Additionally, 55 road-killed sition were analysed by x2 tests and contingency tables snakes were dissected to collect stomach and intestine (Siegel & Castellan, 1988). To avoid pseudoreplication prey items. Mammals were identified using identifica­ of data, no snake was included in the analysis more ilian tion keys based on the internal structure of hairs once. Niche breadth was measured using the standard­ (Teerink, 1991 ). Reptiles, amphibians and invertebrates ized Levin's index (BA) and niche overlap between were identifiedthrough their externalmorphology. Prey ea tegories was measured using Pianka' s symmetrica I items collected from road-killed snakes were measured equation (Ojk) (for details see Ksebs, 1989). The relation (body length), whenever their preservation status al­ between snake size and prey size was detennined by cal­ lowed. culating Kendall's tau (Siegel & Castellan, 1988), TABLE I. Sample sizes, body measures and sexual size dimorphism of Vipera latastei containing prey. SVL , snout-vent length,; HL, head length; HW, head width; HH, head height (all measures in cm); SD, standard deviation; SSD, sexual size dimorphism. See methods for calculation of SSD. -------- Mean Mean Mean Mean 11 SVL±SD HL±SD HW±SD HH±SD Males 62 36.3±9.1 2.02±0.36 1.38±0.26 0.76±0.13 Feniales 39 37.7±8.3 2.12±0.37 1.40±0.30 0.79±0.
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