Lizard Podarcis (=Lacerta)

SHORT NOTES

SHORT NOTES In common with most of such high energy boulder/ pebble beaches there was little sign of animal lifesave

HERPETOLOGICAL JOURNAL, Vol. 5, pp. 200-203 (1995) for substantial numbers of an isopod, Ligia italica Fabricius. L. italica is a large (<16 mm body length in MELANISM AND FORAGING females; Vandel, 1960) plant and debris-eating isopod and browses particularly (in daylight hours) upon li­ BEHAVIOUR IN AN INTERTIDAL chens, microbial films and, on large boulders and fixed POPULATION OF THE MADEIRAN rocks, upon green macroalgae (especially C/adophora LIZARD PODARCIS (=LACER TA) sp.). Lizards that were black dorsally and blue-grey DUGESII (MILNE-EDWARDS, 1829) ventrally were seen in the intertidal zone during most daytime visits to the beach at Canii;:o, thoughsig htings were rare beforemid-morning. They occurred fromthe JOHN DAVENPORT* AND THOMAS DELLINGER** top of the shore down to about low water neap tide level though they were never seen nearer to the water's edge *University Marine Biological Station, Millport, Is le of than about 1-2 m. Cumbrae, Scotland, KA28OEG Melanism. To determine whether the 'intertidal' popu­ lation differed in colour and morphology from the **Universidade da Madeira, Largo do Colegio, P-9000, 'terrestrial' population, 115 lizards (58 'intertidal'; 57 Funchal, Portugal, Madeira 'terrestrial') were collected (between 15.11. 93 and 16.2.94) in fruit-baitedbucket traps set in the intertidal The Madeiran lizard Podarcis (=Lacerta) dugesii zone and nearby terrestrial area (pyroclastic pebbles (Milne-Edwards) is a common lacertid 'wall' lizard and earth, with some vegetation). Measurements were occurring throughout the Madeiran archipelago, par­ made to the nearest mm: snout-vent length (SVL), tail ticularly in the orchards and vineyards of the main length (TL; only for specimens with undamaged island where it occurs in large numbers and has been tails.); and to the nearest 0.1 mm: head width (HW), regarded as an agricultural pest for centuries by the right forearm length (RFAL) and length of 4th digit of Madeirans (Forster, 1777). P. dugesii is omnivorous, right forelimb (FIL; 'finger length'). The colour of liz­ eating vegetation, fruit and even nectar as well as in­ ards was ranked on a scale from 0-5 (5 being sects and other terrestrial invertebrates (Elvers, 1977, completely black, 0 being sand-coloured). This study 1978; Sadek, 1981; Beyhl, 1990). confirmed that intertidal lizards were almost black P. dugesii is a polymorphic, adaptable species that (mean score = 4.4, SD = 0.5, n=58), while terrestrial occurs in a variety of morphological types and colours lizards were dark brown (mean score = 3.1, SD = 0.6, (Cook, 1979; Crisp et al. , 1979) and is foundat all alti­ n=57); the difference was statistically highly signifi­ tudes in Madeira, from sea level where air cant (Mann-Whitney U-test: U = 2293, P<0.001 ). temperatures range between about l 8°C in winter and Intertidal lizards were also significantly bigger (mean 32°C in the summer, to mountainous areas up to 1850 SVL=58.0 mm SD=7.0, n=58) than terrestrial lizards m where summer temperatures may be as low as 20°C (mean SVL= 51.8 mm, SD = 9.4, n=57; Mann­ and snow falls inthe winter. Whitney U-test: U = 2293, P<0.001). There was a The investigation reported here was carried out on a weak, but significant, influenceof body size on colour melanic population of P. dugesii that was seen to be rank (colour score = 0.05 SVL + 1.16; r2 = 0.24, exploiting the intertidal zone, an unusual ecological P<0.001 ). However, ANCOVA established that, for niche particularly given the high-energy (i.e. exposed the same body length, intertidal lizards were always to heavy wave action), stony nature of Madeiran darker than terrestrial animals (P<0.001 ). To correct shores. for body size, slopes and elevations of the regressions The study was carried out at a beach at Canii;:o, ofTL, HW, RFAL and FIL on SVL were compared for Southern Madeira, about 15 km east of Funchal. The both lizard groups. No differences in slopes were de­ whole beach, about 1 km long, consists of small out­ tected (ANCOVA, homogeneity of slopes) for any of crops of dark coloured lava interspersed with long the measurements. However, for a given SVL, inter­ stretches of mixed dark grey boulders, cobbles and peb­ tidal lizards had longer tails and longer fingers than bles. These latter portions of the beach were regularly terrestrial lizards as there were significant diffe rences resorted by the advancing and receding tides since in elevations for TL (ANCOVA: F = 11.6, n = 105, Madeira features strong breezes and heavy oceanic P<0.00 1) and FIL (ANCOVA: F = 18.9, n = 113, swells. The jumble of boulders, cobbles and pebbles P<0.001). The regression equations are listed in Ta­ was generally at least 0.3 m thick and overlaid coarse ble 1. gravel. Although the uppermost layers of this substra­ These data show that the intertidal population dif­ tum rapidly dried out on sunny days, the deeper layers fers significantly in both colour and morphology from were always damp. Most of the dampness was due to its terrestrial near-neighbours. The diffe rences in mor­ seawater, but there were some freshwater inputs to the phology might, perhaps, have been explained by a beach too. difference in sex ratio, since Sadek (1981) reported SHORT NOTES 201

TABLE I. Regression equations of tail length (TL), head The single animal dissected was a mature male width (HW), right forearm length (RFAL) and finger length (overall length 134 mm; snout-to-vent length 61 mm). (FIL) on snout-vent length (SVL) for intertidal and On dissection the stomach contents were foundto con­ terrestrial lizards or, if ANCOV A stated no diffe rence sist wholly of five large, partly digested Ligia. The between regressions, for pooled data. isopods were in pieces; all of the heads were separate from the bodies. The droppings of the three other cap­ tured lizards revealed the following material: Variable Slope Constant r1 p Lizard I: Parts of the exoskeleton of a large cockroach (attracted into the banana-baited trap?), several parts TL(intertidal) 1.41 13.04 0.49 <0.001 of Ligia exoskeleton, particularly the eyes, plus a small TL (terrestrial) 1.62 9.09 0.64 <0.001 piece of polythene sheet. Lizard 2: Droppings contained Ligia exoskeleton HW (pooled) 0.08 0.98 0.58 <0.001 pieces, two dipteran pupal cases, and parts of a large cockroach. RFAL(pooled) 0.20 -0.44 0.67 <0.001 Lizard 3: Beetle elytrae and large numbers of Ligia FIL(intertidal) 0.10 0.56 0.50 <0.001 eyes. The three animals held captive after trapping took FIL(terre strial) 0. 10 0.21 0.83 <0.001 specimens of Ligia offered to them immediately after collection of droppings (i.e. when they had fasted for 24 hr). They were subsequently held for several that females have relatively shorter tails and hind legs months during which they were fed exclusively on a than males. However, although the lizards of this study diet of Ligia and talitrid amphipods. were not sexed directly, the head width data were ana­ In conclusion, the observations recorded in this lysed (c.f. Sadek, op. cit.) and this analysis showed that study indicate that there is a separate population of there was no significant difference in sex ratio between melanic P. dugesii that forages entirely or predomi­ the populations, reinforcing the hypothesis that the in­ nantly between the tidemarks at Caniyo, Madeira, the tertidal population of lizards is morphologically most important item of diet being the large, amphibi­ distinct. ous isopod L. italica. The population appears to be Foraging behaviour. Specimens of P. dugesii were readily separable from nearby terrestrial populations regularly seen along the whole of the beach; direct by virtue of its black coloration, though there are subtle visual observation of their behaviour was carried out morphological differences too. Although Sadek ( 1981) whenever possible over a period of two weeks in the included 'beach' populations in his study of the diet of summer of 1993. Four melanic specimens of P. dugesii P. dugesii on Madeira, he makes no referenceto mela­ were trapped in a banana-baited trap at mid-tide level. nic lizards on the main island of Madeira, and it seems The banana bait was wrapped in perforated plastic film likely that he caught supralittoral/terrestrial predators so that it could be smelt, but not eaten. One of the ani­ rather than intertidal foragers, particularly as the diet mals was sacrificed humanely for dissection and of his 'beach' specimens was dominated by ants, bee­ analysis of stomach contents; the others were held tles, flies and amphipods (which may well have been separately unfedfor 24 hr so that their droppings could terrestrial talitrids, common on Madeira (Dahl, be collected. These three animals were then offered 1967)). Both Crisp et al. ( 1979) and Bischoff et al. specimens of L. italica and their behaviour was ob­ ( 1989) also fail to mention melanic lizards on the served. main island of Madeira. On several occasions intertidal lizards were seen to Other carnivorous lizard species are known to for­ eat discarded fruit remnants (particularly banana age entirely within the intertidal zones of islands: skins) leftbehind by tourists. However, they were also Ano/is agazzi and Diploglossus haycocki which both seen to exhibit alertness whenever specimens of Ligia eat intertidal crustaceans on the Isla Malpelo, Colom­ were visible nearby. Often they were seen to stalk bia, and the skink Cryptoblepharus boutoni which eats isopods, but usually disappeared from view in the jum­ insects, crustaceans and juvenile fish on Madagascar ble of beach pebbles before food capture was observed. (Fricke, 1970), migrating up and down the shore in However, on the afternoon of 17.8.93 two black lizards similar fashion to P. dugesii. Although no other were seen to capture and eat Ligia. One caught a single lacertid lizard has been reported to use the intertidal isopod (capture and hand ling taking 5 s ), but the other zone in this fashion, such behaviour might be expected first chased two large groups (n>20) of isopods without from other wall lizards living on islands (e.g. in the success, ceasing pursuit within 0.2-0.4 m. It then eastern Mediterranean) where L. italica is to be found. chased, caught and swallowed a large isopod (catching The followingfactors may have encouraged the de­ and handling time I 0 s), rested for 60 s beforejumping velopment of intertidal foraging on Madeira. On most in the air to catch a dipteran fly. It ate another Ligia Atlantic coastlines the productivity of cobble/pebble before disappearing into the pebble substratum in pur­ beaches is dominated by strandlines at the top of the suit of another isopod. shore where rotting wrack supports a diverse talitrid 202 SHORT NOTES

amphipod and insect (dipterans and beetles) fauna. moregulatory advantage; Crisp et al. ( 1979) have al­ Such strandlines are less important on a volcanic is­ ready provided good experimental evidence dismissing land like Madeira whose seaweed beds are very limited this possibility forthe melanic P. dugesii living on the in extent. Instead, the dominant animals are large L. Desertas islands. The differences in size and propor­ italica that feed on green algae on boulders and in tions exhibited by the intertidal population of P. pools, but also eat detritus, lichens and microbial films dugesii are more difficult to understand, although the as they move up and down with the tide. On most east­ Galapagos marine iguana (also melanic) has similarly ern north Atlantic shores a different member of the well developed toes and claws for maintaining pur­ genus is found, Ligia oceanica L. (see Vandel, 1960 chase on a slippery substratum (Bell, 1825; Trillmich for discussion). However, L. oceanica is a nocturnal & Trillmich, 1986). species (Nicholls, I 93 I) that hides under stones Acknowledgements. The authors acknowledge the as­ throughout the day, whereas L. italica (at least on Ma­ sistance of travel fundingprovided to J. Davenport by deira) is a day-active isopod that hides at night when the British Council and the Junta Nacional de predatory grapsid crabs (mainly Pachygrapsus Investigacao Cientifica e Technologica in Lisbon. maurus) are numerous and active on the wet rocks. They are also grateful to Dr D. Holdich of the Univer­ Lizards are diurnal visual foragers, too slow-moving in sity of Nottingham for help with identification of the cool of the night to catch fast-moving animals, so isopods. the daytime coincidence of P. dugesii and its isopod prey is important. Additionally, in most coastal inter­ REFERENCES tidal environments there is an alternation ofpredatory influences with fish and crabs dominating when the Bannerman, D.A. & Bannerman, M. (1965). Birds of the tide is in, and sea birds dominating when the tide is Atlantic islands, Va l 2: A history of the birds of out, at least during the day. On Madeira the influence Madeira, the Desertas and the Porto Santo Islands. of birds on the intertidal zone is very limited since the Edinburgh & London: Oliver & Boyd. number of resident seabirds is low (e.g. Jepson & Bell, T. ( 1825). A new genus of Iguanidae. Zoological Zonfrillo I 988), there are no members of the crow fam­ Journal 2, 204-208. ily on Madeira (Bannerman & Bannerman, 1965) and Beyhi, F. E. ( 1990). Observations on the flower-visiting even migrant waders are sparse (Zonfrillo, pers. lizards of the island of Madeira, Portugal (Reptilia: comm.). Both P. dugesii and its prey avoid predation Sauria: Lacertidae). Courier Forschung-Institut at high tide and/or night by moving offthe shore or Senckenberg 129, 153-158. hiding amongst the pebbles of the substratum; they can Bischoff, W., Osenegg, K. & Mayer, W. (1989). both exploit the intertidal zone during the day with lit­ Untersuchungen zur subspezifischen Gliederung der tle or none of the risk that they would face during the Madeira-Mauereidechse, Podarcis dugesii (Milne­ day ifthere were more wading birds, or if corvids were Edwards, 1829). Salamandra 25, 237-259. present. Cook, L. M. ( 1979). Variation in the Madeiran lizard The reliance of a population of P. dugesii on a diet Lacer/a dugesii. Journal of Zoology, London 187 of intertidal isopods poses a number of interesting 327-340. questions regarding salt and water regulation. No data Crisp, M., Cook, L. M. & Hereward, F. V. (1979). Color concerning the osmoregulatory capacity of L. italica and heat balance in the lizard Lacerta dugesii. Copeia appear to have been published, but the related L. 1979, 250-258. oceanica, L. occidentalis and L. pallasii are all power­ Dahl, E. ( 1967). Land amphipods from the Azores and ful osmoregulators at low salinities and are actually Madeira. Boletim do Museu Municipal do Fun cha I hyperosmotic to full sea water (Parry, 1953; Wilson, 21, 8-23. 1970). This suggests that the body fluids of the isopods Elvers, I. ( 1977). Flower-visiting lizards on Madeira. will have osmolarities of at least 1000-1100 m Botaniska Notiser 130, 23 1-234. Osmoles kg-1, some 2-3 times the likely plasma osmo­ Elvers, I. ( 1978). The Madeiran lizard-flower connection larity (probably ea 350-400 m Osmoles kg-1) of the observed in a natural habitat. Botaniska Notiser lizards. Although the ready availability of Ligia will 131, 159-1 60. provide abundant foodfor intertidal P. dugesii, the liz­ Forster, G. (1777). Voyage round the world in His ards must incur a considerable salt load when Brittanie Majesty's sloop Resolution, commanded by exploiting this resource. Whether the lizards cope with Capt. James Cook, during the years 1772, 3,4 and 5. this salt loading behaviourally by seeking out fresh­ In Georg Forsters We rke, Vol l: A voyage round the water for drinking, or physiologically by extra-renal world. (ed. G. Steiner). Berlin: Akademie-Verlag. salt secretion is as yet unknown. Fricke, H. W. ( 1970). Die okologische Spezialisierung The melanic nature of the intertidal P. dugesii is der Eidesche Cryptoblepharus boutoni cognatus undoubtedly cryptic, given the unrelieved black/grey (Boettger) auf das Leben in der Gezeitenzone colour of the rocks and stones of the Madeiran inter­ (Reptilia, Skinkidae). Oeco/ogia (B erlin) 5, 380-391. tidal zone. The small body size of the species makes it Jepson, P.R. & Zonfrillo, B. (1988). Bird notes from unlikely that the black colour provides any ther- Madeira, summer 1986. Bocagiana, 117 1-10. SHORT NOTES 203

Nicholls ( 1931). Studies on Ligia oceanica. I. A. Habitat Vandel, A. ( 1960). Ligia italica Fabricius, 1798. pp. 122- and effect of change of environment on 127 In Faune de France vol. 64. isopodes terrestres respiration. B. Observations on moulting and (p remiere partie) . Liechtenstein: Kraus. breeding. Journal of the Marine Biological Wilson, W .J. ( 1970). Osmoregulatory capabi Ii ties in Association of the United Kingdom 17, 655-674. isopods: Ligia occidentalis and Ligia pallasii. Parry, G. (1953). Osmotic and ionic regulation in the Biological Bulletin. Woods Hole, Massachusetts, 138, isopod crustacean Ligia oceanica (L. ). Journal of 96- 108. Experimental Biology 30, 567-574. Sadek, R. A. (1981). The diet of the Madeiran lizard Lacer/a dugesii. Zoological Journal of the Linnean Society 73, 313-341. Trillmich, K. G. K. & Trillmich, F. (1986). Foraging strategies of the marine iguana Amblyrhynchus cristatus. Behavioural Ecology & Sociobiology 18, 259-266. Accepted: 9. 7. 94