The structure of a mediterranean lizard community⁽¹⁾

Doñana, Ac a Ve eb a a, 2 (2): 145-160, 1975 !he s ue u e o a Medi e aaean U.a d eommuni y Cl) J. MELLADO, F. AMORES, F. F. PARREÑO AND F. HIRALDO 145 The s uc u e o liza d communi ies has ecen ly been he objec oí nu- me ou$ s udies in which cha ac e is ic p ope ies o animal communi ies, such as sha ing o esou ces, niche b ead h and o e lap, ' as well as s a e- gies u ilized by he di e en species o educe compe i ion, ha e be en ana- lyzed. Mos o hese s udies (Rand, 1964; Schoene , 1968; Schoene and Go man , 1968; Hillman, 1969; Pianka, 196~a; Schoene , 1970; Huey e al., 1974:) ha o been made on g oups o sympa ic oongene ic speci'es, whe eas Pianka (1967; 1969b; 1971; 1973) and mo e ecen ly Sage (1974) analyzed en i e communi- ías o liza ds. He e We analyze he s uc u e o a liza d communi y ep esen a i e o ha Eu opean Medi e anean zone, an a ea as ye uns udied excep o a p &- cu so y in es iga ion ca ied ou by Val e de (967). We also compa e his communi y o hose desc ibed by Hu ubia and Di Cas i (1973) and Saga (1973) o he Medi e anean zones o he New Wo ld. Ma e ial and Me hods 610 specimens belonging ' o eigh specíes o liza ds we e collec ed a i egu- la in e als hl'ougholl a pe iod oi 35 mon hs (Oc obe1' 1971 o Augus 1974). These samples we e cap u ed p incipally by shoo lng 01' by hand and p ese cd in 70 pe cen Alcohol, excep iol' mos Lace- a. lepid4 whicb we e dona ed by he Ownel'S o p í a e es a es whe e hey a e usually killed, being conside ed as ha m- ul o hun ing; hese samples we e p ese ed in 10 pe cen o ma Un. Tlle buck o all ma e ial used he1'e has been dona ed o he Biological S a- ion o Doñana, Se illa, Spain. (1): This papel' was o iginally deli e ed in a Symposillm on he «Ecosys em s uc u e and s abili y" in No embe 1974 a Se illa, Spain. 146 J. MELLADO, F. AMORES, F. F. PARREÑO AND F. HIRALDO Time be ween cap u e and analysis a ied g ea ly, om immedia ely a e - wa ds ill se e al mon hs la e o Samples collec ed p o ided he ollowing mo phological da a (measu ed on he p ese ed animaD : SVL (snou en leng h) : Measu ed om he end o he snou o he dis- al edge o he anal pla eo ea . HL ( Head lengh ): F om he end o he snou o he an e io bo de o he HW (Head wid hl: Maximu n wid h o he head . W (Weigh ): Only in esh specimens. Al! neasu emen s a e in millime e s, weigh s in g ams . Only s o m ac h con en s we e conside ed in he s udy o ood. P ey i ems we e coun e d Ol" each s om a ch and clasi ied in o 21 ood c a eg o i es lis ed in able II. The l eng h o in ac p ey was me asu ed o he ne a es .1 o a milli- me e l·. Fi e ld ob se a íon s we e m ade a i egula in e als, a leas once a mon h (exc ep Sep e mh e ), a nd du l ng a minimum p e l od o i e days ea ch m on h, a di e e n ime s o day an d nigh . W hen poss ibl e, he ollo win g d a a we e no ed in ie ld obs e a i o ns : species , ap o x íma e size a nd age , ime , locall y and cx ac loca ion as we ll as ac i i y wh en i s sigh ed. Sludy A es Field obse a ionsand ma e ial employed in his s udy come om an ex ensi e a ea si ua ed a he mos wes e n pa o he Sie a Mo ena moun ain ange in he P o ince o Huel a (S. W. Spa n, ap oxima ely 38° N by 7° W). This zone comp ises lines o moun ainous coun y o oeunded p o- ile, a low al i ude (a e age ele a ion o 500 me e s abo e sea le eD, 'which al e na es wi h wide alleys and peniplanes ( o a de ailed desc ip ion o his zone, see Lau ensach, 1967). The clima e is o he ype «Mesomedi e-- mnéen accen ué» as desc ibed by Embe ge e al. (1963) wi h 75 o 100 ' bio- logi'cally d y days. Annual p ecipa ion a ies om abou 800 o 1,300 mm. o ain which alls mainly om No embe o Ma ch . Tempe a u es below ze o, a e in equen , and he o al numbe o hou s pe yea bel ow ze o does no e a ch he h ee pe cen indica ed by Asch- mann (1973) as a limi o Medi e I' anean clima es. F os ls sca ce and snow excep ional on his land, as any snow all usually mel s immedia ely. WHh ega d o ege a ion, h is o iginally co nsis ed in a dense e e g een scle ophylous o es o Que cus ilex and/o Q. sube wi h so ne isola ed s ems o Q. lusi anica in dampe a eas . This Q. es possessed a dense unde . b ush made up chie ly o Ulex spp. and se e al species o Cis us as well as A bu us unedo. In he b ooks and go ges galle ies o Alnus glu inosa and F axinus sp. occu wi h hick bushes o Rubus and Ne ium oleande . This ege a ion has been adically ans o med by he hand o man in wide ZQ_ Dañan :>., Ac a Ve eb a a, 2 (2), 1975 The $ uc u e 01 a Medi e anean liza d communi y 147 nes, and due o degene a ion o he o .es , wo main ypes o habi a ha e appea ed: Chapa al.-This consis s basically o Cís us ladanije us, E ica aus al s and E. umbella a, oge be wi b Genis a sp., A. unedo and E. a bo ea appea- ing in damp a eas. This unde b ush o ms a e y dense con inuous blan· ke oí up o ou me e s bigh in some places, and gene ally occupie·s high a eas o moun ain anges. E e g een oak wood.-A hin o es o Q. ilex and/o Q. sube in which be soíl has been comple ely clea ed o unde b ush. This habi a occupies he alleys and plains. Ex ensi e pa k~ oí Cas anea sa i a oí simila s uc· u e a-ppea in he coole and dampe a eas. T o hese mus be added a e y peculia o ma ion consis ing oí ou · c ops o g ani ic ocks de e mining ue so·called «Plu onic bu onsll (see Lau ensach, 1967). These a- e ex emely ocky a e as wi h some s ems o ooks , and unde b ush o Pis ada sp ., Mi us sp . and Cis us . ResuU. A maximum o eigh liza d species can co·exis in he egian s udied. These a e (2): Tm Ta en ola m. mau i aniea Be Blanus e. cine eus Lh Lace a hispanica auche i Ll Lace a l. lepida Pa Psammod omus a. algi us Ph Psammod omus h. hispanicus Cb Chalcides bed iagai pis aciae Ce Chalcides chalcides s ia us Gekkonidae Amphisbenidae Lace ida.e )) )) )) )) )) )) Scincidae » » In he es o his pape , he symbols p eceding each liza d name lis ed aboye will be used. Table 1 shows he main biome ic cha ac e is ics o he di e en spe- cies, and igu e 1 p esen s he dis ibu ion o snou ' en leng hs (SVL). Th oo pai s o congene ic speci'es occu in sympa y; hese di e om one ano he wio hin each pai , in aH measu emen s aken in o conside a ion. (2): Following Me e ns and We mu h, 1960, excep o Chalcides bed iagai (see Val e de, 1966). Doñana, A. c a Ve eb a a, 2 (2), 1975 148 J. MELLADO, F. AMORES, F. F. PARREÑO AND F. HIRALDO All hese di e ences a e s a is ica11y signi ican (P< .05; - es ) excep be ween he wo species o he genus Chalcides whe e none o he di e en- ces we e s aUs ica11y signi ican . Table 1 Mo phological s a is cs used in bis s udy SVL W HW HL m c n m c n m c n m c n Tm 54·04 14.1 21 6.6 4.4 15 9.66 2.4 18 15.0 3.6 18 Be 160.05 30.3 37 4.6 1·7 15 4.82 1.9 28 7.8 Vi 28 Lh 46.U3 4.7 131 2.0 1.0 45 5.87 1.0 56 10.3 1.8 .'l9 Ll 138.46 53.7 167 71.7 8.4 35 15.9 9.8 73 32.8 4.3 74 Pa 58.88 14.6 141 7.7 2.4 65 10.6 1.9 39 15.9 2.2 39 Ph 40.05 5.S 57 1.1 1.1 42 5.54 4.0 34 9.8 1.4 35 Cb 66.14 10.6 21 4.4 1.2 11 6.71 1.8 21 8·0 .9 21 Ce 95.08 31.0 22 4.9 3·8 21 5.90 1.2 22 8.5 1.7 19 SVL = Snou - en leng h; HW = Head wid h; HL = Head leng h; W = Weigh . Food A o al o 511 s omachs we e analyzed. Table 2 summa izes he main ea u es o he die o he eigh liza d species in eg a ed in he communi y. Mos liza d species a e ecclec ic p eda o s, eeding on a wide ~ange o in e eb a e p ey (see e.g. Pianka, 1973, and included e e ences). In e- quency o appea ance o p ey üems in s omachs analyzed and equency oí each p ey i em in he o al, he mos abW1dan g oups in he die we e bee- Ies, spide s and .insec 18i ae,amonga wide a ie y o g· oups o in e e- b a es. Seeds 8ind e eb a e p ey a e also included in he die o se e al species. Table 3 shows he p ey size dis ibu ion and he mean p ey size o each liza d species. Liza d size i:; gene ally co ela ed wi h ha o he p ey inges ed. In his case, mean p ey size was col1 ela ed bo h wi h snou - en leng h and he8id dimensions o he liza d o 8111 species. O hese compa í- sons, only mean p ey size s. mean head wid h was s a is ically signi ican ( : .720; P< .05; n: 8). Doiíana, Ae a Ve eb a a, 2 (2), 1975 The s uc u e 01 a Medi eTTanean liza a communl 'll 1.9 • Be Ll Ce - Cb ~ Pa Tm Lh 40 Ph o 20 100 , .o . 180 220 mm. 60 Fig. 1. F equency dis ibu ions o snou - en leng hs (millime e s) lo he eigh liza d species comp'lsing he communi y . Doñana, Ac a Ve eb a a, 2 (2), 1975 150 J. MELLADO, F. AMORES, F. F. PARREÑO AND F. HIRALDO T ophic di e si y was calcula ed by Pielou's me hod (1966), using he B iUouin o mula: H=(1/N) (logIa Ni! -I loglO Ni!) This me hod was o me ly applied o he analysis o die o p eda o s by Hu ubia (1973). Plo ing accumula ed ophic di ' e si y (Hk) s. o al numbe o s omachs (k), di e si y cu es a e ob ained (Figu e 2). These cu o es a e highe o la ge species, in pai s o congene s. Duo o sample he e ogenei y, including s omachs om e y di e en pe- iods, locali ies and habi a s, i is di icul o de e mine he s abiliza ion poin ( ) o he cu es, and so he di e si y o a11 he s omachs (Hz) o each Table 2 Pe cen age compm:i ion, by numbe s, o he die o Uza d species composing he communi y The numbe oC s omachs and p ey i ems on which hese pe cell ages a e based, as well as he a e age ophic di e si y (Hz), a e al so shown. Oligoehae a Isopoda Gas e opoda Miliapoda A ane ae halangidae Seo pionidae O hop el'a De map e a 'BJa la ia Hemyp e a Fo micidae O he hymenop e a Lepidop c a Dip e a Coleople a Lepidop e a Ia ae O he Ia ae Seeds Ve eb a e i ems lTniden i ied P eys i ems S omaehs T ophic di e si y (Hz) Tm 2.85 2. 85 20.0 1.42 10.0 7.14 1.42 8.57 1.42 10.0 17.13 8.57 4.28 2.85 70 18 .975 Be 4.9 .98 2.94 12.74 .98 Lh 3.24 .36 1.79 24.46 1.08 2·16 .72 1.08 2.16 Ll .47 1.12 .42 1.26 0.5 ·08 1.77 .28 .56 16.66 9.35 1.85 3.95 2,4 .03 7. 19 . 98 9.80 20.87 68.42 l7.64 9.71 5.0 28.40 6.47 9.36 2.94 103 23 .760 1.5 .36 .25 5.09 .90 268 2919 9S 158 1.059 1.087 Doñana, Ac a Ve eb a a, 2 (2), 1975 Pa Ph Cb l.19 9.4 22.5 .39 2.5 1. 38 2.3 5.0 7,14 24.8 20.0 .39 8.53 .19 .19 1.98 12.60 10.11 2. 97 38.07 8.92 3.76 1.1 .5 15.3 1.1 2.3 23.9 5.9 8.8 7.5 35.0 7.5 Ce 10.3 2·06 18.55 6.18 8.25 3.10 1.03 4.12 24.73 10.jO 7.21 3.10 .19 1.78 504 3.5 169 48 40 79 132 1.225 .852 15 22 .707 .960 The s uc u e 01 a Medi e anean liza d communi y 151 species, is used as indica o o ophic di e si y; alues ob ained appea in able II. A s a is ically signi ican cOI1 ela ion exis s be ween ophic di e - si y and head wid h o he liza d species ( s: .643; P<.05; n: 8. Spea man ank co ela ion coe .icien . Siegel, 1956). Finally, wi h espec o hun ing s a egy, Pianka 0966; see also 1973) Table 3 T ophic esou ce ma ix. Rows ep esen s p ey size i ems a 5 mm. in e alo To al numbe o p ey measu ed (N), and mean p ey size (MPS) a e also shown. Tm Be Lh Ll Pa Ph Cb Ce O- S 9 15 64 10 39 60 11 16 6- 10 23 47 51 660 63 54 15 28 11- 15 7 2 25 389 34 9 2 15 16- 20 3 4 7 255 33 2 4 21- 25 1 4 55 7 3 26- 30 ~ 5 1 66 11 31- 35 5 23 36- 40 1 10 4 41- 45 1 1 3 46- 50 4 51- 55 1 56 · 60 1 61- 65 3 66- 70 2 71- 75 1 76- 80 1 81- 85 86- 90 91· 95 1 96-100 2 N 46 76 158 1486 197 125 30 68 MPS 10.06 11.35 8.72 13.72 12.9 6.0 7.16 9.72 -de ines wo basic ones: «si -and-wai » and «ac i e o aging». In ou case only one species (Tm) is a «si -and-wai )), while he emainde seem o be <mc i e o aging» hun e s. .. ¡ Doñana, Ae a Ve eb a a, 2 (2), 1975 152 J. MELLADO, F. AMORES, F. F. PARREÑO AND F. HIRALDO Ac i i y The p esence oí a well-de ined wi . e la eney has been a ibu ed as a undamen al cha ae e is ie o he auna o ep iles o he Tempe a e zone (Sain -Gi ons, 1956). Sain -Gi ons (1953) a ibu es wo diapauses o he Me' di e anean zones, he one in win e and a summe la ency, due o he ex· Hk 11 .9 7 , , .5 ..- . I -' / .. .... '" ........ ', ..... Ce Cb Ll Pa Ph .3 .1 o 5 10 15 5 10 15 5 15 20 K Fig. 2. Accumula e ophic di e si y (Hk) cu es s . o al numb o s omachs (k). Compa ison be ween congene ic pai s oi species. eesi e d yness. Da a o e ed by ou sel es in his espee , al hough ineo n- pIe e o so ne species, do howe e , pe ni us o d aw a en ion o wo ypes oí beha iou . Tablc 4 Spa ial esou ce ma ix. Symbols heading he able Indica es mic ohabUa ypea d awn in de ail in he ex o A B e 2 3 1 2 3 Tm o 22 o o o o o Be o 20 o o 17 o Lh () 40 o o 6 o o Ll 27 24 o o 3 o o Pa o o o 141 3 o o Ph 32 o o o o o (j Cb o o o 1 o 21 o Co o o o o o o 22 Doñana, Ac a Ve eb a a, 2 (2), 1975 The s uc u e 01 a Medi e anean liza d communi ll 153 A leas wo species (Ll and Pa) show a ma ked diapause, wi h an inac- i e pe iod o some ou mon hs (No embe o Feb ua y), al hough he smaH- e membe s o hese species a e ac i e h oughou he Whole yea . In o he species (Tm and Lh), bo h adul s and young a e ac i e aH yea . As a as dai1y ac i i y is concemed, only one species (Tm) is noc u - nal, while he emainde a e essen ially diu nal. Howe e , his pa e n oí ac i i y is subjec o app eciable changes h oughou he yea , wi h he noc- u Il8l species becoming diu nal du ing win e mon hs , whe eas some diu - na! species (Ll and Pa) ha e o en been obse ed a nigh du ing ho e mon hs (Mellado and Hi aldo, unpublished). Mie ohabi a p ele enee AH place loca ions obseJ' ed o liza ds when i s sigh ed we e classi ied in o se en weH-di e en ia ed ca ego ies o de ine he spa ial eS<Ju ce sha.- ing o he communi y. We conside only obse a ions e e ing o eeding beha iou o he mo egula o y ac i i ies. These ca ego ies include: A Zones o sca ce o no plan co e . B Zones o dense plan co e . Wi hin each o he abo e-inen ioned ypes he oHowing di isions a e made: 1 On he g ound. 2 On ocks, ees and buildings. 3 Unde s ones, ee unks, e c. In adi ion o hese ca ego ies, ano he one mus be added . I is: e In small hick pas u es in alleys and pJains. One o he species (Cc) is es ic ed o C; mo phologically i is a snake- like liza d which U e ally «swimsll in he pas u es i inhabi s . The es ol he species a e limi ed o a special kind o subs a um: h ee species (Ll, Pa and Ph) li e on he g ound su ace, one o hem (Pa) in places we e he e íS a hick unde b ush, whe eas he o he wo (Ll and Ph) a e ound on open g ound (Ll needs aes and ocks o p o ec ion). Be is a ypical sub e a- nean species, mo phologicallya wo m-like liza d. Cb li es unde s ones in hls egion. Finally, Lh and Tm li e on ocks and l ee unks. Doñana, Ac a Ve eb a a, 2 (2), 1975 160 J. MELLADO, F. AMORES, F. F. PARREÑO AND F. HIRALDO I.&VINS, R. (1968): E olu ion in changing en iTonmen s. P ince on Uni . P ess: 120 pp. MERTENS, R, and WERMUTII, H. (1960): Die Amphibien uoo Rep ilien EUTOpa& Ve lag Waldema K ame , F ank u a. M. PIANKA, E. R (1966): Con exi y, dese liza os and spa ial he e ogenei y. Ecolog¡¡ 47: 1.055-1.059. - - (1967): On liza d species di e si y: No h Ame ican la land dese s. Eco· logy 48: 333-351. - - (1969a): Habi a speci ici y, specia ion and species densi y in Aus alian de- se liza ds. Ecology 50: 498-502. ~ -(1969b): Sympa y o dese liza ds (C eno us) in Wes e n Aus alia. Eco- logy 50: 1.012-1.020. - - (1971): Liza d spacies densi y in he Kalaha i dese o Ecology 52: 1.024-1.029. - - (1973): The s uc u e o liza d co nmuni ies. Ann. Re . Ecol. Sys . 4: 53-74- - - (1974): Niche o e lap and di use compe i ion. P oc. Na . Acad. Sci. USA 71: 2.141-2.145. PIELOU, E. C. (1966): The measu emen 01 di e si y in di e en ypes o biolo- gical collec ions. J. Theo e . Biol. 13: 131-144- RAND, A. S. (1964): Elcological dis ibu ion in anoline liza ds o Pue o Rico. Eco- logy 45: 745-752. SAGE, R D. (1973): Ecological con e gence 01 he liza d aunas o he chapa al communi ies in Chile and Cali o nia, p. 339-348. In: Medi e an!lan ype ecosys em. F. Di Cas i and H. A. Mooney eds., Sp inge -Ve lag, Be lin- Heildelbe g-New Yo k. - - (1974): The es uc u e o liza d 1auna : compa a i e biologies o liza ds in wo A gen ina dese s. Ph. D. hesis, Uni . o Texas a Aus in. Diss. Abs . 35,1. SAINT-GIRONS, H. (1953): No e su les pe iodes de la ence des ep iles au Ma oc. Bull. Soco Zool. F ance 78: 377-381. - - (1954): Cycle d'ac i i é e he mo egula ion chez les ep iles <leza ds e se pen s). Vie e MiHeu 7: 133-226. SCHOENER, T. W. (1968): The Anolis liza d o Bimini: esou ce pa i ioning in a complex auna. Ecology 49: 704-726. - - aud GORMAN, G. C. (1968): So ne nicha di e ences among h ee species o Lesse An illeau anoles. Ecology 49: 819-830. SCHOENER, T. W., and GoRMAN, G. C. (1970): Nonsynch onous spa ial o e lag o li- za ds in pa chy habi a s. Ecology 51: 408-4118. - - and SCHOENER, A. (1971a): S uc u al habi a s o Wes Indian Anolis li- za ds I. Lowland Jamaica. B e io a 368: 1-53. - - - - (1971b): S uc u al habi a s o Wes Indian Anolis liza ds n. Pue o Rican uplands. Ibid. 375: 1-39. - - (1974): Resou ce pa i ioning in ecological communi ies. Science 185: 27- 39. SIEGEL, S. (1956): Nonpa ame ic s a is ics OT he beha io al sciences. McG aw Hill Book Company, Inc., New Yo k-To on o-London. VALVERDE, J. A. (1966): Sob e las sub especies de Chalcides bed iagai (Basca). Bol. R. Soco Española His . Na . (BioU, 64: 169-170. - - (1967): Es uc u a de una comunidad de e eb ados e es es. C. S. . C. Mad id. J. MELLADO, F. AMoRES, F. PARRKÑO and F. HIRALDO Cen o Biológico del Su Reina Me cedes, 17. Se illa-12 ESPAÑA (SPAIN). :Doiíana, Ac a Ve eb a a, 2 (2), 1975