1 1 Scietas Eurpaea Lepidpterlgica; dwnlad unter http://www.bidiversitylibrary.rg/ und www.zbdat.at Nta lepid. 23 (2): 147-172; 01.VII.2000 ISSN 0342-7536 n the ethlgy and eclgy f a small and islated ppulatin f the Dusky Large Blue Butterfly Glaucpsyche (Maculinea) nausithus (Lycaenidae) Manfred Alban Pfeifer. Ulf Jsef Settele" Rbert Andrick 2. Wlfgang Frey 3 & Bahnhfplatz 5, D-67240 Bbenheim-Rxheim. Germany : Mannheimer Str. 12. D-67665 Kaiserslautern. Germany 3 Universität Kaiserslautern, Fachgebiet Wasserbau und Wasserwirtschaft. D- 67663 Kaiserslautern. Germany 4 UFZ - Centre fr Envirnmental Research Leipzig-Halle. Department f Cnservatin Bilgy and Natural Resurces. Permserstraße 15. D-04318 Leipzig. Germany (crrespnding authr: e-mail: settele@pr.ufz.de) Summary. During a study f a small, relatively islated ppulatin f Glaucpsyche (Maculinea) nausithus (Bergsträsser. 1779). nearly every new adult entering the ppulatin was individually marked in summer 1990. Marked individuals were regularly recaptured. The ppulatin assessment was nearly cmplete, as shwn by a cmparisn with estimates based n the Jlly-Seber-mdel. The average adult residence time was estimated at 2.3 days. Individual bservatins were made f the diurnal distributin f the fllwing activities: resting, flight, nectaring. vipsitin. cpulatin, and [ther] mvements. ur results shw sex and age specific differences. n the day f marking, which was nrmally the day f eclsin, all activities were cmparatively lw. lder individuals f bth sexes were generally active fr the same amunt f time per day. but differed with regard t activity prfiles. Males undertk lnger flights t search fr females within the habitat, while the females flew mre ften but ver much shrter distances, resulting in a shrter ttal flight time. Instead, females stayed n the flwer buds f Sanguisrba fficinalis the nly larval hst plant and main adult nectar surce fr much lnger, and laid eggs. Females cpulate directly after hatching and then immediately start t vipsit. Zusammenfassung. Eine kleine, islierte Ppulatin vn Glaucpsyche (Maculinea) nausithus (Bergsträsser. 1779) wurde im Smmer 1990 nahezu vllständig erfaßt. Aufgrund der intensiv angewendeten Fang-Wiederfang-Methde wurden im Habitat Tiere mit hher Wahrscheinlichkeit am Tag des Schlüpfens markiert. Immigratin knnte weitgehend ausgeschlssen werden. Dadurch kennen wir mit grßer Sicherheit das Alter der Tiere, an denen ethlgische Messungen durchgeführt wurden. Vn Individuen dieser Ppulatin liegen zeitliche Messungen der Aktivitäten Ruhen. Fliegen. Saugen. Eierlegen. Kpulieren und Bewegen (Sammelgruppe für weitere Aktivitäten) im Tagesgang vr. 147
Unsere Ergebnisse zeigen, wie sich die Aktivitäten geschlechtsspezifisch unterscheiden und wie sie sich im Verlauf des Falterlebens verändern. Am Tag der Markierung, der in den allermeisten Fällen mit dem Tag des Schlüpfens zusammenfiel, waren die Aktivitäten vergleichsweise gering. Beide Geschlechter verbrachten ungefähr gleich viel Zeit aktiv, allerdings mit deutlich verschiedenen Schwerpunkten. Während die Männchen längere Suchflüge nach Weibchen im Habitat unternahmen, waren die Flüge der Weibchen zwar zahlenmäßig häufiger, insgesamt verbrachten sie aber deutlich weniger Zeit fliegend. Dafür bewegten sich die Weibchen länger auf den Blütenköpfen der Saug- und Eiablagepflanze Sanguisrba fficinalis und legten Eier. Weibchen kpulierten unmittelbar nach dem Schlüpfen und begannen direkt danach mit der Eiablage. Résumé. À l'ccasin de l'étude, l'été 1990, d'une petite ppulatin relativement islée de Glaucpsyche (Maculinea) nausithus (Bergsträsser, 1779), presque chaque individu nuveau entrant dans la ppulatin a été marqué individuellement, puis régulièrement recapturé. L'évaluatin de la ppulatin était presque cmplète, cmme l'a mntré la cmparaisn avec le mdèle de Jlly-Seber. La myenne de la durée de séjur des adultes est de 2.3 jurs. L'bservatin des individus est basée sur la répartitin des activités diurnes: reps, vl, alimentatin, cpulatin, vipsitin et autres. Ns résultats mntrent des différences seln le sexe et l'âge. Le jur du marquage, qui est nrmalement celui de l'émergence, tutes les activités snt cmparativement réduites. Les vieux individus des deux sexes snt généralement actifs pur la même durée chaque jur, mais diffèrent par leurs prfils d'activités. Les mâles passent de lngues pérides en vl à la recherche des femelles dans le milieu, alrs que les femelles vlent mins suvent et sur des distances plus réduites, résultant dnc en une durée de vl ttal plus curte pur celles-ci. À la place, les femelles se repsent sur les inflrescences de Sanguisrba fficinalis la seule plante-hôte des chenilles et la principale surce de nectar pur des durées plus lngues et dépsent leurs œufs. Les femelles cpulent immédiatement après l'émergence et cmmencent la pnte des œufs juste après. Key wrds: Lepidptera, Lycaenidae, Glaucpsyche {Maculinea) nausithus, diurnal behaviur, age dependent behaviur, eth-chrngramme, ppulatin structure, dispersal, residence time, MRR-study, Rhineland-Palatinate, S. Germany. 1. Intrductin The eclgy f the Dusky Large Blue Glaucpsyche {Maculinea) nausithus (Bergsträsser, 1779) has been quite well studied. This is due mainly t its high level f vulnerability as expressed in Red Lists (fr Germany e.g. Pretscher, 1998) and the annexes f the Habitat Directive f the EU and in the Bern Cnventin (Gruttke, 1996; van Helsdingen et a/., 1997), and als t its fascinating Maculinea-type eclgy. The present study aims t imprve knwledge n the adult ethlgy and ppulatin eclgy f the species with special respect 148
t small and islated ppulatins. An example f such a ppulatin was studied in 1990 thrughut the flight perid. As the individuals f G nausithus fly slwly cmpared t ther butterfly species, they are easily fllwed. The species may reach high densities in its habitats, and is clsely assciated with its principal nectar surce and larval hst plant Sanguisrba fficinalis. Fr a mark-releaserecapture study, it is therefre sufficient t search fr patches with flwering plants. 2. Materials and methds 2.1. The species Glaucpsyche nausithus. G. nausithus is univltine. Eggs are exclusively depsited n Sanguisrba fficinalis, where the first three larval instars develp. Hereafter the caterpillar reaches the grund t be adpted by ants f Mynnica rubra L., the nly knwn hst ant f the butterfly. Within the ant nest the larvae live predaciusly and later pupate. n average, 2.5 pupae are fund in ne nest. This and further recent infrmatin n the eclgy f the species are t be fund e.g. in SBN (1991), Elfferich (1998), Thmas et al. (1998), Thmas & Elmes (1998) and Wynhff (1998). 2.2. Study sites. The study was cnducted n a ca. 2000 irr fallw grassland within the Mswieser Tal (Mswieser Valley, site M hereafter) suth f the city f Kaiserslautern, between the settlements Aschbacherhf and Weiherfelderhf at an elevatin f 300 m abve sea level. The site was characterised by a high density stand f Sanguisrba fficinalis, unlike the meadws lcated bth t the East and t the West f the site. The nrthern and suthern limits f the site are dense pine frests f the Pfälzerwald (Palatine Frest). Thus, the area f suitable habitat was relatively islated. The nly pssible immigratin f G nausithus frm its immediate surrundings was frm the margins f meadws lying west f the study site, where sme S. fficinalis plants flwered during the flight perid and where sme individuals f G nausithus were fund. N hst plant was fund t the East f the site. This relatively islated site was chsen fr the study because it seemed pssible t mark a high percentage f all individuals at the day f emergence with the manpwer available and t identify their age fr the behaviural analysis. 149
The site was divided int tw parts. The nrthern sub-area f the Aschbach (a little creek) was drier with Sanguisrba fficinalis dminating the vegetatin, while the wetter sub-area suth f the Aschbach held high densities f Lythrum salicaria L. and sedges, but nly ccasinal expsed S. fficinalis plants. An additinal site Hagelgrund (in the Eselsbachtal, site H hereafter), nrth f the city f Kaiserslautern, was als studied. This was used mainly t test ur methd f using small prtable cmputers (see 2.5). In sme cases we later used data btained frm Hagelgrund t enlarge the basis f ur study (see Tab. 1 belw). There the first G. nausithus culd be bserved n 13 July, ne day befre the first male n site M. As the next bservatins at site M have nly been made frm 17 July nwards, we made sme mre studies at site H until that day. 2.3. Marking. Adult butterflies were caught at rest with ur fingers. A waterprf pen was then used t write numbers n the underside f bth hindwings s that the individual culd be easily recgnised while fllwing it. The site in the Mswieser Tal was visited almst daily during the flight perid (see Fig. 1), and all unmarked individuals encuntered were marked and previusly marked specimens were recrded. The S. fficinalis stands f the mre western meadw margins were visited less ften. All newly encuntered individuals were als marked and we searched fr marked adults which might have emigrated frm ur main study site. 2.4. Estimatin f ppulatin size. ne way t quantify the size f a ppulatin is t sum the minimum number alive (MNA). As the name implies, this gives the minimum number f animals that are present in the ppulatin at ne particular time. T reach that number, the number f marked individuals registered (r marked) in the particular time step (e.g. day) and the number f individuals marked befre that time step and recaptured again after the time step, but nt registered at the time step itself, are added. Individuals that have left the ppulatin in the meantime, as well as thse which were verlked at the time step and were nt re-sighted later (because they have died, emigrated r been verlked repeatedly) d nt cntribute t the MNA f a particular time step. The ttal number f individuals present (fr a particular time step, 150
like ne day r the whle time f a species activity, e.g. 5 weeks) must therefre be at least as high as this minimal number alive, but nrmally is much higher. As the number directly depends upn the mark recapture effrts invested and the size f the ppulatin, its applicatin is mre apprpriate in small ppulatins f animals with a cmparatively high recapture rate, but it cannt be reliably used t cmpare different ppulatins (Settele et ah, 2000). In rder t cmpare ppulatin sizes, the Jlly-Seber methd might be suitable t get a first rugh idea n daily as well as ttal ppulatin sizes (see e.g. Seber, 1982, and Pllck et a/., 1990, fr details). Fr the latter ne may e.g. sum up the 5-values (i.e. the individuals added t the ppulatin between tw ppulatin estimates) resulting frm the Jlly-Seber mdel (see Settele et ah, 2000 fr an example and further details with respect t butterflies). Hwever, ne has t keep in mind, that Jlly-Seber estimates are nt very rbust with respect t ppulatin size. Table 1. rigin f additinal data fr the diurnal eth-chrngrammes (Figs 3, 4), which served as a basis fr the cmputing f the age dependent eth-chrngramme (Fig. 5) (M - Mswieser Tal (Aschbachtal), H - Hagelgrund (Eselsbachtal)). Cmpleted classes rigin f data Age class Perid 66 99 befre 10:50 6 6 1-2, befre 10:50, M 9 9 1-2, befre 10:50, M 10:50-11:40 6 6 0, 10:50-11:40, M+H 9 9 0, 10:50-11:40, M+H 11:40-12:30 6 6 0, 11:40-12:30, M+H 15:50-16:40 6 6 0, 15:50-16:40, M+H 3-10 15:50-16:40 9 9 1-2, 15:50-16:40, M 16:40-17:30 9 9 1-2, 16:40-17:30, M after 17:30 9 9 1-2, after 17:30, M Strictly spken, nly fr the age class 1-2 days a cmplete diurnal eth-chrngramme culd be cnstructed. Thus, in sme cases additinal data have been used frm the site Hagelgrund t cnstruct an age dependent eth-chrngramme ut f the diurnal ethchrngramme. Data f the class 1-2 day ld butterflies Have been used, if still the pled data frm bth sites (M and H) have nt been sufficient. There have been n data f bth sexes f age class fr the daily time befre 10:50 h. Therefre data f 1-2 day ld butterflies have been used. Fr the perid 10:50 h - 11:40 h f the day ld butterflies f bth sexes the data f M and H had t be cmbined t achieve enugh bservatin time. Fr the males it als has been necessary t use data frm H fr the ther age classes. Fr the 3-10 day ld females data f 1-2 day ld females have cmpletely been taken fr the last 3 time classes in rder nt t miss the age dependent eth-chrngramme. 151
00 cd cb 00 00 00 00 00 00 ös J cri d i- 1 «st n T- i- i- C\J CM date (1990) Fig. 1. Minimum daily ppulatin sizes (minimal number alive; fr days withut bservatins n entries have been made). 3 4 5 6 7 10 residence time (days) 152
2.5. Ethlgical bservatins. 2.5.1. Definitins. The activities flight, mving, nectaring, vipsitin, each individual. cpulatin, and resting were recrded separately fr Registratin was perfrmed with small prtable cmputers, using a prgramme especially written fr ur study. The bserver randmly chse a butterfly and entered its number int the cmputer. Fr each activity an abbreviatin, cnsisting f ne letter, was used. At the beginning f each activity the bserver pressed the relevant letter, which was saved tgether with the time. The duratin f ne activity was btained frm the difference between the start f the present activity and the start f the next ne. The activity mving cnsisted f diverse activities, which were registered separately in the field, but have been lumped here. These activities were: walking, turning (e.g. females turning n the flwer heads t lk fr suitable egg laying niches), cleaning, fluttering while sitting, and the characteristic lycaenid behaviur f parallel up and dwn mvement f the clsed wings. Thus, turning when lking fr a suitable egg laying niche was nt cunted as egg laying behaviur, as it culd nt be clearly separated frm nectaring. A female lking fr an egg laying lcality and turning therefre n the flwer, every nce in a while is nectaring. vipsitin was defined as the time while the butterfly has put its abdminal tip between individual flwers f a flwer head. Smetimes females rapidly changed between different single flwers, perfrming abdminal bends and tuching the surface f the flwer head with the abdminal tip. Each f these events was regarded as ne vipsitin behaviur (sensit Figurny & Wyciechwski, 1998). Fig. 2. Relative bserved residence time and residence time accrding t the Jlly-Sebermethd. There are hardly any differences between males and females. The residence prbability within the ppulatin is 74% fr each age, accrding t Jlly-Seber. This means that 74% f all individuals reside fr 1 day, 55% fr 2 days,... 5% fr 10 days. This expnential dependence nly is valid if residence prbability is age independent. As the bserved number f residing individuals decreases expnentially as well, age independence can be assumed. The figure prves the intensive assessment f the ppulatin, because therwise the Jlly-Seber-curve wuld be clearly abve the bserved values. 153
Nectaring was scred as the time when the butterfly put its prbscis int the flwers r when it rapidly mved frm ne flwer t the next lking fr the best nectar surce. Abut 150 hurs f bservatins were registered n ur prtable cmputers, cnsisting f abut 1 1,000 events. 2.5.2. Analysis f behaviural bservatins 2.5.2.1. Prcedure and classificatin. The data have been analysed s that n the ne hand the diurnal distributin f the single activities can be described (diurnal eth-chrngramme), and, n the ther t quantify hw the sexes differ in their behaviur and hw behaviur changes with age (age dependent eth-chrngramme). The data were therefre classified as fllws: sex: male, female age: days, 1-2 days, 3-10 days time intervals: 10:00-10:50, 10:50-11:40... 17:30-18:20 activities: flight, mving, nectaring, Qgg laying, cpulatin, resting The classes were kept as small as pssible. Even s, the activity cpulatin caused prblems in analysis, as there have been relatively few events lasting fr relatively lng time. This prcedure resulted in every day cnsisting f 10 classes f 50 minutes each. The ttal bservatin time within each class (i.e. the ttal f all single events) had a length f at least 3,000 secnds (= 50 min.). Hwever, because few activities ccurred early and late in the day, the first class summarised all results befre 10:50 Central Eurpean Summer Time (= GMT r 9:20 lcal time), starting with the first bservatins at arund 9:30; the last class summarised results after 17:30 (GMT), with last bservatins arund 19:30. Mst data f the first class are indeed frm 10:00-10:50 and f the last class frm 17:30-18:20. Sexes were treated separately and three age classes frmed (see als results) with an apprximately even distributin f ttal bservatin time. This resulted in the age classes 0, 1-2, and 3-10 days. 2.5.2.2. Diurnal Eth-Chrngrammes (Figs 3, 4). bservatin time (t) f ne activity was summarised within ne class (e.g.: males, age: 1-2 days; time: 10:50-11:40): 154
^0 Scietas Eurpaea Lepidpterlgica; dwnlad unter http://www.bidiversitylibrary.rg/ und www.zbdat.at ^ tfiight. l-2days. 10:50-11:40. all males" tflight, l-2days. 10:50-11:40. male 1 + tflight, l-2days, 10:50-11:40, male 2- + + tflight. l-2days. 10:50-1 1:40. male n \*-) Then the relative prprtin (t) f ne activity cmpared t the ttal duratin f all activities was calculated. This was dne as in the fllwing example fr flight f the 1-2 day ld males fr the perid 10:50-11:40: r flight. 1-2 days. 10:50-11:40. all males = ^ tflight. 1-2 days. 10:50-11:40. all males' \^ tflight, 1-2 days. 10:50-11:40. all males + 2- tmving. 1-2 days. 10:50-11:40. all males + ^ «nectaring, 1-2 days, 10:50-11:40, all males + 2* tcpulatin. 1-2 days. 10:50-11:40. all males + ^ t resting. 1-2 days. 10:50-11:40. all males) \^) Figs 3 and 4 shw the diurnal eth-chrngrammes f 1-2 day ld males and females. Within the ther age classes, data were insufficient t prvide a cmplete diurnal distributin. Nevertheless, diurnal eth-chrngrammes were cmputed as these were needed in the age dependent eth-chrngrammes. 2.5.2.3. Age dependent Eth- Chrngrammes (Fig. 5). The age dependent eth-chrngrammes shw hw the distributin f activities changes with increasing age and hw the sexes differ. Abslute values (in minutes) were calculated frm the relative prprtins f the time classes f the diurnal eth-chrngrammes. In ther wrds, we calculated hw lng a virtual butterfly f a certain age class perfrmed each activity during a whle day. The cmputatin is shwn fr the ttal flight time f 1-2 day ld male butterflies: 2- tflight. l-2days. all males = (tflight. 1-2 days, befre 10:50. al males + ^ flight. 1-2 days. 10:50-1 1:40. all males +... + Tflight, 1-2 days, after 17:30. all males) ' ITlin \D) In sme classes the data were insufficient (bservatin time in the 50 minute class was less than 3,000 secnds) t reach a balanced distributin f activity. In these cases, additinal data were included frm a nearby lcality (H - Hagelgrund, see chapter 2.2). Even s, data fr the last three time classes f 3-10 day ld females were still t few (see Tab. 1). Since the values fr the first six time classes (10:50-15:50) f 3-10 day-ld animals were very similar t thse fr the diurnal eth-chrngrammes f 1-2 day ld insects, we assumed that the values f the three remaining classes were nly slightly different frm the values f 1-2 day ld females. Althugh this is nt entirely crrect, fr the sake f btaining a 155
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cmplete picture f the age dependent eth-chrngrammes we decided t include values f 1-2 day ld females fr cmputing values f 3-10 day ld females. In the first class (befre 10:50) f the day ld insects, n data were available, as the insects had just been marked. T achieve a cmplete picture, the values f the 1-2 day ld butterflies were als used in the first time class f the day ld nes. Fr an verview f the manipulatins perfrmed fr the cmputatin f the age dependent eth-chrngrammes see Table 1. 2.6. Discussin f methdlgy 2.6.1. Handling and behaviur. Mrtn (1984) states that handling in sme butterfly species can affect recapture prbability. He als mentins that different species react quite differently in this respect. Hwever, fr G. nausithus there is n indicatin fr a handling effect. Fr example, neither Geißler-Strbel (2000) nr Binzenhöfer & Settele (2000) detected any behaviural change after handling this species; sme individuals even cpulated immediately after the handling and females cntinued t lay eggs (Binzenhöfer & Settele, 2000). 2.6.2. bservatin and behaviur. The presence f (generally) three bservers n a relatively small site might have influenced adult behaviur, especially flight activity and emigratin. Hwever, ur bservatins suggest that the insects were nt affected. We bserved the butterflies frm an average distance f abut 2 meters, where they did nt shw any reactin, althugh ne can apprach as clse as few decimetres withut nticeably affecting behaviur. Althugh the S. fficinalis plants were ccasinally shaken, this was greatly utnumbered by the number f natural disturbance events e.g. by males perfrming curtship behaviur r by ther insects. 2.6.3. Limits f bservatin time due t methdlgical cnstraints. The insects were bserved fr as lng as pssible. Hwever, we smetimes lst track f flying adults, especially males, leading t an underestimate f this activity. 159
Table 2: Residence times f G. nausithus n the main research site (n number f butterflies marked) ttal Number f days 1 2 3 4 5 6 7 8 9 10 Males (n = 68) 26 Females (n = 57) 25 13 10 7 3 7 4 8 7 1 1 3 2 4 3 1 Table 3. Average residence time f the Maculinea species. Species Sex Average residence time [days] Reference Glaucpsxche {Maculinea) nausithus â 1.9 present paper 3 G. (M.) nausithus 9 2.0 present paper 3 G. (M.) nausithus S 9 2.3 present paper" G (M.) nausithus 6 5.4 Geißler-Strbel (2000) c G. (M.) nausithus 9 3.3 Geißler-Strbel (2000> G. (M.) nausithus S 1.3 Laux(1995) b G. (M.) nausithus 9 0.8 Laux (1995) G b (M.) nausithus S ca. 4 Seiler (1991)" G. (M.) nausithus V ca. 3 Seiler ( 1991 )" G. (M.) nausithus S 9 ca. 0.7-2.3 Wynhff(1998) cd G. (M.) teleius (Beresträsser) S 9 0.7 Laux(1995) G b (M.) teleius Ö* 9 ca. 0.3-2.8 Wynhff (1998) cd G (M.)arin(L.) ct 9 1.8 Pauler^ö/. (1995) ac G. (M.) rebeli (Hirschke) 6 1.8 Kckelke ef a /. (1994) ac G (M.) rebeli (Hirschke) 9 2.5 Kckelke é>f a/. (1994) ac a Average residence time: difference between first and last sighting. h Average residence time based n Jlly-Seber estimates. c The authrs mentin ne day mre, as they regard an individual at the day f first sight as ne day ld already. Thus the data have been crrected by -1 fr this cntributin, t make cmparisn f data pssible. d data frm 1991 until 19%. Table 4. Number f eggs laid per female per day, based n mean daily behaviural prfiles. Age class day 1-2 days 3-8 days Number f eggs laid (estimated*) 80 94 81 accrding t ur estimate; we assume that in abut 95% f all events eggs have really been laid (cmpare text). vipsitin behaviur 160
( 10:00 Scietas Eurpaea Lepidpterlgica; dwnlad unter http://www.bidiversitylibrary.rg/ und www.zbdat.at Table 5. Average number f flights and average flight duratin f 1-2 days ld males and females f Glaucpsyche naushhus during ne day f bservatin t 18:20 h Central Eurpean Summer Time). Sex events/dav/individual duratin/dav/individual duratin/event 161 1 min 26 sec 223 38 min 10 sec 3. Ppulatin eclgical results and their discussin 3.1. Ppulatin dynamics. The flight perid f the ppulatin lasted frm 13 July until 28 August 1990 (at the site Mswieser Tal ). Weather cnditins were favurable fr nearly the whle perid, with the exceptin f tw rainy days and ne further day with maximum temperatures belw 20 C. Thus, perids f reduced abundance were neither expected nr bserved during the curse f the ppulatin curve (based n the minimal number alive: see Fig. 1). The prterandric phenlgy f G. nausithus is clearly visible in Fig. 1. Fr example, the date when half the males in the whle ppulatin had been bserved (29 July) is clearly (5 days) earlier than the date fr half the females (3 August). In G. nausithus. prterandry has been bserved in many instances (Geißler-Strbel. 2000: Garbe. 1991: Laux. 1995) and is a cmmn phenmenn in ther butterfly species (see e.g. Wiklund & Fagerström. 1977). ur ppulatin reached its maximum n 3 August 1990. with a minimum number (see chapter 2.4) f 18 males and 13 females (see Fig. 1). This was als the day f maximum male numbers, while that f females (with 15 individuals) ccurred n 7 August. As the ppulatin curve is asymmetric (left sided), the date at which half f the ttal annual ppulatin appeared is earlier (29 July fr males and 3 August fr females, see abve). 3.2. Ppulatin size. In ttal. 68 males and 57 females were marked n the main site. A further 18 males and 20 females were marked elsewhere, f which 28 butterflies were encuntered near the settlement Alte Schmelz. >1.5 km away. Nne f these marked specimens were fund entering the main site, nr was any specimen fund utside after beine marked n the main site. 161
Due t the intensive effrt, nearly every adult entering the ppulatin was marked n the day it emerged. Thus, an estimate based n the daily estimates and a summatin f the ^-values (i.e. the individuals added t the ppulatin between tw ppulatin estimates) f the Jlly-Seber mdel (see e.g. Seber, 1982, and Settele et al, 2000) resulted in 122 specimens, three specimens less than the 125 marked individuals. The use f this methd was nt ideal fr this study because the small ppulatin size and shrt residence time f individuals resulted in few recaptures n sme days. Thus, fr a mre thrugh statistical analysis, nly the central 18 days f field assessment culd be used (mitting the first 5 and the last 7 days). Fr this perid the ttal ppulatin size is 119, with a 90% cnfidence interval f +/- 16 (and a MNA f 1 12; resulting in a ppulatin size between 112 and 135; calculatin accrding t Settele et al., 2000:181, Tab. 5.14a; but see "http://www.ufz.de/spb/nat/settele/tagfalter.html" fr crrectins). Estimates f ttal ppulatin size supprt ur ntin that the ppulatin was almst cmpletely assessed n the main site. In cntrast, the number f individuals utside the main site was undubtedly larger than the marked 38 specimens (due t lw intensity f field wrk and thus lw numbers f marked and recaptured specimens Jlly-Seber estimates can nt be perfrmed). But, as n individual amng thse marked utside the main site was recaptured within the site and vice versa, we assume that the number f individuals immigrating int the site must have been very lw. Additinally, nne f the specimens freshly marked n the site shwed bvius signs f extensive previus flight activity (like lss f scales r fringed wing margins), which wuld have suggested it was an immigrant. 3.3. Dispersal and islatin f the ppulatin. ur assumptin f lw emigratin and immigratin is cnsistent with the statements f lw mbility f the species by Bink (1992) r Weidemann (1995). New studies hwever reveal that it is much less sedentary (Settele et al, 1996). The lngest dispersal distance bserved s far is 5100 m (Binzenhöfer & Settele, 2000). Detailed studies n the species' mbility (e.g. Geißler & Settele, 1990; Binzenhöfer & Settele, 2000) have mainly been perfrmed in landscapes with numerus lcal ppulatins and rather large habitat patches. There- 162
fre these distances might be due t mbility within habitats r between habitats, using stepping stnes (e.g. singular Sanguisrba stands) in the latter case. The lw density f habitats in the study regin and the islated situatin f the study site (due t a large prtin f surrunding frests) makes it less prbable, that butterflies immigrate frm ther ppulatins in large numbers. As in the study f Laux (1995), high recapture rates indicate cmparatively sedentary behaviur in these types f envirnments. Sme emigratin as well as immigratin can f curse never be excluded. 3.4. Residence time. It fllws frm 3.3 that the vast majrity f butterflies must have hatched frm the site itself. We can als be cnfident that the majrity f animals has been marked n the day f hatching, s their date f eclsin is knwn. Cnsequently, it was generally pssible t identify the exact age f a butterfly at each activity r recapture. The average residence time was estimated based n Jlly-Seber (Seber, 1982; Pllck et al., 1990) estimates f the ppulatin size and residence rate ((p). The average residence rate equals -(\nçp)' 1 and resulted in a value f 0.74, which is a weighted average (with respect t the daily ppulatins size), cvering the whle flight perid. Because f the small ppulatin size, bth sexes were cmbined, which des nt intrduce bias s lng as bth sexes have similar recapture rates, as here (61.8% fr males, 56.1% fr females; cmpare Tab. 2, which als gives an indicatin fr the frequency f capture as assessment was made nearly every day and the z-values fr the Jlly-Seber-estimate are mstly 0). The residence time f the different age classes (in days) can thus be expressed as functin f(age) = 0.74 age (Fig. 2). The average residence time is the average time ne individual spends in the ppulatin r the time when the ppulatin is reduced by 50%, i.e. f(age) = 0.5. In ur case, the average residence time was 2.3 days, which is nly slightly higher than the average value resulting frm the difference between the day f marking and the day f last bservatin f all butterflies (2.0; cmpare Fig. 2 and Tabs 2, 3). This is a functin f the cmprehensiveness f ur bservatins. As ur ppulatin has bth been studied intensively and experienced lw lsses due t emigratin, we can regard the average residence time as a gd apprximatin f the average life time. If we 163
additinally regard the average residence times f ther authrs (see Tab. 3), we can state that the lngevity f adult G. nausithus is rather shrt cmpared t ther butterfly species (cmpare e.g. Sctt, 1973). Lking at ther Maculinea species (Tab. 3), this seems t be typical fr the whle taxn (but nt exclusive, as shwn fr many ther temperate-zne Lycaenidae: Väisänen et al., 1994; Arnld, 1983; Warren 1992). The bserved age independent survival prbability f the species als indicates that the butterflies reach their ptential physilgical age nly in few exceptinal cases, as in all free living rganisms. n ur study site we had 6 cases f spider casualties (web r crab spiders). 2 butterflies were killed by cars n a nearby rad. ne specimen just died and drpped ff a Sanguisrba stem fr n bvius reasn. Despite ur intensive study, we thus knw the reasns fr the death f nly 8 ut f 125 marked specimens. We nly can assume the high activity f ptential predatrs (high densities f damselflies, dragnflies and birds alng the nearby creek; further spiders with undiscvered prey, r ncturnal predatrs). 3.5. Availability f Sanguisrba flwer heads as nectar surces. The number f flwer heads was partly cunted. Stands with cunted flwer heads were used t estimate further Sanguisrba stands by eye. The number f heads increased frm ca. 400 in the early flight days t ca. 1600 twards the end f the emergence. The number f flwer heads must have been sufficient thrughut the flight perid, as it prbably has nly t prvide the carbhydrates needed fr flight, as is the case with mst shrt lived butterflies, which nrmally acquire the resurces needed t develp eggs during the earlier larval stage (capital reserves, cmpare Sibly & Calw, 1984, 1986). 3.6. Availability f vipsitin sites and emigratin. Eggs are exclusively laid n Sanguisrba fficinalis. As shwn by Thmas (1984) and intensively studied by Figurny & Wyciechwski (1998), clsed r nly slightly pened flwer heads are preferred. Nearly all flwer heads n the study site were in the preferred stage. The ttal number f flwer heads then steadily increased, reaching a maximum f abut 1600 twards the end f the flight perid, althugh the number f clsed r nly slightly pened nes was lwer than in the beginning. 164
Based n ur behaviur data (see belw) we tried t estimate the average number f eggs laid per female per day. Laux (1995) stated, that in G. nausithus nly an egg laying behaviur that lasts at least 3 secnds results in an egg being laid. In mst cases she registered ne egg per egg laying event. In ur study, in 95% f all cases, egg laying behaviur lasted at least 3 secnds. In fact, nly in 22 f 53 egg laying events (42%) bserved by Laux (1995) were eggs actually laid. Hwever, we were unaware f this bservatin when we cllected ur data, nr did we interrupt the time registratin when a female quickly inserted her abdmen between single flwers, which led t the high value f 95%. n the ther hand, this presumed verestimate will be cmpensated fr, as several egg laying behaviurs sensu Laux (1995) have been regarded as nly ne. This has t be kept in mind when interpreting the fllwing rugh estimates f eggs laid. (Precise empirical data n egg prductin and its dynamics are lacking fr all Maculinea species. Such data are urgently needed in rder t evaluate the usefulness f published estimates, like in Bink (1992), as well as fr parameterizing mdels.) Accrding t ur estimates, just under 10,000 eggs were laid n ur study site (assuming 60 females with an average life expectancy f 2 days and an egg prductin f 85 per day; see Tab. 4). The number f Sanguisrba flwer heads was arund 1 600 at the end f the flight perid, many f which were nt available during the peak f ppulatin density at the main flight perid. ther flwer heads never reached the size necessary fr G. nausithus females t lay eggs n (accrding t Figurny & Wyciechwski, 1998, the flwer head has t have a length f at least abut 6.5 mm). If we assume that abut three quarters f all heads were used fr vipsitin, we wuld have 1200 flwer heads used. This wuld give an average f 8 eggs per flwer head. This is f the same rder f density reprted by Fiedler (1990), wh fund an average f 7.65 eggs per head in habitats with high density f butterflies but relatively few Sanguisrba plants (with a maximum f mre than 20 per head). Fiedler (1990) fund that mre than 3 larvae seldm survived in ne flwer head. Larger numbers f eggs per flwer head thus have lw survival prbability, which means that the number f egg-lay - 165
ing pssibilities may limit ppulatin size. G. nausithus, accrding t Figurny & Wyciechwski (1998), des nt seem t be able t recgnise whether a flwer head already cntains eggs. Hence, the increased tendency fr dispersal within the secnd half f the flight perid, which was bserved by Garbe (1991), may be stimulated by decreasing number f flwer heads available in a suitable stage. As in ur study, the number f preferred nn-flwering flwer heads (cmpare Figurny & Wyciechwski, 1998) was (even abslutely, nt nly relatively) decreasing twards the end f the phenlgy, in spite f an increasing ttal number f flwer heads, we culd als imagine such effects t be f relevance fr ur ppulatin. Hwever, due t the small number f marked butterflies, we have n prf fr differences in residence times during the phenlgy f the species. This merits further study. 4. Results and discussin f behaviural bservatins 4.1. Precnditins fr the interpretatin f behaviural bservatins. As the ppulatin results shw, mst adults were marked n the day f hatching. Thus we culd identify the age f the butterfly at a later recapture. We cnsidered that there was n lack f fd, resting sites r similar resurces. We assume that the flwer heads ften receive mre eggs than they can supprt, as the females cannt identify whether a head is already ccupied. Thus the behaviur f the adults wuld nt have been affected. 4.2. Handling. ur results shw that <1 day-ld individuals are less active than lder nes (Fig. 5). Fr the age class 3-10 days (diagrams nt shwn) the diurnal eth-chrngrammes fr bth sexes are very similar t thse f the 1-2 days ld adults (Figs 3, 4). In cntrast, the results fr age class (diagrams nt shwn) are very variable and shw n clear maximum f activity arund nn. As stated, Binzenhöfer & Settele (2000) fund n handling effect fr this species (see 2.6.1). ur data shw that insects are less active n the day f marking than n ther days. This might be explained as natural fr the species r as a handling artefact. Hwever, as males and females have been affected in a very similar way, we regard the cmparisns between bth sexes as justified. 4.3. Cpulatin and vipsitin. Fr the 1-2 day-ld males, the diurnal distributin f activities is strngly influenced by the 166
ccurrence f cpulatins (see Fig. 3). Althugh we bserved nly 2 cpulatins, these have a strng influence because f their lng duratin. The females f this age class nrmally had n cpulatins (Fig. 4). In ttal, three cpulatins were bserved fr their entire length, which lasted between 24 and 34 minutes. If bservatins frm Eselsbachtal are included, we knw the age f 12 females and 10 males f a ttal f 13 cpulating cuples. Females prbably cpulate nly nce, in all but ne bserved case n the day f emergence. f the 13 cpulatins, 11 were in the mrning r early afternn (until ca. 14:30 h). nly twice was cpulatin bserved after 17:00 h. These results indicate that females cpulate as early as pssible. nly in ne female at Eselsbachtal was cpulatin bserved n the next mrning after the day f marking (but then earlier than all ther cpulatins with newly hatched females; at 10:27 h, when the cuple was sighted, cpulatin was already ging n). The 10 males f knwn age at cpulatin were up t 7 days ld. The sexual activity f males seems t last fr their whle lifetime. Tw f the males were bserved cpulating twice. Accrding t SBN (1991), lder Maculinea males are mre ften invlved in cpulatins than yunger nes, which als fits t the bserved lwer activity f day adults in ur study. And if males mate less ften when they are yung, it wuld nt be surprising as they first have t reach sexual maturity (sclertizatin f genitalia, maturatin f testes), which may take lnger in males f prtandrus species (wn unpubl. data; Fiedler, pers. cmm.). In ne case, egg laying behaviur was bserved as sn as 17 secnds after the end f cpulatin, althugh we did nt see whether an egg was actually laid. In a secnd case, a female did nt lay fr 35 minutes, and was then lst frm sight; hwever, after 1 hur and 44 minutes, the same female was resighted laying eggs. The cpulatin f females immediately after eclsin and mating may be seen as cnsequence f their shrt residence time. Befre leaving the ppulatin (due t death r emigratin) they leave as many eggs as pssible in their habitat f rigin. Accrding t Fiedler (pers. cmm.) this has t be regarded as a remarkable exceptin frm all ther Plymmatinae lycaenids, where nticeable egg prductin starts abut 1-2 days after mating and reaches its peak anther 1-2 days later. 167
4.4. Flight. Flight in bth sexes peaked shrtly after 12:00 h lcal time (i.e. 13:30 h central Eurpean summer time) when the sun was already in its highest psitin, but temperature had nt yet reached its maximum (cmpare Figs 3 and 4). Diurnal changes in activities were mre prnunced in males (Fig. 3) than in females (Fig. 4). In the males they are clearly paralleled with the time spent fr nectaring, while in females the times are similar but less clear. During a whle day, males fly twice as lng as females (Fig. 5), althugh there has been an underestimatin in the latter (cmpare 2.6.3). The number f flights per day hwever is greater in females (Tab. 5), resulting in a much lwer duratin per flight. Thus, within the habitat females mainly undertake very shrt flights (i.e. frm ne flwer head r plant t the next in search fr vipsitin sites). In bth sexes, sme flights serve t reach nectar surces. Females als fly t reach egg laying sites. Ptential nectar and Qgg laying plants can nrmally be reached in very shrt distances. Males repeatedly undertake lnger flights searching fr females. The mre a male flies, the higher his prbability f finding an unmated female. n each male there is selectin t be mre active in flying and thus mre successful in finding unmated females than ther cmpeting males. Thus, we can expect a higher reprductive success in mre actively flying males, which explains the bserved differences in sexes in terms f flight. In sme butterfly species there are temperature-dependent differences in male searching behaviur fr females, which might indirectly lead t a diurnal switch. In Cennympha pamphilus (L.), fr example, Wickman (1985) bserved that males tend t perch within their territries at lwer temperatures, while at higher temperatures they patrl. Als in G. nausithus males, the greatest flight activity is arund nn, but temperature dependent change in behaviur (e.g. frm patrlling t perching) was neither bserved nr expected. As females cpulate directly after eclsin and are nt active flyers, it wuld represent a pr strategy fr males t perch rather than patrl and actively seek newly eclsed females (see Rutwski, 1991, fr a review n male mate-lcating behaviur). Because they patrl and mate with freshly emerged females, males have t stay within their habitat t reprduce successfully 168
rather than undertake lng distance flights. The latter risk des nt exist fr females, which makes them mre suitable agents fr the (genetic) exchange between ppulatins and the clnisatin f new habitats (independent f the numbers f males which als leave the habitat). This fits t the bserved higher flight distance f females and the bservatins that females have, perhaps temprarily, left the site mre ften than males. Dispersal f males frm ne habitat t anther, as bserved by Binzenhöfer & Settele (2000) and Geißler- S trbel (2000) cannt be disregarded. Its relevance depends much n the distance between habitats r the degree f islatin f the riginal habitat. Nearby habitats might be reached s easily that individuals might be regarded as belnging t the same ppulatin (cmpare discussins f gradual changes frm subppulatins t prper ppulatins and finally metappulatins; e.g. Hanski 1999, Settele 1998). In such cases, male dispersal is expected. 4.5. Mvement and nectaring. Nectaring is strngly assciated with flight, especially in males (Fig. 3). Females f all age classes shw mre prnunced mvement n flwer heads than males (Figs 3, 4, 5), as females have t mve fr nectaring as well as between each egg laying event. In females, nectaring and egg laying may be cmbined as bth activities ften happen within a shrt time n the same flwer head. Males and females bth spend less time nectaring n the day f eclsin than when lder (Fig. 5). The strng decline in nectaring f 3-10 days ld males is prbably due t the strng influence f the few but lng lasting cpulatin events. Mst prbably in males and females, nectaring has the same duratin, as e.g. als bserved fr Anthcharis cardamines (L.) by Wiklund & Âhrberg (1978). As shwn in Fig. 5, bth sexes are active fr the same perid f time, althugh the type f activity is very different: males spend mre time n lng patrlling flights while females invest mre time visiting flwer heads and egg laying. It can be assumed that time f activity is temperature limited. Hwever, the butterflies ften rest arund nn (Figs 3, 4), presumably t recver r t avid verheating. 169
Acknwledgements Fr the cnstant and patient field bservatins we are very much indebted t Frank Lhner, Karin Bink, Carla Kleinschmidt, and Mari Nenn. Fr rganisatinal assistance we want t thank Uwe Kch and Michael Werner. The cmputers fr field data registratin have kindly been supplied by Dr. de Kramer (BASF cmpany, Ludwigshafen). The Fachschaft Bilgie f Kaiserslautern University gave financial supprt fr equipment. Fr general supprt and c-peratin we thank Sabine Geißler-Strbel. Critical and very cnstructive cmments n earlier versins f this manuscript have been prvided by Irma Wynhff and Knrad Fiedler. Jeremy Thmas kindly made final cmments n cntent and language. References Arnld, R. A., 1983. Eclgical studies f six endangered butterflies (Lepidptera, Lycaenidae): Island bigegraphy, patch dynamics, and the design f habitat preserves. Univ. Califrnia Publ., Ent. Ser. 99: xii+161 p. Bink, F. A., 1992. Eclgische Atlas van de Dagvlinders van Nrdwest-Eurpa. Schuyt & C, Haarlem, 512 biz. Binzenhöfer, B. & Settele, J., 2000. Vergleichende autöklgische Untersuchungen an Glaucpsyche (Maculinea) nausithus Bergstr. und G. (M.) teleius Bergstr. (Lepidptera, Lycaenidae) im nördlichen Steigerwald. In: Settele, J. & Kleinewietfeld, S. (Hrsg.). Ppulatinsöklgische Studien an Tagfaltern 2 UFZ-Bericht 2/2000 (in press). Elfferich, N.W., 1998. New facts n the life histry f the dusky large blue Maculinea nausithus (Lepidptera: Lycaenidae) btained by breeding with Myrmica ants in plaster nests. Deinsea 4: 97-102. Fiedler, K., 1990. New infrmatin n the bilgy f Maculinea nausithus and M. teleius (Lepidptera: Lycaenidae). Nta lepid. 12(4): 246-256. Figurny, E. & Wyciechwski, M., 1998. Flwerhead selectin fr vipsitin by females f the sympatric butterfly species Maculinea teleius and M. nausithus (Lepidptera: Lycaenidae). Ent.gener. 23: 215-222. Garbe, H., 1991. Zur Bilgie und Öklgie vn Maculinea nausithus Bergstr. (Lepidptera, Lycaenidae). Diplma Thesis, University f Marburg (unpublished). Geissler-Strbel, S., 2000. Autöklgische Untersuchungen zu Maculinea nausithus im Filderraum bei Stuttgart. In: Settele, J. & Kleinewietfeld, S. (Hrsg.). Ppulatinsöklgische Studien an Tagfaltern 1 UFZ-Bericht 1/2000 (in press). Geissler, S. & Settele, J., 1990. Zur Öklgie und zum Ausbreitungsverhalten 170
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