© Entomologica Fennica. 3 October 2012

Can brief marking campaigns provide reliable dispersal estimates? A Nickerl’s Fritillary (Melitaea aurelia, Lepidoptera: Nymphalidae) case study

David Novotný, Martin Konvièka & Zdenìk Fric

Novotný, D., Konvièka, M., Zdenìk, F. (2012) Can brief marking campaigns provide reliable dispersal estimates? A Nickerl’s Fritillary (Melitaea aurelia, Lepidoptera: Nymphalidae) case study. — Entomol. Fennica 23: 155–164. Functions expressing dispersal probability decays with increasing distance are widely used in studies of animal movements. The inverse power function (IPF) exhibits the property of self-similarity, and hence should perform robustly against variation in marking efforts, allowing comparisons across studies. We in- vestigated this function property using dispersal data of Nickerl’s fritillary (Melitaea aurelia), a little studied checkerspot butterfly which is currently ex- panding in Central Europe. During mark-recapture in South Moravia, Czech Re- public, a single researcher worked for the entire flight period in 2005, while in 2006 five researchers worked for just 5 days. Slopes of the fitted functions did not differ between the two seasons, illustrating the robustness of the function and suggesting the possibility to obtain reliable dispersal estimates even from brief marking campaigns. For both years, it was predicted that approximately one indi- vidual per one thousand would cross 10 km distance, the maximum distance sep- arating the most isolated colonies in the region. D. Novotný*, M. Konvièka & Z. Fric, Biology Centre of the Academy of Sciences of the Czech Republic andUniversity of South Bohemia, Faculty of Science, Branišovská 31, 37005, Èeské Budìjovice, Czech Republic; *Corresponding au- thor’s e-mail: [email protected] Received28 November 2011, accepted26 January 2012

1. Introduction cesses, unless saved by occasional colonisation (Hanski 1999). Studies on model groups such as The awareness of insect losses from habitat frag- butterflies have provided ample evidence that in ments surrounded by human dominated land- such isolated populations losses indeed occur scapes, such as the intensive farmlands of West- (Wenzel et al. 2006, Leidner & Haddad 2011). ern and Central Europe, has raised interest in in- The data amassed in empirical butterfly dispersal sect dispersal, or mobility related to gene flow studies, using a variety of approaches from classi- (Habel et al. 2009, Hanski & Mononen 2011). cal mark-recapture to molecular and electronic Losses from habitat fragments are explicable by tools, are now allowing broad generalisations the metapopulation theory, which predicts that which link dispersal to species’ life histories and small local populations are prone to eventual ex- aim to assist practical conservation (Hovestadt & tinctions due to deterministic or stochastic pro- Nieminen 2009, Stevens et al. 2010, Sekar 2012). 156 Novotný et al. • ENTOMOL. FENNICA Vol. 23

Fig. 1. Schematic rep- resentation of dánický les hills, Czech Repub- lic, showing the study site, the distribution of xeric grasslands, and their occupancy by Melitatea aurelia within the mostly intensively farmed, or forested, area.

Checkerspot butterflies (Nymphalidae: Meli- 2002), but is currently re-expanding to histori- teaeinae) represent a popular model in population cally occupied regions (Germany: Eichel & ecology (e.g., McLaughlin et al. 2002, Hellmann Fartmann 2008, the Czech Republic: Spitzer & et al. 2003, Ehrlich & Hanski 2004, Bulman et al. Beneš 2010). It hence offers an excellent compar- 2007, Zimmermann et al. 2011). They often form ison with much more studied, but recently declin- spatially distinct colonies, are easily detectable ing, checkerspots. both as larvae and adults, are rather poor fliers Mobility studies based on capture-mark-re- and occur in high densities. At the same time, they capture often miss rare long-range movements display a remarkable diversity of life history traits (Baguette & Schtickzelle 2003, Zimmermann et (Ehrlich & Hanski 2004), which transfers into al. 2011), instrumental in the recolonisation/ex- varying levels of threat from intensive land use. pansion processes. Franzen and Nilsson (2007) Some European species are among the most en- recommended that a sampling area as large as 100 dangered butterflies on the continent (e.g., Eu- km2 is necessary to estimate lepidopteran phydryas maturna, cf. Freese et al. 2007), while (re)colonisation ability. A way to overcome this others vary in status regionally (e.g. Melitaea logisticlimitation is to use dispersal kernel func - athalia, cf. Hodgson et al. 2009). An interesting tions, predicting long-range movements from case in Central Europe is Melitaea aurelia short-range ones. This does not fully eliminate Nickerl, 1850. This xeric grasslands specialist the problem with undetected long-range dis- has declined severely during the 20th century persal, but at least allows comparisons among (e.g., Van Swaay & Warren 1999, Beneš et al. species, sexes, regions and seasons (Fric et al. ENTOMOL. FENNICA Vol. 23 • Dispersal estimate in brief marking campaign 157

2010). Contributing to the debate (e.g., Baguette corded to date, including 17 nationally threatened et al. 2000, Kuras et al. 2003) on the appropriate ones, according to unpublished Czech Butterfly shape of dispersal kernel functions, Fric and and Moths Recording data), complex land-hold- Konvièka (2007) showed that a particular func- ing patterns have so far prevented legal protec- tion form, the inverse power function (IPF), is tion. A part of the dry grassland area is actively particularly robust to variation in the marking ef- managed by conservation volunteers. fort, owing to the scale invariance of the function slope; i.e., the parameter expressing decay of dis- persal with distance. 2.2. Study species The scale invariance of IPF offers an attrac- tive possibility to collect reliable dispersal data Melitaea aurelia is a univoltine butterfly, dis- with reduced marking effort and to compare dis- playing colonial distribution at short-bladed xeric persal data collected under varying study circum- grasslands with a high cover of its host plant, stances. We examined this by performing a two- Plantago media L. (Eichel & Fartmann 2008). seasons mark-recapture study targeting move- Females lay small egg batches on the underside of ment patterns in a strong M. aurelia population host plant leaves, larvae are gregarious until au- inhabiting xericgrasslands in the CzechRepub - tumn. Its total distribution stretches from France lic. In the first season, we followed a traditional through Central and Eastern Europe, Transcau- approach, marking the population during the en- casia, the Southern Urals and Siberia up to the tire flight period. In the second season, we sub- Tian Shan region (Tolman & Lewington 2008). stantially increased the study area, restricted In the Czech Republic, a decline during the 20th marking to peak adult flight, and increased per- century has restricted its distribution to the warm- day marking intensity. Here, we (1) describe the est regions of the country, but recent records doc- basicdemography and movement patterns in the ument a return of the species to parts of its former population of M. aurelia, and (2) assess the ro- range. bustness of the movement estimates obtained while following different marking protocols. 2.3. Mark-recapture data We worked in four adjoining valleys with a total 2. Material and methods dry grasslands area of 51 ha, divided into eleven subsites for the purpose of dispersal analysis (Fig. 2.1. Study system 1). Two adjoining valleys (32 ha) were covered by marking in 2005, and two others (19 ha) were The study was carried out in a system of valleys at added in 2006. the southern slopes of the Zdanicky les hills In 2005, the marking covered the entire adult (49°05’N, 17°02’E, altitude 245–330 m), South- flight period (25. VI–28. VII), every day with fa- ern Moravia, SE Czech Republic. While most of vourable weather, from 10:00 to 17:00 (CEST). Southern Moravia consists of intensively farmed Each netted butterfly was assigned a numeric lowlands, Zdanicky les is a ridge (maximum alti- code and released at its point of capture. A single tude 437 m), formed by base-rich Carpathian person carried out the marking, working for 32 flysch and covered by oak-hornbeam and beech person-days in total. forests. Its south-facing valleys, historically cov- In 2006, the marking was limited to the peak ered by a patchwork of gardens, hay meadows, of adult flight season (5.–9.VII), but covered an orchards and pastures, were largely abandoned expanded area. Five persons did the marking each several decades ago, and now are in varying day, resulting in the total marking effort of 25 per- stages of succession. The prevailing vegetation is son-days. Daily marking intensity, i.e. the num- classified as Bromion erecti dry grasslands, Ge- ber of marking persons per day and unit area, was ranion sanguinei forb-rich mantles, and Berbe- 3.6 times higher in 2006 than in 2005 (5-fold in- ridion xericscrub(Chytrý et al. 2001). Despite crease in persons marking, but covering an area high bioticdiversity (e.g., 74 butterfly speciesre - 1.37 times larger). 158 Novotný et al. • ENTOMOL. FENNICA Vol. 23

Fig. 2. Daily numbers of captures of Melitaea aurelia males and fe- males.–a.In2005. – b. In 2006. The ar- rows in panel (a) show the 2005 days pheno- logically corresponding to the five marking days in 2005.

2.4. Demography estimates ments, lumped them into 50-m distance classes and dispersal analysis (e.g., Drag et al. 2011), and fitted the inverse power function, expressing the probability den- Population size in 2005 was estimated using the sity (I) of movements as Jolly-Seber method, POPAN parametrisation suitable for open populations with births, deaths, I=C•D–n (1a) and migration, as implemented in the program MARK v. 4.3. The residence rate (Phi) combin- or, in logarithmicform, ing deaths and emigration, capture probability (p) and proportional recruitment (pent) are POPAN ln I =lnC – n(ln D) (1b) primary parameters, which may be constant for sexes and time (·) sex-dependent (g); or factori- To compare the 2006 (shorter term, expanded ally (t), linearly (T) or polynomically (Tx) time- area) data with the 2005 data, we used two 2005 dependent; additive (g+t)orinteractive(g×t). datasets. One consisted of all available data. For POPAN derived parameters are daily recruitment the second, we selected movements on five con- (Bi), daily population size (Ni), and total popula- secutive days with a sex ratio corresponding to tion size (Ng) (Schtickzelle et al. 2002, Vlasanek 2006 data (see Fig. 2a). et al. 2009). We fitted IPFs on these three datasets and To compare mobility between years and compared obtained slopes and intercepts using sexes, we used data on patch-to-patch move- the Tukey HSD test (Zar 1996). ENTOMOL. FENNICA Vol. 23 • Dispersal estimate in brief marking campaign 159

Table 1. Overview of mark-recapture data collected during the study for Melitaea aurelia males (M) and females (F).

Marking Individuals Individuals Capture Movements days M/F captured M/F recaptured M/F events M/F M/F

2005 32 / 29 304 / 108 318 / 98 649 / 230 146 / 20 2005 reduced 5 / 5 113 / 55 62 / 11 191 / 67 29 / 8 2006 5 / 5 827 / 468 455 / 110 1375 / 598 274 / 151

3. Results 1). The total population size for 2006 was not re- alistically identifiable from the data, due to too For the 2005 marking (Table 1), the best demog- low number of marking days, but probably raphy model was Phi(t)p(g+t)pent(g*T2)N(g), reached a few thousand individuals. containing 50 parameters (AICc = 2162.58). The The mean / median flight distances in 2005 second-ranking candidate model differed by were 307±36 SE m / 200 m for males and 203±6 DAICc> 3.0. Melitaea aurelia residence Phi thus m / 100 m for females; in 2006, 377±23 m / 200 m responded factorially to the day of marking, inde- for males and 398±8 m / 250 m for females. Based pendently of sexes; capture probability p varied on the reduced 5-day data, the mean / median dis- among marking days and was always higher in tances were 256±10 m / 200 m for males and males; and recruitment pent was approximated by 132±4 m / 100 m for females. In both years, the domed quadraticpolynomials, differing in shape longest detected movements were those of males between sexes and revealing protandrous recruit- (Table 2). ment (Fig. 3). The total estimated population size All fitted IPFs approximated the dispersal was 490 (±35 SE) males and 270 (±36 SE) fe- data significantly (Table 2), with all R2 >0.7(Fig. males. 4). Comparing the functions across years and In the 2006 marking, we marked 3.1 times sexes did not reveal any overall difference in in- more individuals than in 2005. Based on sex ratio tercepts (F = 0.25, df =1,5,p = 0.618) or slopes on capture, the marking period in 2006 (5.–9. (F = 0.08, df =1,5,p = 0.775). In other words, the VII) corresponded to 10.–14.VII in 2005. The short-term marking in 2006 detected identical males had already passed their annual peak, dispersal parameters as the entire flight period whereas females were just peaking (Fig. 2). The marking in 2005. Post-hoccomparison of all pos - number of butterflies marked in those phenologi- sible combinations, however, suggested a differ- cally corresponding 2005 days was 168 (Table ence in slopes between the total data of females in

Fig. 3. Estimated daily population sizes (± standard er- rors of the estimates) for Melitaea aurelia males and females accordingtobest-fit- ting model for 2005. Black arrows show the phenological phase correspond- ingtothefivemark- ing days in 2006. 160 Novotný et al. • ENTOMOL. FENNICA Vol. 23

Table 2. Test statistics for the inverse power functions linear regressions and derived estimates of Melitaea aurelia movement probabilities to distances longer than those covered by the marking.

d.f.; F; p Predicted movements (%) Max. 1km 2km 5km 10 km distance

2005 males 1,15; 66.4; <0.001 2.94 1.03 0.25 0.09 1,250 m females 1,7; 38.7; 0.004 3.12 1.36 0.46 0.20 600 m 2005 reduced males 1,11; 20.8; <0.001 2.49 1.14 0.41 0.18 750 m females 1,3; 47.8; 0.002 1.06 0.52 0.2 0.01 450 m 2006 males 1,13; 51.4; <0.001 1.79 0.73 0.22 0.09 2,000 m females 1,14; 38.2; <0.001 4.18 2.26 0.99 0.54 1,050 m

2005 and males in 2006 (at p = 0.05), but this sin- data. On the other hand, the increased marking in- gle difference was likely influenced by the differ- tensity in 2006 expanded the data set, but none of ent sample sizes in the two regressions. the two procedures changed the original (i.e., to- The IPF-derived predictions of movements to tal data 2005) IPF kernel. 5 (10) km distances suggest that a population con- Melitaea aurelia adult demography patterns taining 1,000 (10,000) individuals will likely do not deviate from those found in other checker- contain a few individuals moving to such dis- spots (Ehrlich & Hanski 2004, Konvièka et al. tances. The occupancy pattern in a wider region 2005, Schtickzelle et al. 2005, Hodgson et al. (Fig. 1) shows that patch-to-patch distances 2009), or other single-brood nymphalids (e.g., among occupied xeric grassland patches are with- Proclossiana eunomia: Schtickzelle et al. 2002; in this distance range. Boloria aquilonaris: Baguette & Schtickzelle 2003). Neither does the increase in numbers be- tween 2005 and 2006 appear unusual, as abun- 4. Discussion dance fluctuations by an order of magnitude oc- cur frequently in butterfly populations (Hellmann Estimation of the parameters of the inverse power et al. 2003, Schtickzelle et al. 2005, Èizek & function dispersal kernel using the M. aurelia Konvièka 2009). In species with communal lar- mobility data obtained in two consecutive years val nests, such changes may be particularly prom- resulted in very close function parameters, and inent as entire broods are easily destroyed by ca- hence predicted identical migration rates. The tastrophes, parasitoids or pathogens (Ehrlich & functions were identical despite major differ- Hanski 2004, Schtickzelle et al. 2005), which ences in data collection in the two years: while in might be responsible for the rapid patch occu- 2005 we covered the entire flight period, in 2006 pancy turnover in checkerspot metapopulations we marked the butterflies only during the peak (Hanski 1999). flight, albeit with higher marking intensity. Mean and median movements also fell within The unchanged function performance sup- the ranges reported in previous checkerspot stud- ports the conjecture by Fric and Konvièka (2007) ies i.e. a few hundred meters for mean distances that given the scale invariance of IPF, it should and 1–2 kilometres for maxima (e.g., Wahlberg et predict identical dispersal rates under a great vari- al. 2002, Baguette 2003, Hovestadt & Nieminen ety of marking protocols. The authors showed 2009, Zimmermann et al. 2011). The 2006 mark- this by subsampling data for two species, Euphy- ing returned longer mean distances, which could dryas aurinia (Rottenburg, 1775) and Parnassius have been caused by either the increase of mark- mnemosyne (Linnaeus, 1758). Here, we in fact ing area (e.g., Schneider 2003, Zimmermann et subsampled the 32-day 2005 data to 5-day 2005 al. 2011), or by density-dependent emigration ENTOMOL. FENNICA Vol. 23 • Dispersal estimate in brief marking campaign 161

Fig. 4. Fitting of inverse power functions (IPF) expressing the probability density of Melitaea aurelia movements to certain or longer distances against the distances (ln-ln scale). Short-term intensive marking in 2006 returned identical dispersal kernels (i.e., slopes and intercepts of the function) as did the marking in the whole flight period in 2005.

(Nowicki & Vrabec 2011); the data do not allow the values with other checkerspots suggests a tax- distinguishing between the two alternatives. The onomicconservatismin mobility. This justifies mean flight distances obtained with the reduced using data from closely related species, such as 2005 data were by one-third (males) or even by various checkerspots, for such purposes as mod- one-half (females) shorter compared to the total elling or reserve design (cf. Wahlberg et al. 1996, 2005 data, which is attributable to a lower chance Fric et al. 2010), however, the further verification of detecting long individual displacement during on other taxonomicgroups is needed. Differences the shorter marking time. It is hence more encour- between sexes were not statistically significant, aging that IPF did not differ among the three data although male slopes tended to be steeper, indi- sets. cating fewer long-distance movements compared The ranges of fitted IPF slopes (–1.1 to –1.5 to females. This pattern, explicable by different for males, –0.9 to –1.2 for females) were also sim- fitness consequences of dispersal for sexes ilar to other checkerspot studies (e.g., Fric et al. (Zonneveld & Metz 1991), is frequent in butter- 2010, Zimmermann et al. 2011). The overlap of flies (Scott 1975, Baguette & Nève 1994, Hanski 162 Novotný et al. • ENTOMOL. FENNICA Vol. 23

& Thomas 1994, Gutiérrez et al. 1999). Female et al. 2010). Dispersal is a probabilisticprocess dispersal results in establishment of new colo- and the chance to reach more distant sites in- nies, and ultimately, in range expansions. Identi- creases with abundance at the source colonies, cal movements in butterfly sexes, however, were highlighting the need for appropriate conserva- also reported for some checkerspots (Wahlberg et tion management of as large as possible stretches al. 2002, Wang et al. 2004). of suitable habitats (Eichel & Fartmann 2008). In Germany, M. aurelia often forms small colonies at calcareous grassland islets (Eichel & Acknowledgements. We thank the following colleagues for Fartmann 2008). Habel et al. (2009) showed that help in the field: K. Zimmermann, J. Michálek, E. Patáèová, A. Matoušù, T. Tyml and J. Hedrich. Comments despite living in small colonies, it maintains high by J. C. Habel and an anonymous referee much improved levels of genetic polymorphism by occasional the manuscript. This study was supported by the Czech long-distance dispersal. Our IPF-derived move- Ministry of Education (60077665801), Czech Science ment probabilities (Table 2) predict that one indi- Foundation (P505/10/2167), and the University of South vidual per thousand reaches a 10 km distance. Bohemia (04-144/2010/P). Given the xericgrasslands distribution in the re - gion (Fig. 1), this should maintain a mutual inter- References connection of the system. 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