cynthiaBulletin of the Catalan Monitoring Scheme 2011 - no. 11

Cover The CBMS network Centre in a typically Pyrenean environment domina- Detail of the underside of Southern White Admiral Current situation (2011) of the ted by – including Bromion-rich hay mea- Limenitis reducta (photo: A. Miquel). Butterfly Monitoring Scheme dows and pastureland and damp grasslands – with Large Skipper Ochlodes venata feeding on excrement in Catalonia, and the subalpine forest in a few sections. The counts in the (photo: J.C. Vicente Arranz). Balearic Islands area began a few years ago but due to the complexity and diversity of the butterfly communities present the Editorial In all, 67 stations provided complete data time needed for the counters to learn all the species Indicators of climatic change during the 18th CBMS season. Three new was extended. This walk is extraordinarily interesting stations were incorporated into the Scheme, in a CBMS context given that subalpine environ- limatic change is without doubt one of the factors while at a further six stations regular ments have traditionally been lacking in the network. C that will most affect the world’s biodiversity in preliminary counts were undertaken. The The subalpine meadows at Planes de Son are well-pre- the coming years and in Catalonia many organisms – majority of the stations in the BMSAnd served and are host to populations of singular species including – are already responding to this network and the four on Menorca and such as Lesser Marbled (), Pearl-bordered phenomenon. Biodiversity monitoring programmes Eivissa continued to function. In total, (Boloria euphrosyne) and Small Pearl-bordered (B. sele- such as the CBMS provide highly valuable informa- 134,803 butterflies belonging to 159 species ne) Fritillaries, Amanda’s Blue (Polyommatus amanda) tion that enable us to analyse in detail the type and were counted in 2011. and Bright-eyed Ringlet ( oeme). The counts, magnitude of these responses. In recent years a num- the work of Francesc Rodríguez and Montse Ballbé, ber of climatic change bioindicators, whose aim is uring the 2011 season, of the 67 active stations have also detected some of the classic Pyrenean spe- to synthesize all this information into a manageable D all but one (Sadernes) provided sufficient data to cies such as Apollo (Parnassius apollo) and Small Tor- form, have been developed using data from bird and calculate the annual indices of the species found (fig. toiseshell (Aglais urticae), among others. butterfly monitoring programmes. This number of 1). As well, preliminary counts continued at Moià Castellbell Meander (Bages, 150 m). This station Cynthia contains an article that explains which indi- (el Bages, 700 m) and new counts were set up at is situated in a meander of the river Llobregat in a cators are in use and what they are telling us about Puig Graciós (Cingles de Bertí, Vallès Oriental), Vi- typically Mediterranean environment that includes a this question on European and Catalan scales. One ladecans (Baix Llobregat), Betrén and Bagergue (Val certain amount of agricultural land. A fairly diverse such indicator has revealed the existence of a so-ca- d’Aran) and in the Tres Turons Park in El Carmel butterfly community flies here that is totally domi- lled ‘climatic debt’, although in the Mediterranean in Barcelona (Barcelonès). The Betrén and Bagergue nated by Mediterranean species such as Southern area a second indicator that also takes into account itineraries are of special interest for the CBMS as ( cecilia) and Spanish (P. bathseba) Gatekee- water availability provides a better reflection of the they are the first to be established in La Val d’Aran pers, Dusky Heath ( dorus) and False changes in species’ populations that are occurring as and will provide more data from the hitherto poorly Ilex Hairstreak (Satyrium esculi). As well, there are a result of climatic change. The growing frequency represented Pyrenean habitats. The station of Tres good populations of well-known species that include and intensity of droughts, above all, seems to be se- Turons is poor in species but is of special interest as a Marsh Fritillary (Euphydryas aurinia) and Provence riously affecting the butterflies that are most typical reasonably well-preserved habitat located within the Chalkhill Blue (Polyommatus hispana), without for- of Mediterranean environments, as the sharp fall in heart of the city of Barcelona. getting somewhat more locally distributed species their numbers in recent years testifies. The available annual series are shown in figure 2. such as Escher’s Blue (Polyommatus escheri), Dingy We are also delighted to be able to publish in It is worth noting that there are now more than 30 Skipper (Erynnis tages) and Berger’s Clouded Yellow this edition the first serious attempt to establish a stations that have provided data continuously for 10 ( alfacariensis). Gerard Farré is in charge of the list of common vernacular names in Catalan for all or more years. The number of stations with seven or counts. our butterflies. The proposal has been drawn up by more years of data is 53, a figure that gives an idea Compared to 2010, the number of stations fell in a group of experts from the Granollers Natural His- of the possibilities that the CBMS database offers for 2011 but the total number was still over 60 and was tory Museum and the Catalan Lepidopterological the analysis of trends operating over time. one of the best years since the CBMS began. In 2011 Society over the past two years. We hope that it will counts were discontinued at seven stations: Vallgras- be well received and that these names will slowly be- New transects sa, Punta de la Móra, Granja d’Escarp, Aiguabarreig, gin to be used by butterfly enthusiasts to refer to the Llobera (Solsonès, 850 m). This walk takes place in Gerri de la Sal, Les Alberes-1 and Les Alberes-2. The different species present in our country. We believe a dry and fairly open oakwood with sub-Mediterra- station at Vallgrassa alternates annually with those that this is an excellent way of further popularizing nean herbaceous plant communities dominated by of Olesa de Bonesvalls and Olivella. In the other six this group of amongst the general public. the flower blue aphyllanthes Aphyllanthes monspe- cases those in charge of the counts were unable to Finally, in the regular sections of the journal you liensis in most sections. As well, the walk incorpo- continue, a loss that could be merely temporary at will find that ‘The butterfly’ discussed is the Glan- rates areas of bare rock, stands of rushes and damp La Granja d’Escarp, Aiguabarreig and Gerri de la Sal. ville Fritillary, a species that has become one of the . From a point of view of the butterflies, Even so, the loss of these three itineraries is a blow world’s best-known butterflies in recent years. It was Llobera is an extremely diverse itinerary and during since the first two represent an extreme habitat that is selected by Ilkka Hanski and his team as a model the first year with accurate data 62 species were de- under-represented in the CBMS network (arid wes- for the study of metapopulations and has contri- tected. Nevertheless, this figure will undoubtedly tern Catalonia), while the latter at Gerri is the most buted greatly to the development of this important increase in coming seasons. The communities of diverse of all CBMS stations and provides data for a concept, thereby earning itself an important place in Lycaenidae are extremely rich and there is a mix of vast range of species. On the other hand, counts were modern ecology. We believe that this is an excellent Pyrenean species such as Mazarine (Polyommatus se- restarted at Olivella and Pineda, in the latter case by example of the role that butterflies can play in the miargus), Amanda’s (P. amanda), Chalkhill (P. cori- the rangers of the Montnegre-Corredor Park, which advancement of science. don) Blues and other species such as Osiris (Cupido should guarantee their continuity. osiris), Chapman’s (Polyommatus thersites) and Cata- lan Furry (P. fulgens) Blues that are more representa- Habitats represented tive of upland Mediterranean areas. Nympalinae and The main environments and plant communities communities are also diverse and there are represented in the 2011 counts are detailed in Ta- well-established populations of locally rare species ble 1. In all, 75% of the itineraries are located in such as Twin-spot Fritillary (Brenthis hecate). The Mediterranean areas with an important presence of counts are carried out by Mati Morales. holm oak woodland (55% of itineraries). Counts in Planes de Son (Pallars Sobirà, 1540 m). This iti- upland montane habitats continue at around 20% nerary runs around the Planes de Son Interpretation of all itineraries and, with the incorporation of the II Cynthia

Planes de Son, subalpine habitats now appear in 6% Eighteenth year of CBMS cies (fig. 2). This graph shows a considerable decline of the counts. Balance of 2011 season in numbers in many species of Catalan butterflies from 2007 onwards and, indeed, the last five years Species present In terms of abundances, the 2011 season (2007–2011) of the CBMS were the worst of any The list of butterflies detected in 2011 and in pre- was similar to the previous year and the years of the whole data series. This negative tendency vious years can be found in Table 2. In all, 159 spe- tendency of the last five years for low is undoubtedly closely related to the severe droughts cies were detected in 2011, four fewer than in the population levels in many species continued. in 2006 and 2008. As been noted recently, in cer- previous year but 19.1 more than the average for the This situation is exemplified by many of tain Iberian forest ecosystems, extreme droughts not period 1994–2010 (fig. 3). In 2011 no new species the Satyrinae that appear in the counts, only seriously affect the physiological state of trees for the CBMS network were detected, although whose numbers continue to fall, and in 2011 but also have a knock-on effect on higher trophic le- Mountain Clouded Yellow (), Ga- species such as Spanish () vels such as the herbivorous insects that depend on varnie (Erebia gorgone) and Lefèbvre’s (E. lefebvrei) and Southern (Pyronia cecilia) Gatekeepers them. Furthermore, these effects may last for many Ringlets (three true Pyrenean species) and Spanish and Grayling (Hipparchia semele) reached years2. Unfortunately, the slight recovery in counts Marbled White (Melanargia ines) (typical of western their lowest registers since the CBMS counts that was intuited in 2010 did not continue in 2011, Catalonia), all absent from the counts in 2010 and began. An exception to this rule are the which was the third worse season since counts began general poorly represented in the network, were all species that hibernate as adults, which in in 1994. refound. general increased in number. This fact, Constantí Stefanescu coupled with the positive trends the past Changes in abundance: population fluctuations two decades for many of these species, Apart from Camberwell Beauty (Nymphalis antio- 1 Folch i Guillèn, R., 1981. La vegetació dels Països explains the highest-ever CBMS counts in pa) and Large Tortoiseshell (N. polychloros), whose Catalans. Ketres Editora, Barcelona. 2011 for Peacock (Inachis io) and Nettle-tree numbers dropped slightly, the 2011 season was par- Butterfly Libythea( celtis). For the second ticularly favourable for species that winter as adults. 2 Karsholt, O. & Razowski, J., 1996. The year in a row, counts of migrant species were The other hibernating species such as the Nettle- of Europe. A Distributional Checklist. Apollo Books, very low. tree Butterfly (Libythea celtis) and Peacock (Inachis Stenstrup. io) reached by far their highest ever levels since the Weather and counts beginning of the CBMS (Table 1). The fluctuations Fig. 1. Geographical situation of all the stations that The year 2011 was hot throughout most of the cou- in the numbers of Nettle-tree Butterfly in Catalonia have ever participated in the CBMS network (1994- ntry and average annual temperatures were slightly are truly spectacular since its increase has been not 2011), with their official number and name. Also above or just below the averages for 2009, 2006 and only continuous but also widespread over much of shown are the generally accepted boundaries of the 2003, to date the warmest years on record in Cata- Catalonia. The increase in the twoGonepteryx spe- biogeographical regions present in Catalonia1. lonia (see www.meteocat.com). Nevertheless, rainfall cies, Cleopatra (Gonepteryx cleopatra) and Brimstone was very irregular: 2011 was wet in most of eastern (G. rhamni), has also been notable, both reaching in Fig. 2. Distribution of the complete annual series Catalonia (above all along the central coastline and 2011 their second highest figure for the whole data available for all the different stations that have the mountains of El Montseny, Les Guilleries, El set. Even a species such as the Small Tortoiseshell participated in the project (1994–2011). Collsacabra and the eastern Pyrenees) but dry in (Aglais urticae), whose overall tendency was negati- the western half (above all in the upper valley of the ve, increased markedly in 2011. As well, 2011 was Fig. 3. The number of species detected annually in river Segre, the Tremp Basin, the Lleida Plains and also a favourable year for common ‘whites’ () the CBMS network (1994–2011). the Ebro Delta). The previous autumn was genera- such as the Large (Pieris brassicae), Small (P. rapae) lly cold, wet at the beginning but much drier at the and Green-veined (P. napi) Whites, above all in the Table 1. Habitats and plant communities represented end. Winter was characterized by normal or slightly former (Table 2). in the CBMS in 2011, with the number of stations warmer-than-average temperatures (e.g. February), Nevertheless, the majority of Satyrinae and two they appear in. Classification of the vegetation zones but was dry or very dry in much of Catalonia (with fairly common skippers, Small (Thymelicus sylvestris) and plant communities as per reference 1. rainfall 90% below climatic averages). The begin- and Lulworth (T. acteon) Skippers, all of which feed ning of the CBMS season coincided with one of the on various species of grass, suffered severe declines in Table 2. Butterfly species recorded in the CBMS hottest springs of the past decade, above all April, 2011. Spanish (Table 1) (Pyronia bathse- network over the last 10 years (2002–2011). The month in which new temperature records were set ba) and Grayling (Hipparchia semele) reached their number of stations at which a species has been for many Catalan weather stations. Yet spring was lowest ever totals since the beginning of the CBMS. recorded is indicated (out of a possible total of 41 also wet (above all, March) everywhere except along Other species such as Southern Gatekeeper (Pyronia in 2002, 46 in 2003, 51 in 2004, 52 in 2005, 64 in the northern coast and in the north-west of the cou- cecilia), Dusky (Coenonympha dorus) and Pearly (C. 2006, 70 in 2007 and 2008, 66 in 2009, 69 in 2010 ntry, which was generally dry. There was a respite arcania) Heaths and Western Marbled White (Me- and 67 in 2011). as per reference 2. from these high or very high temperatures during lanargia occitanica) declined to a lesser extent, even most of the summer: no heat waves occurred and though their general tendency in recent years is Photo 1. The itinerary at Llobera, in an open dry oak July 2011 was the coldest in Catalonia since 1997. clearly negative and in 2011 their populations con- forest with sub-Mediterranean grassland, generates In the north-east summer was wet, a situation that tinued to fall. Although a more rigorous analysis still data that are very representative of the butterfly was reversed in the second half of the year, which was has to be conducted, it would seem that these falls fauna of the Lleida pre-Pyrenees. A highly diverse marked by a long dry period from August to October in numbers mainly affect univoltine early-flying Sa- group of butterflies fly here, that include noteworthy and a warm dry month of September. tyrinae. Another group of species that declined cons- species such as Twin-spot (Brenthis hecate), High Overall, during the 2011 season only 3.8 weekly picuously in 2011 were the polyvoltine Lycaenidae Brown (Argynnis adippe), Lesser Spotted ( counts were lost per station, practically the same as such as Common Blue (Polyommatus icarus), Spanish trivia) Fritillaries, Marbled Skipper (Carcharodus in 2010. This figure enables us to calculate reliable Brown Argus (Aricia cramera) and Small Copper lavatherae) and many ‘blues’ (photo: J. Jubany). annual indices for all species for the vast majority of (Lycaena phlaeas), all of which reached their second stations. In fact, 87% of itineraries lost six or fewer lowest ever totals for the CBMS period. Photo 2. Despite being widespread in the Pyrenees, counts and in only two cases were more than 10 wee- The year 2011 was not a good year for migra- outside of this area Amanda’s Blue Polyommatus ks lost (fig. 1a). The most difficult periods for the tory species, which in general declined. For instan- amandus is only found in isolated sites in the pre- counts were in spring (second week of March and ce, for the second consecutive year since the great Pyrenees, mountains of the Serralada Transversal, third of April) and, surprisingly, in summer (begin- Painted Lady year of 2009, this species was rare in El Montseny and some of the highest mountains ning of June and end of July) during countrywide Catalonia. The three other well-known migrants in in the province of Tarragona. The Llobera station wet spells (fig. 1b). Catalonia, Red Admiral (Vanessa atalanta), Clouded has a well-established population that is of great Yellow (Colias crocea) and Bath White (Pontia biogeographical interest (photo: J. M. Sesma). Changes in abundances: tendencies daplidice) were all fairly scarce as the fact that all Although the number of both species and indivi- three reached their second worst ever CBMS totals Photo 3. Spanish Marbled White (Melanargia ines) is duals in 2011 fell slightly compared to 2010, these demonstrates. Although a few migrant Plain Tigers a typical Mediterranean butterfly that flies in North differences were not statistically significant. The -spe (Danaus chrysippus) did reach L’Illa de Buda in the Africa and the Iberian Peninsula. Although it can be cies richness (mean and standard deviation calcula- Ebro Delta at the end of July, the species did not abundant in southern , in Catalonia it is rather ted for the 61 stations with comparable data) were appear at any BMS station, unlike the two previous rare and only found in semi-steppe environments in 48.0 ± 18.5 species in 2010 and 47.3 ± 17.8 species years – above all 2009 – in which it was observed the west of the country. In the CBMS network it is in 2011 (Student Test for paired samples, t = 1.17, P at six CBMS stations. As exceptions to this general present in small numbers at the stations of Sebes = 0.25). For the abundance, the values were 2249.2 rule for migrants it is worth mentioning the cases and Timoneda d’Alfés. Adults fly in April and the ± 2097.2 individuals in 2010 and 2045.4 ± 1493.5 of Long-tailed (Lampides boeticus) and Lang’s Short- beginning of May in grassland with feather grasses in 2011 (t = 1.55, P = 0.25). The declines in many tailed (Leptotes pirithous) Blues, which increased (Stipa spp.), their caterpillars’ food plants (photo: species are reflected in a lower position for the 2011 notably in number in 2011 (above all the latter). J.M. Sesma). season in the historical rankings that are calculated This increase could be due in part to the recovery using the annual indices of the 66 commonest spe- of local populations that survived the mild winter Cynthia III

rather than the result of the arrival of large numbers Habitat management and conservation Pasha (Charaxes jasius) (STI = 14.77ºC), Chapman’s of migrant individuals. Indicators of climate change in Green Hairstreak (Callophrys avis) (STI = 14.57ºC) Finally, despite the warm and favourable spring Catalan butterflies and Southern Gatekeeper (Pyronia cecilia) (STI weather, it is worth highlighting the lack of chan- = 14.06ºC) are much higher than those of species ge in many of the typical spring species such as This article discusses the climate change such as Silvery Argus (Aricia nicias) (STI = 4.14ºC), Orange-tip (Anthocharis cardamines), Moroccan indicators that are being developed Mountain Clouded Yellow (Colias phicomone) (STI Orange-tip (A. euphenoides), Green (Callophrys rubi) using data from the CBMS network. The = 6,76ºC) and Pearl-bordered Hairstreak (Boloria and Provence (Tomares ballus) Hairstreaks, Panoptes Community Temperature Index (CTI) has euphrosyne) (STI = 6,95ºC) that have more boreal Blue (Pseudophilotes panoptes) and Spanish Festoon been used to analyse in conjunction data distributions. In a second phase, the temperature (Zerynthia rumina). from both bird and butterfly monitoring indices for the species at a specific locality (for exam- Constantí Stefanescu programmes, and has detected the existence ple, a CBMS station) are used to calculate an average of a so-called ‘climatic debt’. Nevertheless, temperature for the community, weighted according 1 Greatorex-Davies, J.N. & Roy, D.B., 2001. The given that it does not include data on water to the contribution of each species in terms of its Butterfly Monitoring Scheme. Report to recorders, availability, this index behaves abnormally abundance. The result is the community tempera- 2000. 76 pàg. Centre for Ecology and Hydrology, in Catalonia. A second indicator taking into ture index (CTI), a simple way of measuring the rate Natural Environment Research Council, Huntingdon. account this factor would thus seem to be of change in community composition in response to a more appropriate way of understanding global warming that has been adopted as an indica- 2 Carnicer, J., Coll, M., Ninyerola, M., Pons, X., the effects of climate change in the tor of climate change by the European Union3. Sánchez, G. & Peñuelas, J., 2011. “Widespread Mediterranean. Recently, this methodology has been used simul- crown condition decline, food web disruption, and taneously with the vast databases generated by Eu- amplified tree mortality with increased climate n recent years the generation of indicators that ropean bird and butterfly monitoring programmes change-type drought.” Proc. Nat. Acad. Sci. USA, I can evaluate and quantify changes in natural to examine the consequences of global warming on 108: 1474-1478. systems has increased spectacularly. In many cases, these two bioindicator groups. Data from 9,490 and these indicators are calculated using data on popula- 2,130 bird and butterfly monitoring stations, respec- Fig. 1. (a) Coverage of the counts at the different tion trends in bioindicator species that are generated tively, have been analysed from , Sweden, the CBMS stations, and (b) distribution of the lost by biodiversity monitoring programmes. The data United Kingdom, the Netherlands, the Czech Re- counts during the official 30 weeks of the 2011 obtained from the various different European BMS public, and Catalonia. The results we discuss recording season (1 March–26 September). networks, for example, have been used to calculate a here are the fruit of this collective work11. European Butterfly Indicator for Grassland species, As was to be expected, the CTI varies at spatial Fig. 2. Ranking of the CBMS seasons in terms of the which is now used by the European Environment level in both birds and butterflies, with lower values general abundance of the 66 commonest butterflies Agency as one of its reference indices for trends in (that is, with a predominance of species with lower in the CBMS network. The best year to date was biodiversity in the European Union1,2.This indicator, STIs) at higher latitudes (fig. 1a,b). This relationship 1995, closely followed by 2002, and the worst 2008. along with its equivalent based on bird monitoring is linear and highly significant in both groups: simi- Calculations were carried out using the methodology programmes3, reveals how changes in land use (prin- lar losses of 1.47 ± 0.08 x 10-3 ºC and 1.26 ± 0.01 x described in reference 1; annual indexes were cipally, due to the abandoning of traditional agri- 10-3 ºC are observed in butterfly and bird commu- calculated with the TRIM programme. cultural and stock raising methods, and agricultural nities, respectively, for every kilometre further north intensification) are affecting negatively biodiversity we go (fig. 1a,b). Table 1. Evolution of the annual indexes for the at a European scale. Thus, the alarm has been raised Nevertheless, in the same way that the CTI varies 66 commonest butterflies in the CBMS network regarding the need to adopt new strategies in envi- spatially, variation also occurs over time as a result of (2002–2011), based on an arbitrary value of 1 for ronmental policies in the European Union that will climate change. As temperatures increase, the CTI 1994. Annual indexes were calculated with the TRIM help reverse this trend3. at a site should also increase because the number of programme. Aside from changes in land use, within the scien- species that require higher temperatures will also rise. tific community a broad consensus exists to the effect The data accumulated during two decades of Euro- Table 2. Sum of the annual indexes and ranking that climatic change is the other principal motor of pean monitoring programmes (1990–2008) confirm of the abundance of the 20 commonest species global change affecting biodiversity. Changes in the this hypothesis: there has been an unmistakeable in- from the 2011 CBMS season compared to the distribution and phenology of many species have crease in the CTI of both butterfly and bird commu- corresponding figures from the 2010 season. been well documented in recent years4,5 and Catalan nities that mirrors temperature increases, once again butterflies are no exception6,7. Such changes can lead with highly significant linear relationships: 9.3 ± 0.5 Drawing 1. Peacock Inachis io has increased notably to asynchrony of different elements of an ecosystem x 10-3 ºC per year for butterflies and 2.6 ± 0.19 x 10-3 in the last two decades, and in 2011 it attained and cause species’ populations to decline, as has been ºC per year for birds (fig. 2a,b). During this period, its highest ever levels since the beginning of the shown by a number of different studies8,9. The im- the overall temperature in Europe has increased sig- CBMS. Adults come out from hibernation at the end portance of this phenomenon in natural systems has nificantly in a linear sense by 5.50 ± 0.61 x 10-2 ºC of winter or very early in the spring, this species motivated in recent years the establishment of indi- per year (fig. 2c). being one of the first recorded in the season’s cators of climate change. In this article we describe Using the spatial gradients shown in figure 1, it counts. Larval nests on nettles are also very easy to two such indicators and discuss the results obtained is possible to transform these trends in temperatures detect along many transects, both in the spring and from the CBMS data. over time into spatial tendencies. For temperatures, summer (drawing: M. Miró). the increases recorded in the period 1990–2008 are The Community Temperature Index and the equivalent to a northwards movement of 249 ± Drawing 2. Spanish Gatekeeper Pyronia bathseba is climatic debt 27 km, whilst the increases in the CTI are the equi- one of the commonest butterflies in Catalonia. In As commented above, the increase in temperatures valent of northwards movements of 114 ± 9 km for the CBMS is only absent from sites in the Pyrenees, that many species are experiencing as a result of glo- butterflies and 37 ± 3 km for birds. The difference along the sea-coast and in Minorca and Eivissa, in bal warming is obliging many species to shift their between the movement of the temperatures and of the Balearic Islands. Althought is normally one of ranges northwards in an attempt to maintain opti- the CTI is known as the climatic debt: even if both the most abundant butterflies in open Mediterranean mum conditions vis-à-vis environmental temperatu- bird and butterfly populations respond as predic- scrub, the last years have seen a strong decline of res. Presumably, these distribution changes will lead ted, their rhythm of change is not sufficient to keep its populations due to the negative effect of drought to changes at local scale in community structures, up with the temperature change and these (drawing: M. Miró). which will progressively become dominated by spe- thus begin to accumulate a growing ‘debt’ in rela- cies that are better adapted to higher temperatures. tion to their optimum temperatures. Currently, this A few years ago, Devictor and his team developed debt stands at 135 km and 212 km for butterflies a method for testing this hypothesis and for quanti- and birds, respectively. The existence of this climatic fying changes in bird communities as a response to debt will presumably make populations more global warming10. vulnerable to perturbation of all types. Furthermore, They used data from long-monitoring program- these two indicator groups are not responding to cli- mes whose predictions were tested initially on mate change at the same speed, which could lead to French bird communities. It is based on the idea that loss of synchrony in predator-prey interactions bet- both individual species and communities in a certain ween butterfly and bird populations. location can be characterized by means of a tempe- rature index. Thespecific temperature index(STI) The Mediterranean paradox is calculated using data from the distribution of the and a new climate change indicator species in question, that is, the average temperature The results that we mention here are applicable at of the area in which the species is present. For ins- a European scale. Yet, if we look at the results for tance, in the case of European butterflies, the STI for each country individually a surprise is in store: the typically Mediterranean species such as Two-tailed pattern that we have identified fits central and nor- IV Cynthia

thern European countries well, but not Catalonia 2 van Swaay, C. et al., 2012. “The European butterfly 16 Giorgi, F. & Lionello, P., 2008. “Climate change (fig. 3). Here, the linear relationship is not signifi- indicator for grassland species 1990-2011”. Report projections for the Mediterranean region”. Glob. cant for either birds or butterflies and, furthermore, VS2012.019, De Vlinderstichting, Wageningen. Planet Change, 63: 90-104. in the latter group there is even a negative gradient that would seem to suggest that there is a slight ten- 3 European Environment Agency, 2012. “Streamlining Fig. 1. Spatial tendencies in the CTI of butterflies dency for communities to ‘cool down’, that is, to be European biodiversity indicators 2020: building and birds in Europe. Each point corresponds increasingly dominated by species with lower STIs. a future on lessons learnt from the SEBI 2010 to a monitoring station in the year 2005. The This result has been confirmed by an analysis of the process”. EEA Technical report No. 11/2012. temperature is the average in Europe in March– CBMS stations with the longest annual series12. September in 2005. The X-axis indicates the distance This apparent paradox can be resolved in large 4 Parmesan, C., 2006. “Ecological and evolutionary of sites from the extreme south of Catalonia. part by using a new climate change indicator that responses to recent climate change”. Annu. Rev. takes into account not only temperature but also Ecol. Evol. Syst., 37: 637-669. Fig. 2. Trends over time of the CTI of bird and other climatic variables such as water availability. butterfly communities and of European temperatures This new indicator, initially developed using Euro- 5 Hickling, R., Roy, D. B., Hill, J. K., Fox, R. & in the period 1990–2008. The temperature anomalies pean bird monitoring data13, calculates the diver- Thomas, C. D., 2006. “The distributions of a have been calculated using differences from the gence in population trends between (a) species that wide range of taxonomic groups are expanding 1961–1990 average (according to ref. 11). should benefit from and (b) species that should be polewards”. Global Change Biol., 12: 450–455. negatively affected by climate change according to Fig. 3. Trends in the CTI of the bird and butterfly the predictions of the so-called ‘climate envelope’ 6 Parmesan, C., Ryrholm, N., Stefanescu, C., Hill, communities in the different countries used in the models. A positive tendency in the indicator means J.K., Thomas, C.D., Descimon, H., Huntley, B., study. In all cases except for Catalonia tendencies that the populations of the species benefitting from Kaila, L., Kullberg, J., Tammaru, T., Tennent, W.J., are significantly positive. climate change – the ‘winner’ species – will have Thomas, J.A. & Warren, M.S., 1999. “Poleward increased over time more than those of the ‘loser’ shifts in geographical ranges of butterfly species Fig. 4. Climate change indicator for Catalan butterfly species. The climate envelope models explain the cu- associated with regional warming”. Nature, 399: populations calculated using the method of Gregory rrent distribution of species on the basis of a combi- 579-583. et al. (2009). Thin continuous line: trends in species nation of just a few variables that define the principal that, according to climate envelope models, will abiotic limitations that organisms have to confront 7 Stefanescu, C., Peñuelas, J. & Filella, I., 2003. increase their distribution area; dashed line: species (e.g. minimum winter temperatures, available ener- “Effects of climatic change on the phenology of whose ranges will shrink; thick line: Climate Change gy during the growing season and water availability), butterflies in the northwest Mediterranean Basin”. Indicator calculated as the ratio between the and project future distributions bearing in mind the Global Change Biol., 9: 1494-1506. two previous groups. The trend of the indicator is values for these variables projected under general cir- significantly positive for the period 1994–2011. culation climatic models (GCM). 8 Both, C., Bouwhuis, S., Lessells, C.M. & Visser, When the climatic indicator of Gregory et al. M.E., 2006. “Climate change and population Photo 1. The Provençal Short-tailed Blue Cupido (2009) is calculated for Catalan bird populations declines in a long-distance migratory bird”. Nature, alcetas lives in scrub and grassland in the northern using the SOCC dataset, a significant increase in 441: 81-83. half of Catalonia, with its densest populations found the period 2002–2011 occurs that confirms the in La Garrotxa. Climate envelope models predict expected response of Catalan bird communities to 9 Saino, N., Ambrosini, R., Rubolini, D., von that its area of distribution will expand under a climatic change. If the same procedure is applied to Hardenberg, J. Von, Provenzale,A., Hüppop, K., scenario of climate change, a possibility that could the CBMS dataset, using the predictions of changes Hüppop, O., Lehikoinen, A., Lehikoinen, E., Rainio, be heightened by this species’ known ability to in distribution provided by the climatic risk atlas of K., Romano, M. & Sokolov, L., 2011. “Climate disperse. In fact, the Provençal Short-tailed Blue is European butterflies14, a significant increase in the warming, ecological mismatch at arrival and one of the few species that has become commoner indicator for the period 1994–2011 also becomes population decline in migratory birds”. Proc. R. Soc. in Catalonia in recent years and, in doing so, has apparent (fig. 4). B, 278: 835-842. confirmed the predictions of climate models (photo: In contrast to the CTI, this new indicator mat- J. Jubany). ches predictions under a scenario of climate change. 10 Devictor, V., Julliard, R., Couvet, D. & Jiguet, F., We believe that this difference is due to the fact that 2008. “Birds are tracking climate warming, but not the models used by this second indicator to predict fast enough”. Proc. R. Soc. B., 275: 2743-2748. trends in species take into account both tempera- The CBMS at Alzinar de Sant Martí ture and precipitation. The combination of these 11 Devictor, V., van Swaay, C., Brereton, T., Brotons, (Sallent, El Bages) two variables reflects water availability (or degree L., Chamberlain, D., Heliölä, J., Herrando, S., of aridity), which is the key to understanding how Julliard, R., Kuussaari, M., Lindström, A., Reif , J., Located in a lowland Mediterranean area Mediterranean ecosystems work. Thus, temperature Roy, D.B., Schweiger, O., Settele, J., Stefanescu, on calcareous soils, this butterfly walk, acts as a limiting factor for many taxonomic groups C., van Strien, A. van, Turnhout, C., Vermouzek, active since 2000, runs through a growing (including butterflies) in ecosystems at higher lati- Z., DeVries, M.W., Wynhoff, I. & Jiguet, F., 2012. holm oak forest (alzinar) and areas of tudes in more northerly areas of Europe, whereas in “Differences in the climatic debts of birds and mixed scrub. It is a very diverse area and Mediterranean systems aridity is the principal limi- butterflies at a continental scale”. Nature Clim. to date 86 species have been recorded. The ting factor15. Change, 2: 121-124. population of Striped Grayling Hipparchia All climate change models agree that recurring fidiais the densest in the whole of the CBMS droughts and heat-waves in the Mediterranean will 12 Stefanescu, C., Torre, I, Jubany, J. & Páramo, F., network and other Satyrinae species such as have an increasingly greater impact16. Climate enve- 2011. “Recent trends in butterfly populations Dusky Heath Coenonympha dorus and Wall lope models that take into account water availability from north-east Spain and Andorra in the light of Brown Lasiommata megera are also very to explain the distribution of European butterflies habitat and climate change”. J. Conserv., abundant. Of the rarer species, it is worth predict that the species with the highest STIs, that 15: 83-93. noting the presence of Spanish Fritillary is, the most typical Mediterranean species, will have Euphydryas desfontainii and Sage Skipper their distribution areas reduced most of all. This pre- 13 Gregory, R.D., Willis, S.G., Jiguet, F., Vorisek, Muschampia proto. Although the diversity of diction theoretically leads to a fall in the CTIs in the P., Klvanová, A., van Strien, A., Huntley, B., species seems to have increased slightly in areas in which these Mediterranean species domina- Collingham, Y.C., Couvet, D. & Green, R.E., 2009. recent years, overall the abundance of the te and, at the same time, an increase in the climate “An indicator of the impact of climatic change commonest species has fallen. indicator of Gregory et al. (2009). Both predictions on European bird populations”. PLoS ONE, 4(3): are confirmed by the data from the CBMS network. e4678. The transect Amongst the challenges facing us in the near futu- The butterflies counts at Alzinar de Sant Martí near re is that of understanding how climate change and 14 Settele, J., Kudrna, O., Harpke, A., Kühn, I., van the town of Sallent began in 2000 and were the first changes in land use combine to produce a generali- Swaay, C., Verovnik, R., Warren, M., Wiemers, to be carried out in the comarca of Bages. Today, zed fall in butterfly numbers in Catalonia. Once this M., Hanspach, J., Hickler, T., Kühn, E., van however, along the valley of the river Llobregat there conundrum is solved, we must then design efficient Halder, I., Veling, K., Vliegenthart, A., Winhoff, are a good number of stations. The area chosen for proposals to minimize as much as possible this loss I. & Schweiger, O., 2008. “Climatic risk atlas of the walk consists of a broad plateau lying at 520 m of diversity that we are already beginning to suffer. European butterflies”. Pensoft, Sofia-Moscow. a.s.l. where arable land and woodland alternate with the scrub that has developed in areas of abandoned Constantí Stefanescu 15 Stefanescu, C., Carnicer, J. & Peñuelas, J., 2011. agricultural land and areas burnt by the 1986 forest “Determinants of species richness in generalist and fire. Nevertheless, this fire left some areas of the cha- specialist Mediterranean butterfly: the negative racteristic holm oak forest intact and today this forest 1 Stefanescu, C., 2008. “L’indicador europeu de les synergistic forces of climate and habitat change”. is well established and in expansion despite the relati- papallones de prats”. Cynthia, 8: 10-12. Ecography, 34: 353-363. vely small surface area that it covers. Cynthia V

Changes in land use and its News The walk passes through three main types of ha- effects on the butterflies bitat: open scrub (sections 1 and 4) with the grasses Proposal for common names for Brachypodium phoenicoides and B. retusum and blue No important changes have occurred in the land use Catalan butterflies aphyllanthes (Aphyllanthes monspeliensis); woodland of the area during the years the butterfly walk has in sections 2, 3, 5 and 7, with both evergreen holm operated: a few tracks have been repaired and clea- f you flick through some of the back numbers of oak and Aleppo pine (Pinus halepensis) and deci- red, but no animals graze here and the forests are not I the journal Cynthia you will see that the articles duous mixed woodland; and the scrub in section exploited for their timber. No generalised or inten- it contains talk of ‘blavetes’, ‘papallona de l’arboç’ 6, rich in Mediterranean species. Some of the most sive efforts had ever been made to clear the under- or ‘papallona del lledoner’. These are the common important plant species for butterflies either as food growth or the lower branches trees of the forests until vernacular names that are used by many butterfly plants or nectar sources that are common along the the beginning of the 2012 counting season. Without watchers and counters, as well as by other amateur walk include Dorycnium pentaphyllum, bramble doubt it will be interesting to analyse the changes naturalists from other fields. Interest in butterflies as (Rubus ulmifolius), sloe (Prunus spinosa), hawthorn that may occur as a result in relation to other similar a group has grown in recent years, as the number of (Crataegus monogyna), Mediterranean buckthorn stations as a means of evaluting whether the impact CBMS counters testifies. More and more books on (Rhamnus alaternus), wild privet (Ligustrum vulgare), of these type of changes are positive or negative. wildlife – including butterflies – are being published strawberry-tree (Arbutus unedo), Etruscan honeysuc- Initially, an unusual abundance of Marsh Friti- and natural history-related subjects have become kle (Lonicera etrusca), horseshoe vetch (Hippocrepis llary was detected in section 2, possibly the result of commonplace in the mass media in recent years. For comosa), Cephalaria leucantha, a sainfoin Onobrychis the clearance work that could have forced butterflies example, field guides, guides to protected areas and supina and Montpellier milk-vetch (Astragalus mons- to leave section 6 and head elsewhere. On the other local wildlife walks providing information on local pessulanus). hand, in 2012 for the first time in 13 years no Spa- flora and fauna have proliferated and an interest in nish Festoons were counted, although, fortunately, local natural history has become a popular past-time The butterflies the species’ foodplant, Aristolochia pistolochia, were and is no longer seen as the sole preserve of scientists In all, 86 species of butterfly have been detected found to be playing host as per usual to the caterpi- or of impassioned amateurs. along the walk. The annual average is almost 64 spe- llars of this species. Thus, knowledge of the common names of our cies, with a minimum of 59 in 2006 and a maximum The overall data from the 13 years of butterfly animals and plants will become not only important of 69 in 2009. Of the 10 most abundant species at counts reveal that the diversity of the area has in- but necessary, and will complement – and in some Sallent (fig. 1), six are among the commonest 20 spe- creased whilst abundances have dropped notably. A cases, even substitute – the use of scientific names. cies in the whole of the CBMS network (as of 2010). section-by-section study of the data in terms of both This is almost the case already for vertebrates, and in The exceptions are Dusky Heath (Coenonympha absolute and relative abundance and species richness other languages such as English an established body dorus), Striped Grayling (Hipparchia fidia), Marsh show that the most wooded sections and sections of common names for most species has existed for Fritillary (Euphydryas aurinia) and Berger’s Clouded with scrub on dry stoney soil have remained stable many years1. Yellow (Colias alfacariensis), which all have sizeable (sections 3, 4 and 5) over time, although their overall This situation has led to the need for a body of populations in this itinerary. The most notable case is production is relatively low. Sections on which the common names in Catalan to be developed for our that of Striped Grayling, a common and widespread forest grows more in height than in density (which butterflies. A group of experts from the Granollers species in lowland Mediterranean areas that at Alzi- was already there) have clearly suffered over the years Natural History Museum and the Catalan Lepi- nar de Sant Martí reaches its greatest densities in the (section 2), whilst, finally, in sections with scattered dopterological Society has started the ball rolling by whole of the CBMS network. Likewise, the Marsh trees on more humid soils the tendencies are more drafting a list of proposed common names for our Fritllary can be highly abundant in certain years and, encouraging (very positive in section 1, less so in butterflies, which it is hoped will be accepted by tho- as in the case of the Striped Grayling, its patterns section 6). se who have an interest in using them. The adjoining of annual variation closely follow those occurring list contains proposals for all the butterflies found elsewhere in Catalonia. The Alzinar de Sant Martí station in Catalonia and those wishing to learn more about This itinerary has an excellent selection of the lar- in relation to other butterfly studies how these names were chosen should consult the ori- ge summer Satyrinae species: aside from the Striped My interest in butterflies and desire to learn more ginal article in which these names were presented2. Grayling there are also good populations of Wood- about the biology of this group of insects has led me A number of different criteria were used to es- land (Hipparchia fagi), Tree (Hipparchia statilinus) to more detailed field studies of the itinerary at Alzi- tablish the list of names. Initially, the few existing and Great Banded (Brintesia circe) Graylings, com- nar de Sant Martí and in neighbouring areas (e.g. El traditional names, mainly collated by Vilarrúbia3, moner here than in many of the other CBMS sta- Guix, nearer the town of Sallent). I worked for many were all accepted. Wherever possible the new names tions. Of the abundant CBMS species, here too Wall years on a study of the parasitism of the two Euphy- were adapted to the classical binomial nomenclature Brown (Lasiommata megera) and, above all, Dusky dryas species that fly in the area, for which I collected used by Linnaeus such that species in the same genus Heath are common. and bred hundreds of larvae to see which species of have the same basic name, subsequently qualified to This itinerary also boasts a couple of the jewels parasitoids would emerge. The results were specta- identify each species. In some cases, though, a single that are enough to make any entomologist happy. In cular and revealed the enormous amount of unseen name was considered to be sufficient. The catalaniza- a woodland clearing in section 2, there are always life there is behind the world of the butterflies1. As tion of the common names used in other European a number Spanish Festoons (Zerynthia rumina) and well, in autumn-winter of 2001/02 I monitored me- languages or even of the scientific names was another at the beginning of the season their eye-catching ticulously around 150 larvae of the Two-tailed Pasha of the methods employed. Other criteria used inclu- but cryptic coloration is a joy to behold. The other Charaxes jasius in a study of the causes of mortality de references to obvious morphological features of delight is a small population of Glanville Fritillary in this species. The data from the CBMS network the species or its choice of food plant. A further set (Melitaea cinxia) whose caterpillars are often easier to enabled us to show that extremely cold winter of names includes allusions to the species’ choice of detect than the adults. Every year in section 1 we find weather has a severe negative impact on this species 2. habitat or certain characteristic behaviour. the dense webs of their larvae on often rather stunted We hope that this list of names will help increase plants of ribwort plantain (). Also Josep Planes knowledge of and respect for our butterflies. of note are the Spanish Fritillary (Euphydryas desfon- tainii), a singular species that is found in few CBMS 1 Stefanescu, C., Planas, J. & Shaw, M.R., 2009. Francesc Vallhonrat stations and is relatively abundant in the area. Fina- “The parasitoid complex attacking coexisting lly, it is worth commenting on the regular presence Spanish populations of Euphydryas aurinia and 1 Tolman, T. & Lewington, R.. 1997. Butterflies of of Sage Skipper Muschampia proto, not only because Euphydryas desfontainii (Lepidoptera: , Britain and Europe. 320 pp. + 104 pls. Harper this skipper is uncommon in the CBMS network but Melitaeini)”. J. Nat. Hist., 43: 553-568. Collins Publishers, Londres. because at Sallent it reaches its most northerly point of its Catalan distribution. 2 Stefanescu, C. & Planes, J., 2003. “Com afecta 2 Arrizabalaga, A., Stefanescu, C., Vallhonrat, F., As occurs elsewhere, many of the total of 86 spe- el rigor de l’hivern les poblacions catalanes de Dantart, J., Vila, R., Jubany, J., Sesma, J.M., cies such as Small Heath (Coenonympha pamphilus), Charaxes jasius”. Butll. Soc. Cat. Lep., 91: 31-48. Viader, S. & Dinca, V., in press. “Proposta de noms Chapman’s Green Hairstreak (Callophrys avis), comuns per a les papallones diürnes (ropalòcers) Amanda’s Blue (Polyommatus amanda) and White- Aerial photo. The CBMS transect at Alzinar de Sant catalanes”. Butll. Soc. Cat. Lep., 103. letter Hairstreak (Satyrium w-album) have only ever Martí. Its total al length is 1676 m, and there are been detected on a couple of occasions. Nevertheless, seven sections with an average length of 239 m 3 Vilarrúbia, J., 1961. Els nostres insectes. Converses given the context of the local butterfly populations, (range: 106-388 m). de divulgació entomològica. Vol. 1: 1-118; vol. 2: it would be no surprise if species such as Chequered 1-108; vol. 3: 1-112. Col·lecció Popular Barcino, Blue (Scolitantides orion) turned up on the butterfly Fig. 1. Average abundance (average of the annual núms. 191, 192 i 195, Barcelona. walk. Other species such as Small Heath are com- indexes during the period 2000–2012) of the 15 the mon in the area but were only ever counted during commonest butterflies at the Alzinar de Sant Martí the first years of the counts. station.

VI Cynthia

The butterfly be invaded by scrub) and by the intensification of tion larvae that become males8. In general, females The Glanville Fritillary Melitaea agricultural practices. only copulate once, just after emerging, although cinxia, a model for the study of Although in other parts of Europe the caterpillars on occasions females with two spermatophores that metapopulation theory of this fritillary feed on both plantains and plants must have mated twice are found. such as spiked speedwell Veronica spicata3, in Catalo- Over the past two decades, the Glanville nia it would seem to be completely associated with Metapopulational structure Fritillary Melitaea cinxia has become one the former4. The Glanville Fritillary is a paradigm of a species that of the most studied of all animals in the lives in a metapopulation. Its distribution over space field of ecology. This butterfly lives in Biological cycle and phenology is not uniform but rather is fragmented into popu- metapopulations, that is, in populations The Glanville Fritillary is a typical univoltine species lations that occupy well-defined patches of favoura- interconnected by the dispersion of adults. whose larvae hibernate during their fourth instar. ble habitat (in this cases, grassland with plantains). In Catalonia it is relatively common in open Its flight period lasts from April to July, although, These populations exchange individuals via dispersal areas with plantains in uplands, above exceptionally, its populations are bivoltine and the processes and the set of these interconnected popu- all in the northern half of the country. first generation in April-May is followed by a second lations is known as a metapopulation6. Populations In spring, its eye-catching caterpillars – in July-August in the areas of L’Alta Garrotxa and may be somewhat unstable and may often die out black with white-spots, and red feet and Alt Empordà. This behaviour, possibly genetic in ori- due to various causes. Nevertheless, population ex- heads – are easily observed as they feed on gin5, is very localized and is well-documented from tinctions are compensated for by the establishment plantains. the CBMS stations at Sales de Llierca, Darnius and of new populations in unoccupied patches by in- Els Aiguamolls de l’Empordà. Figure 3 shows the dividuals dispersing from other populations. The Geographical distribution flight curves from various stations in the CBMS net- fraction of the habitat patches that are occupied and situation in the CBMS work with differing environmental conditions. In El by a metapopulation depends above all on the size The Glanville Fritillary lives over much of Europe, Montseny and the mountains of La Serralada Trans- of the patches and the distance between them. As North Africa (Atlas Mountains in Morocco to Al- versal the first individuals appear in April, numbers a rule, the larger the patch, the less likely it is that geria), Russia and western Asia as far as the Amur peak in May-June and the generation lasts until the the population that occupies it will die out; and the region1. It is relatively common in the northern third beginning of summer (fig. 3a). In upland areas of smaller the distance between patches, the greater the of the Iberian Peninsula in a more or less continuous the Pyrenees, emergence takes place almost a month probability that individuals dispersing from another area running from the Pyrenees to the Cantabrian later but henceforth the flight curve is more synchro- population will compensate for any extinction that Mountains (including northern Portugal) and the nized. Maximum numbers are seen at the beginning may occur. Metapopulation theory has been greatly mountains of the Sistema Ibèric, as well as in isolated of June and by the second half of July few are still on developed over the past 20 years by Ilkka Hanski and lowland areas. In the centre of the Peninsula it flies the wing (fig. 3b). Figure 3c shows the bivoltine phe- his team on the basis of the Glanville Fritillary po- above all in upland areas (mountains of the Sistema nology at the stations of Darnius and Sales de Llier- pulations on the Åland islands in south-east Finland Central, Serra de Guadalupe and Serrania de Cuen- ca: the first generation flies earlier than in other loca- (fig. 4)11. ca) but becomes much scarcer in the south where it lities and is concentrated in the second half of April The metapopulation model can be generalized is confined to the eastern ranges of Cazorla, Espuña, and the first half of May, while the second is much to other animals (including many butterfly species) Sagra and Alfacar2. poorer in numbers and may in fact only be partial. and has very important implications for biodiversity It is widespread in the CBMS network and ap- The first butterflies of this second generation appear conservation12. It has been repeatedly shown that the pears in almost half of the stations, an accurate re- in July and numbers peak in July–August. After that, persistence of metapopulations is threatened when flection of its distribution in Catalonia (fig. 1). It is numbers fall very quickly and only a very few are changes in land-use lead to the fragmentation and found above all in the northern half of the country ever seen in September. At L’Estany de Sils (in what reduction in the size of habitats. When this occurs, and is common in the Pyrenees (including Andorra), appears to be an isolated population), although the habitat patches shrink and become more isolated, pre-Pyrenees and the mountains of the Serralada first butterflies appear likewise very early in the year, thereby increasing the risk of local extinction13,14. Transversal, Montseny and Sant Llorenç del Munt. there is no second generation, possibly because the Beyond a certain threshold, the number of occupied It is much rarer and somewhat local in the Empor- environmental conditions are unfavourable at such patches becomes insufficient and the metapopula- dà and Girona plains and the northern part of the low altitude due to the summer drought. tion structure collapses, thereby causing the comple- coastal mountains (Les Gavarres and El Montnegre). This species has been studied to an extraordinary te disappearance of the species in the area where it It becomes a rare species further south where local degree in recent decades owing to its role as a model was once present. populations are only known from La Mola de Coll- species in the development of metapopulation theory dejou (Baix Camp) and the mountains of Prades and (fig. 4)6,7. Thus, we have an unmatched knowledge of Natural enemies Els Ports de Tortosa-Beseït (although it has not yet its biology. As in other Melitaeini species, eggs are Parasitoids are especially important natural enemies appeared in the counts from the latter area). Of great laid in dense batches (between 25 and 250 at a time) of the Glanville Fritillary and have been studied interest are the occasional records from the CBMS on the underside of the leaves of its food plant. It has in great detail in Finland15. The most typical such stations at Granja d’Escarp and Sebes, two stations been calculated that a female can lay up to seven egg- parasitoids are two specialist icneumonids, Cotesia in arid areas that would seem to be unsuitable for batches during her life time, with progressively fewer melitaearum and Hyposoter horticola, that attack the the species. Nevertheless, relict populations of this eggs each time8. Eggs are laid preferably on small caterpillars, a variety of generalist chalcididoids that fritillary must survive in these areas of south-western plantains in hot microclimates, often in areas of bare attack the chrysalis (above all, Pteromalus apum and Catalonia. ground. Females also tend to select plants with high P. puparum), and various species of secondary para- In Catalonia this species has an extensive altitu- concentrations of iridoids, a secondary metaboli- sitoids belonging to the genera Gelis and Mesochorus dinal distribution that ranges from sea-level (occa- te in the plantain that accelerates the development that attack the primary specialist parasitoids. The si- sional records from the Aiguamolls de l’Empordà) of the larvae and reduces the impact of its specific tuation seems to be similar in Catalonia, where the to subalpine and even alpine habitats in Andorra, parasitoid Cotesia melitaearum9,10. Caterpillars hatch two primary specialist parasitoids have been detec- from where it has been recorded in the Sorteny and within 2–4 weeks and immediately build a silken ted16. The impact of these parasitoids on Glanville Pessons itineraries. Good population densities with web on the food plant. Once they have moulted out Fritillary populations has been studied on numerous annual indices of 20–50 occur in montane areas (e.g. of their fourth instar (mid-summer), the caterpillars occasions (see reviews in refs 17, 18) and continue to in El Montseny at 700–1,110 m a.s.l.; fig. 2a), but enter into and hibernation, which will last be one of the main lines of research in the metapo- also in lowland areas (e.g. Alta Garrotxa below 300 the whole winter, in a denser web spun at the base of pulation system on the Åland islands. Of particular m a.s.l., fig. 2b) and high mountain areas (e.g. Sant the host plant. Although they are hard to detect in interest is the case of H. horticola, which parasites Maurici at 1,600 m a.s.l.). Nevertheless, in almost a Catalonia, these winter webs are much more visible the host larvae when they are still in the egg (photo third of the stations in which the species has been in northern Europe and are often used in population a). To do so, the females of this icneumonid search detected its densities are extremely low (annual in- surveys. In March–April the larvae finish hibernating for the butterfly egg batches, memorize their posi- dices below 1) and very possibly these individuals (photo b), abandon their winter webs and enter a tion and revisit them periodically until the larvae are correspond to butterflies in dispersion from nearby solitary phase. They are easily seen when warming about to hatch. Normally this icneumonid only pa- populations. themselves on dried up vegetation (photo c). Due to rasitizes a third of the eggs in each batch, but never- their dark colour they are able to increase their body theless is still able to locate almost all the egg batches Habitats and food plants heat substantially and thus speed up their growth ra- in a population. The result is that this parasitoid kills The Glanville Fritillary lives in open areas, above all tes. Once the sixth instar is complete, the caterpillar a more or less fixed proportion – around a third – in grasslands with short swards and xeric in character, pupates. The chrysalis remains hidden suspended in of the Glanville Fritillary populations19. Aside from but also in more ruderal habitats (along tracks and the vegetation (photo d) for around 1–3 weeks until these parasitoids, certain carnivorous insects feed op- paths) wherever there is an abundance of its main the adult butterfly finally hatches. portunistically on this fritillary’s eggs (e.g. lacewings, food plant, ribwort plantain Plantago lanceolata. As When searching for mates, males combine patro- ants and ladybirds) and caterpillars (e.g. bugs)17. such it is totally absent from areas of intensive arable lling and territorial behaviour. As in the other Meli- land and thick forests. It is thus a specialist species taeini species there is a certain degree of proterandry Population trends that has been greatly affected by both the abando- (males emerge before females), which is explained by Due to its metapopulation structure, the Glanville ning of extensive grazing (which allows habitats to the shorter development time in the post-hiberna- Fritillary is particularly sensitive to landscape chan- Cynthia VII

ges, above all if they lead to a reduction in the open 7 Ehrlich, P.R. & Hanski, I. (eds), 2004. On the wings parasitoid of the butterflyMelitaea cinxia”. Oikos, areas where populations are located. Given that the of checkerspots. A model system for population 78: 91-100. loss of these open spaces is one of the main causes biology. Oxford University Press, Oxford. of the overall regression in Catalan butterfly popula- 22 Singer, M.C. & Parmesan, C., 2010. “Phenological tions in recent decades20, a fall in Glanville Fritillary 8 Boggs, C.L. & Nieminen, M., 2004. “Checkerspot asynchrony between herbivorous insects and their populations in Catalonia is also to be expected. reproductive biology”. In: On the wings of hosts: signal of climate change or pre-existing CBMS data indicate that numbers in most of checkerspots. A model system for population biology adaptive strategy?” Phil. Trans. R. Soc. B, 365: the monitored populations fell sharply in the period (Ehrlich, P.R. & Hanski, I., eds). Oxford University 3161-3176. 1994–2008. Subsequently, the species has recovered Press, Oxford, pp. 92-111. somewhat and in 2011 its tendency was classified Fig. 1. Relative abundance (expressed as the values by the TRIM programme as ‘uncertain’ (fig. 5). The 9 Nieminen, M., Suomi, J., van Nouhuys, S., Sauri, P. of the annual index/100 m) of the Glanville Fritillary recovery between 2009 and 2011 could be due to & Riekkola, M.-L., 2003. “Effect of iridoid glycoside Melitaea cinxia in different CBMS sites (1994–2011). a certain relaxation in density-dependent mortality content on oviposition host plant choice and factors such as parasitization by Cotesia melitaearum parasitism in a specialist herbivore”. J. Chem. Ecol., Fig. 2. Two of the typical habitats of the Glanville (which generally only has serious effects when host 29: 823-844. Fritillary in Catalonia. (a) Grassland with annual densities are high)21. Nevertheless, this recuperation plants in an upland holm oak forest at 750 m a.s.l. could also be related to favourable climatic condi- 10 Harvey, J.A., van Nouhuys, S. & Biere, A., 2005. on the Santa Susanna itinerary in El Montseny tions and a series of wet springs and summers could “Effects of quantitative variation in allelochemicals Natural Park, and (b) open habitat, glades and rides have favoured larval survival just before and after in Plantago lanceolata on development of a within evergreen oak forest, in Sales de Llierca diapause. The importance of this latter factor should generalist and a specialist herbivore and their itinerary (photos: a, M. Miralles; b, M. Lockwood). not be underestimated in this species given that va- endoparasitoids”. J. Chem. Ecol., 31: 287-302. rious studies have shown that species with similar Fig. 3. Phenology of the Glanville Fritillary in ecologies are seriously affected when the synchroni- 11 Nieminen, M., Siljander, M. & Hanski, I., 2004. different habitats in Catalonia. (a) Populations in zation between larval and food-plant development “Structure and dynamics of Melitaea cinxia the mountains of El Montseny (itineraries at Santa periods is altered22. The fact that most Glanville metapopulations”. In: On the wings of checkerspots. Susanna, El Puig and Pla de la Calma) and in the Fritillary eggs are laid on plantains in hot microcli- A model system for population biology (Ehrlich, P.R. mountains of La Serralada Transversal (itinerary at mates, which are thus prone to withering, makes this & Hanski, I., eds). Oxford University Press, Oxford, Sant Feliu de Pallerols) at heights of 350–1,100 hypothesis even more plausible. pp. 63-91. m a.s.l. (n = 2,079 individuals, 1994–2011); (b) Come what may, the CBMS data reveal the pos- population at Sant Maurici in a subalpine habitat at sibility of a general decline in this species in Catalo- 12 Hanski, I. & Thomas, C.D., 1994. “Metapopulation 1,600 m a.s.l. (n = 342 individuals, 2006–2011); (c) nia (fig. 5), which would be most evident in isolated dynamics and conservation: a spatially explicit bivoltine populations at Darnius and Sales de Llierca populations and in those situated in areas with the model applied to butterflies”.Biol. Conserv., 68: (n = 1,057 individuals, 1994–2011). most extreme Mediterranean climate. Examples of 167-180. such threatened populations include those at Sils, Fig. 4. The Glanville Fritillary is one of the best those in the mountains in Tarragona and those that 13 Kuussaari, M., Saccheri, I., Camara, M. & Hanski, studied butterflies in the world and has been used are scattered in the western comarques of Segrià and I., 1998. “Allee effect and population dynamics as a model for the development of metapopulation Ribera d’Ebre, whose viability may well be put to test in the Glanville fritillary butterfly”. Oikos, 82: theory. In the books by I. Hanski and P. R. Ehrlich in the mid-term under a scenario of climate change 384-392. the results of this ecological research into this and the growing impact of recurring droughts. species’ metapopulation structure, along with other 14 Saccheri, I., Kuussaari, M., Kankare, M., Vilkman, aspects of its life history, are explained in detail. Constantí Stefanescu P., Fortelius, W. & Hanski, I., 1998. “Inbreeding and extinction in a butterfly metapopulation”. Fig. 5. Population fluctuations in the Glanville Nature, 392: 491-494. Fritillary in the CBMS network in the period 1 Tolman, T. & Lewington, R., 2002. Guía de las 1994–2011 calculated with the programme TRIM. mariposas de España y Europa. 320 pp. + 104 pl. 15 Lei, G.C., Vikberg, V., Nieminen, M. & Kuussaari, Lynx Edicions, Bellaterra. M., 1997. “The parasitoid complex attacking Photos. (a) Egg-batch on a ribwort plantain Finnish populations of the Glanville fritillary Plantago lanceolata being inspected by the 2 García-Barros, E., Munguira, M. L., Martín Cano, J., Melitaea cinxia (Lep.: Nymphalidae), an icneumonid parasitoid Hyposoter horticola; (b) Romo Benito, H., Garcia-Pereira, P. & Maravalhas, endangered butterfly”. J. Nat. Hist., 31: 635-648. group of caterpillars leaving their winter web in the E. S., 2004. “Atlas de las mariposas diurnas de la Pyrenees; in warmer parts of Catalonia the species Península Ibérica e islas Baleares (Lepidoptera: 16 Kankare, M., Stefanescu, C., van Nouhuys, S. & apparently does not spin such conspicuous winter Papilionoidea & Hesperioidea) ”. Monografías Soc. Shaw, M.R., 2005. “Host specialization by Cotesia webs; (c) caterpillar in its sixth and last instar ent. aragon., 11: 1-228. wasps (Hymenoptera: Braconidae) parasitizing warming itself on a dry leaf; (d) chrysalis; (e, f) species-rich Melitaeini (Lepidoptera: Nymphalidae) adults showing the characteristic black spots on 3 Kuussaari, M., Singer, M. & Hanski, I., 2000. “Local communities in north-eastern Spain”. Biol. J. Linn. the submarginal band of the hind-wing, visible on specialization and landscape-level influence on Soc., 86: 45-65. both the upper- and under-wings. (photos: a, S. Van host use in an herbivorous insect”. Ecology, 81: Nouhuys; b, J. Piqué; c, e, J.M. Sesma; d, J.R. Salas; 2177-2187. 17 van Nouhuys, S. & Hanski, I., 2004. “Natural f, J. Jubany). enemies of checkerspot butterflies”. In:On 4 The numerous observations of egg-laying on the wings of checkerspots: A model system for ribwort plantain Plantago lanceolata, as well as population biology (Ehrlich P.R. & Hanski I., eds). post-hibernation caterpillars feeding on plantain Oxford University Press, Oxford, pp. 161-180. Identification in various parts of the Pyrenees, mountains of La Serralada Transversal and El Montseny, indicate 18 Shaw, M.R., Stefanescu, C. & van Nouhuys, How to separate the species that this plant is the main food plant for most S., 2009. “Parasitism of European butterflies of the genus Erebia (4) populations. Nonetheless, a solitary observation of (Hesperioidea and Papilionoidea)”. In: Ecology of Once we reach the heights of the Pyrenean a female egg-laying on spiked speedwell Veronica butterflies in Europe (Settele, J., Shreeve, T.G., peaks and ridges, the commonest ringlets spicata on the Puig itinerary in El Montseny Konvicka, M. & Van Dyck, H., eds). Cambridge (Erebia spp.) are usually Mountain (E. suggests that locally this plant may be used on University Press, pp. 130-156. epiphron), Gavarnie (E. gorgone) and, above occasions, as occurs in northern Europe. all, Silky (E. gorge) and Lefèbvre’s (E. 19 van Nohuys, S. & Ehrnsten, J., 2004. “Wasp lefebvrei) Ringlets. The description of these 5 In breeding experiments under similar behavior leads to uniform parasitism of a host four species completes the revision of the environmental conditions, the bivoltine available only a few hours per year”. Behav. Ecol., Catalan ringlets. populations from L’Alta Garrotxa and L’Alt Empordà 15: 661-665. maintain their marked tendency to produce a f the species that fly at great altitude, only second generation in the laboratory. In the other 20 Stefanescu, C., Torre, I, Jubany, J. & Páramo, F. OMountain (E. epiphron), Gavarnie (E. gorgone) univoltine Catalan populations this phenomenon 2011. “Recent trends in butterfly populations and Lefèbvre’s (E. lefebvrei) Ringlets have ever been is highly exceptional (but not unheard of) (M.C from north-east Spain and Andorra in the light of detected in the CBMS-BMSAnd network ­– and, at Singer, com. pers.). habitat and climate change”. J. Insect Conserv., that, only very sporadically. Of the three, Mountain 15: 83-93. Ringlet has appeared most often in the butterfly cou- 6 Hanski, I., 1999. Metapopulation ecology. Oxford nts due to its wide altitudinal range (1,400 to 3,000 University Press, Oxford. 21 Lei, G.-C. & Hanski, I., 1997. “Metapopulation m a.s.l.); on the other hand, Gavarnie Ringlet only structure of Cotesia melitaearum, a specialist flies above 1,850 m a.s.l. and Lefebvre’s is confined VIII Cynthia

to true alpine habitats above 2,400 m a.s.l., where it LEFÈBVRE’S RINGLET The males of all three species are territorial and often often flies with Silky Ringlet E.( gorge). These four Upperside (general): black, females generally with await females perched on trees or rocks. Their larvae species have similar ranges in the Catalan Pyrenees, post-discal band, but males generally without have been reported to feed on a variety of grasses1 from La Val d’Aran, El Pallars Jussà, El Pallars Sobirà Underside (general): under- and upper-sides (see ref. 2-3 for more details on their life-cycles). through L’Alt Urgell and La Cerdanya as far as El are similar in males, but in females underside is Ripollès, where the easternmost populations of these brownish Constantí Stefanescu species are found1. Gavarnie Ringlet, endemic to the Marked with line: Pyrenees, and Mountain Ringlet fly in subalpine and Post-discal band absent; black eye-spots with white 1 Tolman, T. & Lewington, R., 2002. Guía de las alpine grasslands, while Silky and Lefèbvre’s Ringlets pupils directly on background; often with a rusty mariposas de España y Europa. 320 pp. + 104 pl. are commoner on screes and rocky slopes. All four red post-discal band and black pupiled eye-spots; Lynx Edicions, Bellaterra. species are univoltine, with population peaks in July brownish tones; hind-wing with paler post-discal and August; Mountain Ringlet is the first to fly (at area 2 García-Barros, E., 2000. “Comparative data on the the end of June), while the last to fly is Gavarnie Rin- adult biology, ecology and behaviour of species glet, which is sometimes found at the end of August. These species are easy to separate, although both belonging to the genera Hipparchia, Chazara The biological cycle of Gavarnie Ringlet is unknown, Mountain and Lefèbvre’s Ringlets and, to a lesser and Kanetisa in Central Spain (Nymphalidae: whilst the other three are known to have bi-annual extent, Silky Ringlet have considerable individual Satyrinae)”. Nota lepid., 23: 119-140. cycles due to the rigorous climate of the areas they variation and marked sexual dimorphism. Mountain inhabit2. In terms of food plants, the larvae of Mou- Ringlet is the smallest and is easily recognisable 3 Möllenbeck, V., Hermann, G. & Fartmann, T., 2010. ntain Ringlet feed on Festuca, Poa or Deschampsia, by the pointed apex to its fore-wing and its small “Larval ecology and habitat requirements of the those of Silky Ringlet on Festuca or Poa and those of black eyes-spots that lack any white pupils. In Silky woodland grayling Hipparchia fagi – basics for Lefèbvre’s on Festuca3. Ringlet the broad reddish post-discal bands and the the evaluation of fire management”. Arb. Inst. Jordi Dantart series of four large white-pupiled eye-spots on the Landschaftsökol., Münster, 19: 77-152. hind-wing are distinctive. Gavarnie Ringlet has broad 1 García-Barros, E., Munguira, M. L., Martín Cano, J., reddish post-discal bands that do not stand out well 4 Lafranchis, T., 2000. Les papillons de jour de France, Romo Benito, H., Garcia Pereira, P. & Maravalhas, against the background, while the underside of the Belgique et Luxembourg et leurs chenilles. 448 pp. E. S., 2004. “Atlas de las mariposas diurnas de la hind-wings have a characteristic marbled texture Collection Parthénope, éditions Biotope, Mèze. Península Ibèrica e islas Baleares (Lepidoptera: against which the veins and pale post-discal bands Papilionoidea & Hesperioidea)”. Monografías Soc. are obvious. Lefèbvre’s Ringlet is the blackest of Drawings ent. aragon., 11: 1-228. the Pyrenean ringlets; in addition, in males often only the white pupil of the black eye-spots is visible 2 Ligue Suisse pour la Protection de la Nature, 1987. against the dark background. WOODLAND GRAYLING Upperside (general): dark brown, almost black with Les papillons de jour et leurs biotopes. xi + 512 pp. white postdiscal band

3 Underside (general): similar to upperside Tolman, T. & Lewington, R., 2002. Guía de las Marked with line: mariposas de España y Europa. 320 pp. + 104 pl. Identification Androconial scales with velvety appearance; grey- Lynx Edicions, Bellaterra. How to separate the species white band; more regularly curved white band of the genus Hipparchia (2) Drawings ROCK GRAYLING Two species of the genus Hipparchia, Upperside (general): dark brown, almost black with MOUNTAIN RINGLET Woodland (H. fagi) and Rock (H. hermione) white post-discal band Upperside (general): very dark brown or black, matt Graylings, are found in Catalonia. Their Underside (general): similar to upperside or with the odd metallic reflection similarity is so great that to separate the two Marked with line: Underside (general): similar to the upperside, but species it is essential to examine external Androconial scales with velvety appearance; with paler, greyer or browner tones genital characteristics. Nevertheless, their grey-white band; curved white band with a marked Marked with line: ecological requirements are quite different. dog-leg Quite pointed apex; postdiscal golden-brown band Superficially, they also resemble the Great interrupted by dark veins and blind black eye-spots; Banded Grayling (Brintesia circe) and GREAT BANDED GRAYLING Upperside (general): black, with a white post- series of round golden patches with blind black eye- confusion between them is possible. discal band interrupted on the fore-wing spots; series of black spots directly on background Underside (general): similar to upperside he Woodland Grayling (Hipparchia fagi) inha- Marked with line: SILKY RINGLET Tbits Mediterranean woodlands and is modera- Androconial scales with velvety appearance; marks Upperside (general): brown in the basal, discal and tely common in open holm oak and pine woodland on post-discal band are pure white; additional white post-discal areas throughout much of Catalonia (but is absent from discal band Underside (general): fore-wing predominantly red- the Pyrenees). Its maximum numbers are recorded brown; hind-wing marbled grey and black from the mountains of the Serralada Prelitoral and All three species have broad white post-discal bands Marked with line: dry areas of the pre-Pryenees. On the other hand, on the upper-wings, although in both Woodland Broad postdiscal band, generally reddish and silky in the Rock Grayling (H. hermione, also known as H. and Rock Graylings this band is a smoky grey-white, appearance; two apical black eye-spots with white alcyone) is characteristic of open upland areas and almost yellow, in colour. The Great Banded Grayling pupils; reddish post-discal band with a series of four is well distributed throughout the Pyrenees and the can be separated from the other two species by pupiled black eye-spots; discal area darker; post- pre-Pyrenees, but is only local in La Serralada Preli- the short white band in the discal region on the discal area with a series of pupiled eye-spots toral. Its populations can reach important densities underside of the hind-wing. This feature is easy on peaks and ridges such as on El Turó de l’Home to appreciate when the butterfly is perched with its wings folded. Woodland and Rock Graylings are GAVARNIE RINGLET and Les Agudes in El Montseny. The Great Banded very similar and can be easily confused, although Upperside (general): very dark brown Brintesia circe Grayling ( ) is very common everywhere the former is slightly larger (length of fore-wing in Underside(general): fore-wing reddish mahogany in Catalonia, from sea level to subalpine forests. It Woodland Grayling = 33 mm, but in Rock Grayling brown; hind-wing marbled with a paler postdiscal prefers grassy areas near woodlands. None of these = 32 mm). The males can be reliably separated band three species are found on the Balearic Islands and without dissection by examination of the Jullien’s Marked with line: all three are univoltine. Both the Woodland and organ (see drawing in ref. 4). In Catalonia the two Mahogany postdiscal band that only stands out Rock Graylings appear in June and are commonest species only fly together in the pre-Pyrenees and in poorly from background; small black eye-spots with in August or even September. The Great Banded these sites it is recommendable to regularly confirm white pupils; marbled grey-brown background; pale Grayling, however, appears in May–June and has a the identification of males by an examination of the postdiscal band without obvious spots bimodal flight period since the majority of indivi- Jullien’s organ. duals aestivate in July–August. Eggs are laid from the end of summer through to the beginning of October.

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