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Geographical Variability In, and Temperature Effects On, the Phenology of Maniola Jurtina and Pyronia Tithonus (Lepidoptera, Satyrinae) in England and Wales

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Geographical Variability In, and Temperature Effects On, the Phenology of Maniola Jurtina and Pyronia Tithonus (Lepidoptera, Satyrinae) in England and Wales Ecological Entomology (1987) 12, 139-148 Geographical variability in, and temperature effects on, the phenology of Maniola jurtina and Pyronia tithonus (Lepidoptera, Satyrinae) in England and Wales PAUL M. BRAKEFIELD Department of Zoology. University College, Cardiff ABSTRACT. 1. Geographical variability in. and temperature effects on, the mean date of adult flight period and the SD about this date are analysed for two univoltine, grassland butterflies in England and Wales from 1976 to 1985. Data were collected on the Butterfly Monitoring Scheme for Maniola jurtina (L.) at twenty-nine sites and Pyronia tithonus (L.) at twenty sites. 2. Substantial variability for mean date and SD occurs between years and between sites. Changes in mean date between years tend to occur consistently at different sites. The species show some parallel in variation between sites, especially for mean date. 3. June maximum temperature accounts for 95% and 75% of the varia- tion in mean date between years in M. jurtina and P.tithonus. respectively (r=-0.97 and -0.87). Similar relationships occur for temperatures cumul- ated over the period of post-winter development from March to July or August. 4. Greater geographical variability in phenology, and a generally less synchronized flight period in M.jurtina may be associated with broader habitat preferences than in P. tithonus. 5. The mean date of adult flight period remains at roughly the same date at more northerly latitudes. In M.jurtina the flight period becomes more synchronized, begins later and ends earlier in the north. P. tithonus shows little or no indication of such a response to latitude. This is discussed with regard to changes in season length and factors limiting the species' range. Key words. Maniola jurtina, Pyronia tithonus, butterfly, phenology, geographical variability, temperature, development, emergence, distribu- tion, latitude. Introduction tribution of the British butterfly fauna (e.g. Dennis. 1977; Heath ft til.. 1984). Dennis There is renewed interest in factors which examined isotherms coinciding with the influence the historical and contemporary dis- northern limits of many species .ind showed that Correspondence: Dr Paul M Brakefield. Depart- a numher of ™ogeograph,cal and climatic fac- ment of Zoology. University College. P.O. Box 78. tors- including July temperatures, act .is good Cardiff CF1 1XL. predictors of differences between regions in 140 Paul M. Brakefield species diversity. More recent studies have con- timing of the adult flight period in a population centrated on analysing patterns in 10 km square of a North American butterfly, Papilio glaucus, distribution maps given by Heath et al. (1984). using data describing the degree-days above the Nearly 80% of the variation in species diversity developmental threshold required for complete is accounted for by summer temperature and development. The present study is an analysis of sunshine together (Turner, 1986) or by latitude the influence of temperature during post- (Barbour, 1986). Turner et al. (1986) showed diapause, pre-adult development on the timing that climate was able to explain some part of the and duration of the flight period of two strictly variation in diversity which is independent of univoltine, satyrine butterflies at grassland sites latitude. Barbour ( 1986) also found that patterns throughout England and Wales. The study of extinction since pre-1970 records were correl- species are the meadow brown Maniola jurtina ated with richness of the epiphytic flora and air (L.) and the gatekeeper Pyronia tithonus (L.). pollution levels. Dennis & Williams (1986) Geographical variability in their phenology is emphasized the many covarying factors and the also examined. The analysis was made possible necessity of taking into account the great by data recorded over the period 1976-85 on the variability in the biology of the species of Butterfly Monitoring Scheme (BMS) and butterfly. generously made available by Dr E. Pollard. Ecological studies of small numbers of populations of certain species of butterfly in Britain have indicated that temperature can influence their population dynamics (e.g. Methods Pollard, 1979a; Thomas, 1983; Warren et al., 1986; and see discussion in Thomas, 1984). The methods used in the BMS are fully docu- Pollard's population study and survey work on mented elsewhere (Pollard et al., 1975; Pollard, Ladoga Camilla strongly suggested that a con- 1977). It is based on standardized counts usually traction and subsequent expansion in its range made at least once a week along fixed transects was associated with changes in weather patterns at sites throughout Britain. Sites are visited from and, in particular, in June temperature. Unusual April until the end of September which covers weather frequently results in local extinctions or the whole flight period of P. tithonus and all but marked fluctuations in population size (Pollard, the extreme tail for M.jurtina in a small propor- 1984; see also Ehrlich étal.. 1980). The larvae of tion of its populations. Estimates of the expecta- certain species exhibit thermoregulatory tion of adult life in a meadow habitat near behaviour, absorbing radiant energy from the Liverpool in 1976 and 1977 were similar for each sun to gain some independence from air temp- species, varying from 3.5 to 8.5 days (Bra- erature (Porter, 1982, and see Dennis, 1985). kefield, 1979a, 1982a, b). Both species were also Such behaviour may enable populations to per- quite sedentary. Therefore, the probability of an sist in more northerly latitudes. Geographical individual insect being recorded along a transect variability in the number of annual generations on more than one date is likely to be similar for or changes in voltimsm between years has been M.jurtina and P. tithonus and although the BMS documented for several species in Britain (see counts describe their flight periods they should Heath et al., 1984; Dennis, 1985). Such also be related to adult emergence in a similar phenomena seem to be associated with variation way for each species. The sexes are not recorded in the length of time available for development separately in the BMS. Counts are made only ('season length', sensu Roff, 1980). Systematic when weather conditions meet specified mini- investigations of temperature effects on the mum criteria, for example for sunshine. In a few development rate of British butterflies and cases prominent gaps occurred in the records hence on the timing of emergence or phenology and these were filled with an estimate calculated of natural populations have not been carried out as the average of the single preceding and although such effects are ubiquitous in other following counts. The BMS was started in 1976 insects. An understanding of them is likely to with many more sites being added in 1977 and provide important insights into the constraints of fewer in later years (Pollard, 1984). Two sets of season length on the butterfly fauna. A study by sites are analysed here; those covering the peri- Hagen & Lederhouse (1985) has examined the ods of 1976-85 and of 1977-85 (including those Phcnologv of satvrine butterflies 141 this date (SD). These represent estimates of the mean date of adult flight period and of the degree of synchronization of the flight period at each individual site. In P.tithonus there is no relationship between the site means for SD and total annual counts (r=-0.14). In contrast, for M.jurtina this relationship is quite strong (r=0.61, df=27, P<0.001). However, the latter correlation is entirely due to the five sites with counts substantially higher than the others (excluding the five sites: r=—0.03). Therefore there is some indication of a non-linear relation- ship in M.jurtina but the results described below are unchanged when these five sites are excluded from the analyses. Both species overwinter as early to mid-mstar larvae exhibiting semi- (in M.jurtina) or wholly arrested development Regular night-time feed- ing on grasses commences during March. Larvae begin to pupate in late May (M.jurtina) or June. In both species nearly all adults have emerged by the end of August. Monthly mean daily max- FIG. 1. Map of the BMS sites (circles) in England and imum and minimum temperatures for the post- Wales used in this study. Map numbers refer to silc winter period of pre-adult development during names in Table 1. The large circle shows position of the five sites with the prefix A. Squares indicate March-August inclusive were abstracted from meteorological stations providing temperature data. the Monthly Weather Reports (H.M.S.O., 1976-85) for a representative series of twelve meteorological stations distributed over the region covered by the BMS sites (Fig. 1). recorded in 1976). This choice enables a good Cumulative daily temperatures were also calcul- coverage of England and Wales (Fig. 1) and ated for this period using the average of monthly yields the longest possible periods of continuous means for daily maxima and minima. The latter recording. It also provides a means of examining statistics were used in the absence of the necess- the effect of the exceptionally hot summer of ary data to calculate day-degrees. Statistical pro- 1976. Three of the sequences for individual sites cedures were performed using the MINITAB (two for M.jurtina and one for P.tithonus) package on the mainframe computer at Univer- included a year with no records. These sity College, Cardiff. sequences were completed by substituting the overall means for the appropriate year and species. A total of twenty-nine sites were recorded in M.jurtina and twenty in P.tithonus. Results Their distribution is shown in Fig. 1. The nature of the individual data sets, particularly the The overall means for each species and site are variability between sites in the frequency of given in Table 1. The mean date of adult flight recording visits, means that some caution must period is nearly 10 days earlier in M.jurtina (27 be exercised when comparing estimates of tim- July) but the SD about this date is 50% greater.
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