THE RED ADMIRAL, ILLUSTRATIVE NATURAL HISTORY OF

[Tyrant]

Why do you make such an imposition out of yourself?

 

[Tyrant]

Do you think it's self-aware?

 

[Gagh]

 

[Grammour Boy]

^^ Off-topic. Gagh, Need I remind you that you're in the no-spam zone? Why do you insist on putting on the stupid act? You know full well that spam is prohibited in Armchair Philosophers.

 

[Tyrant]

Why are you spamming a no-spam zone with an off-topic question, Grammour Boy?

 

[Grammour Boy]

Why are you spamming a no-spam zone with an off-topic question, Grammour Boy?

Gagh's poster of the BBW party has nothing to do with the Red Admiral.

(Admins: Can you please clean up this thread and split off the off-topic spam whose sole purpose is to disrupt? Thank you in advance.)

 

[Tyrant]

Why are you spamming a no-spam zone with an off-topic question, Grammour Boy?

 

[Grammour Boy]

The effects of methodological limitations in the study of butterfly behavior and demography: a daily study of Vanessa atalanta (Lepidoptera:
Nymphalidae) for 22 years

Henry F. Swanson and Julián Monge-Nájera
Biologia, Universidad de Costa Rica, San José, Costa Rica
Revista de Biología Tropical, 48:605-614, 2000.​

Abstract

Normally, butterfly behavior and population size are studied intensively for brief periods or occasionally for long periods, not in detail for long periods, producing an incomplete view in both cases. How time limitation affects studies has been unknown for a long time. This paper analyses this problem based on an intensive long term study of Vanessa atalanta (L.) that covered nearly 8000 days, most of them consecutive, for 22 years (April 15, 1977-April 14, 1999), in a subtropical habitat near Orlando, Florida. There is no evidence that ethological studies are affected by their normally brief duration (one year or less), but the analysis of yearly values hid the associations of number of individuals and arrival time with climate. In small areas, isolated population counts lasting less than two weeks are not reliable, according to this study. We found no difference in number of visitors for El Niño years. The daily number of visitors was inversely correlated with temperature and precipitation, but arrival time of the first visitor was positively correlated with both. The number of visitors reaches a peak near the end of Winter. The activity period span is greater than in more seasonal climates. Individuals were active even at 10º C and with 9 m/s winds. Individuals with fresh wing condition were most common from January to June. There were 82 atypical cases of individuals arriving before 12:00 hr. Aerial interactions were seen whenever there was more than one individual in the site (i.e. 41 % of days, N=7634 total days). Only once in these 22 years was predation seen.

For economic and logistic reasons, butterfly biology normally is studied by observing a population intensively for short periods (which is satisfactory to solve ethological problems, Monge-Nájera et al. 1998 ), or through isolated observations over a long time (e.g. Stekolnikov's 1992 study that spanned more than half a century). The effort closest to a long term intensive study may be the British monitoring system (Pollard et al. 1997), but it also has important methodological limitations (Nielsen and Monge-Nájera 1991). In general, we ignore how the lack of long term intensive studies effects our view of butterfly ecology.

The current view of butterfly population ecology is summarized in the following paragraphs.

Density dependent factors (e.g. hostplants): The importance of specific hostplant species, individuals and even plant parts for population size, distribution and genetic variability is controversial (Thompson 1988, Daily et al. 1991 Goulson et al. 1997) but food abundance generally is associated with population increases (e.g. Wickman et al. 1990)

Density independent factors: In tropical areas of varying altitude, warmer and moister periods are associated with butterfly population increases, while in temperate areas, warm weather normally produces premature emergence, additional generations, increased breeding success and larger adult populations (Pollard et al. 1997, Wickman et al. 1990, Steiner 1991).

How butterflies reduce the effect of ecological factors: Butterflies can control density dependent and density independent factors in two ways: (1) locally by behavioral and physiological mechanisms, and (2) ex situ by migration. Behavioral mechanisms include activity pattern coordination with the hostplant's reactions to climate, while physiological adaptations even involve larval monitoring of daylength to control development (Young 1983, Kunte 1997, Nylin 1997).

Migration: Butterfly travelling varies from small changes in microhabitat distribution to long range migration. Migration seems to be genetically determined and relates with lifespan (Baker 1984, Ehrlich 1984). While solar radiation, temperature and wind may (Calvert et al. 1992, Monge-Nájera et al. 1998 ) or may not (Frey et al. 1992) help predict the microhabitats where populations concentrate. It seems clear that topography and the microclimate pattern that topography creates affect both butterfly population size and biodiversity (Gutiérrez 1997).

Mid-range movements: Mid-range movements are associated with isolated habitat patches that predispose a species to the shifting mosaic metapopulation model (Harrison et al. 1998 ), with relatively frequent local extinctions and re-establishments. Stepping-stone gene flow is important at least in some metapopulations (Neve et al. 1997, Peterson 1997 Singer and Thomas 1997).

Long-range migrations: Long-range migrations are known in tropical and temperate butterflies, among them the Red Admiral, Vanessa atalanta L., a species that appears to fit the requirements for survival in urban areas (Swanson 1979, Kitahara and Fujii 1997, Srygley et al. 1997, Thomas 1984). It is a widespread nymphalid found in the American continent from central Canada to Guatemala, often in clearings (Bitzer and Shaw 1979, Tuberville et al. 1996).

This paper reports on an intensive (daily) long term study (22 years) of a V. atalanta population in a subtropical habitat with emphasis on the association of number of individuals and arrival time with climate.


Materials and Methods

Individual butterfly visitation records (and a qualitative microclimatic classification) were made by the senior author in an urban clearing devoid of hostplants in Winter Park, near Orlando, Florida, USA (28° 34'57'' N, 81° 20'04'' W) for 22 years (April 15, 1977-April 14, 1999: 8035 days). Details of the site and the unusual conditions that allowed daily observations for such a long period appear in Swanson (1998 ).

Climatic data are from monthly means provided by government databases available in the Internet (http://water.dnr.state.sc.us/climate/). Besides the inferential statistics presented here, graphs of daily changes in number of visiting butterflies and their arrival time (available in the on-line edition of this journal) were analyzed visually for non-linear associations (Kozlov et al. 1997) but none were found. Non-parametric statistics were used to avoid potential conflict with the less realistic requirements of parametric tests. "Visitation" and "number of daily visitors" are defined as total number of occupants and intruders arriving each day.


Results

Natural history: Individuals were active even at 10° C and with 9 m/s winds. Individuals with wings in fresh condition were most common in the first half of year (January-June) when they represented roughly three quarters of the population. Smaller individuals (general impression: no measurements were taken) were seen from late December to late March. There were 82 atypical cases of individuals arriving before 1200 hr (not included in the graphs). Aerial interactions were seen whenever there was more than one individual in the site (i.e. 41 % of days, N=7634 total days, Fig. 1).

A total of 4 794 visits were recorded, with an overall attendance rate of 59.5 %. During the 2 hr afternoon visit period there were frequently 1-3 individuals in the site, two times there were eight and once (March 15, 1982) there were ten. Only once in these 22 years was predation seen: an individual that flew with difficulty was captured by a bird (Cardinalis cardinalis). Some specimens had missing legs or lacked parts of wings, antennae or proboscis.

Visitation pattern: There are no hostplants on the site. The population is composed of visitors that perch and participate in aerial interactions sensu Bitzer and Shaw (1979). The number of visitors begins to increase in autumn, reaches a peak near the end of winter and decreases almost constantly during the spring and summer (Fig. 2a, 2b). Generally, there are two visitation patterns. Pattern I (1977 to 1985): a marked curve with zero population around October, e.g. Fig. 2a). Pattern II (1986-1996):a flatter curve with 1-5 months without visitors, e.g. Fig. 2b. Arrival time of first individual to enter the area had the same pattern during the study period, with a soft increase from January through June and a soft decline until December (Julian dates, Figs. 2b and 2 d).

The mean daily number of visitors a particular month was inversely correlated with mean monthly temperature (Spearman Rank Correlation, SRC: r =-0.43, p< 0.01, n=261 months; mean monthly values for number of visitors and climatic factors were used in this and following tests) and precipitation (SRC: r = -0.34, p< 0.01, N=261) (Fig. 3A,C). There were occasional absences of 1-14 days that represented gaps in the general trend mostly during stormy weather (Fig. 2 A-D, a few represent the rare instances in which observer was absent). Arrival time of the first visitor was positively correlated with mean monthly temperature (SRC: r = 0.84, p< 0.01, N=238 ) and with mean monthly precipitation (SRC: r = 0.49, p< 0.01, N=231) (Fig. 3B,D). Total daily number of individuals arriving at the site was associated with a subjective classification of climate (Kruskal-Wallis ANOVA, p< 0.0001, N=5658 days; mean ranks: 4229 windy, 3152 good climate, 2614 cold, 743 overcast or rainy) but not with arrival time (Kruskal-Wallis ANOVA 1.7, p=0.63, N=4 681 days).

Yearly mean values were not correlated (SRC values: visitors versus temperature -0.17, visitors versus precipitation -0.40; arrival time versus temperature 0.17, arrival time versus precipitation 0.02; in all cases N=22 years and p> 0.05). (Fig. 4)

There was no significant difference in number of visitors between El Niño (mean 1.13 visitors/day, standard deviation SD 0.21, range 0.90-1.50, N=11, years 1977, 1978, 1982, 1983, 1986, 1987 and 1991-1995) and non-El Niño years (mean 1.25, SD 0.28, range 0.80-1.70 visitors/day).

Discussion

The migratory routes of V. atalanta in North America are poorly known, but a greater number of studies in Europe have shown that the migratory population follows temperature layers (Benvenuti et al. 1996), is subjected to significant wind drift and travels with other species (Hansen 1997); has time-compensated sun orientation, migrates thousands of kilometers, feeds in winter and sometimes reproduces while traveling northward in the warmer part of the year, around June (Baker 1972, 1984, Stefanescu et al. 1996, Hansen 1997, Palmqvist 1998 ). It is now clear that migrations are bidirectional (Benvenuti et al. 1996) but how the travel information is passed among butterfly generations in general is not known (Pasteur 1984) and future studies of V. atalanta should stress this aspect.

In this study, seasonal changes in the number of visitors, with fewer visitors in the moister and warmer months and predominance of fresh individuals from January to June suggest that V. atalanta overwinters in Florida and migrates to northerly regions when high summer temperatures affect Florida (R. Bitzer 2000, pers. comm.). A similar migration from Africa to Europe is known (Larsen 1993). This would fit with the smaller individuals seen from late December to late March: fresher individuals may emerge in Florida as the offspring of the larger butterflies first seen in late October - early November. These larger individuals could have migrated from more northerly regions earlier in the fall, according to the parallel migration pattern of V. atalanta in Europe described by Larsen (1993) and Benvenuti et al. (1996) (R. Bitzer 2000, pers. comm.). For V. atalanta temperature is a more important limiting factor than host plant availability (Bryant-Simon et al. 1997, Maier and Shreeve 1995).

The activity period span in Florida is greater than in the more seasonal climate of Iowa, where V. atalanta is active from April to October (Bitzer and Shaw 1979, 1995). The greater number of visitors on windy days suggests that the site was selected because it was surrounded by vegetation and buildings (see Swanson 1998 ).

The El Niño Migration Model predicts that overpopulation increases butterfly migration to outbreak level during El Niño years (Larsen 1984, Myres 1985). We found no difference in number of visitors for El Niño years, apparently because V. atalanta is a regular rather than an outbreak migrator.

The limited temperature range that characterizes sites where V. atalanta occurs (Bryant-Simon et al. 1997, Maier and Shreeve 1995) and butterflies' inability to fly under heavy rain can explain why individuals arrive at their perching spots later in the hot, rainy months. V. atalanta males establish courtship territories 2-4 hours before sunset (Wiltshire 1997, Bitzer and Shaw 1979, 1995) and use neural motion detectors adapted to great speed (O'Carroll et al. 1996) in aerial interactions (Bitzer and Shaw 1979). Territories have distinct characteristics (sunlit clearings and ridge tops) and are used by different individuals over the years (Bitzer and Shaw 1979, Wiltshire 1997). Despite its methodological difference with previous studies, this study produced the same conclusions about territorial behavior, and also agrees with the ecological model of perching behavior which proposes that perching is associated with relatively high temperature, lekking and long life (Wickman 1992a,b, van Dyck et al. 1997a,b, van Dyck and Matthysen 1998. No data are available regarding the wing color and body proportion elements of the model).

If our results can be generalized (and that will be unknown until more long term intensive studies become available) behavioral studies of butterflies do not suffer from the general impossibility of daily observation for long periods, but at least for small areas, occasional population counts covering less than two weeks are unreliable: even total absence of individuals for such periods can simply represent gaps in the general trend. Furthermore, the analysis of yearly values hid the associations of number of individuals and arrival time with climate that were found when a smaller scale was used. Long term studies based on yearly samplings should be re-evaluated for this aspect.


Acknowledgments

We thank Shirley Gregory and Pat Miller for assistance and support, María Isabel González for the statistical analysis, Marisol Rodríguez Pacheco and Ana Araya Anchetta for data digitalization and Royce Bitzer, Department of Entomology, Iowa State University, William Ramírez, Museo de Insectos, Unversidad de Costa Rica, Per-Olof Wickman, Dpt. Mathematics and Science, Stockholm Institute of Education and one annonymous reviewer, for suggestions that greatly improved the manuscript.


References

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Wickman, P.O. 1992b. The influence of temperature on the territorial and mate locating behaviour of the small heath butterfly, Coenonympha pamphilus (l.) (Lepidoptera: Saturidae). Behav. Ecol. Sociobiol. 16: 233-238.

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[Tyrant]

Oh.

 

[Grammour Boy]

 

[bad dog]

This will get rid of the bugs.

 

[Grammour Boy]

Increased abundance of the red admiral butterfly Vanessa atalanta in Britain: the roles of immigration, overwintering and breeding within the country

Pollard & Greatorex-Davies
Biological Records Centre, Institute of Terrestrial Ecology, Monks Wood, Cambridgeshire, U.K. ​

Abstract
The migratory butterfly Vanessa atalanta increased in abundance at monitored sites in Britain from 1976 to 1996. Three possible causes of the increase are improved winter survival within Britain, greater breeding success within Britain, and increased immigration.

Trends during most of the season were similar to those of immigrant or overwintered individuals in spring; thus the evidence does not support greater breeding success in Britain. As abundance in spring was not correlated with abundance in the previous autumn, when trend was taken into account, it seemed unlikely that overwintering in Britain was important. Thus the increase in abundance was probably due to increased immigration. Incidental to the main study, the mean index per site per year was closely correlated with the collated index, the usual measure of annual fluctuations. This agreement suggests that the mean index may be a useful check for trends in monitoring data for other wide-ranging organisms.

Introduction
The Butterfly Monitoring Scheme (BMS) ( Pollard & Yates 1993) has monitored the abundance of butterflies at sites in Britain since 1976. During this period, although butterflies that were already rare have tended to decline further, several species which were already common and widespread within their ranges have increased in abundance ( Pollard et al. 1995) and expanded their ranges ( Pollard & Eversham 1995). Included amongst the species which have increased markedly in abundance at monitored sites is the migratory butterfly Vanessa atalanta (L.), the red admiral.

The main larval food-plant of V. atalanta is nettle Urtica dioica L. Two nettle-feeding butterflies resident in Britain, Polygonia c-album (L.) and Inachis io (L.), have also increased in abundance from 1976 to 1996 and, particularly in the case of the former, have expanded in range ( Pollard & Yates 1993). It is therefore of interest to see whether the increase in abundance of V. atalanta was associated with increased breeding success in Britain. If this was the case, it might indicate, amongst other possibilities, that a change in the abundance or quality of the food-plant in Britain is implicated in the changes.

Vanessa atalanta migrates to Britain each spring from further south in Europe, although the areas from which it originates are not known in any detail. Adults occasionally survive the winter in Britain ( Frohawk 1934) and there are many recent records of adults in January and February (e.g. Allen 1996; Gann 1996; Gardiner 1996); occasionally larvae may also overwinter successfully ( Tucker 1996). Nevertheless, there is little doubt that most spring adults have immigrated rather than hibernated here ( Thomas 1986; Emmet & Heath 1989). Elsewhere in Europe, it is a common resident as far north as central Germany and migrates much farther north, occasionally even close to the Arctic Circle ( Emmet & Heath 1989).

The eggs are laid on nettles Urticadioica and occasionally on other related plants. Larvae are rarely found before June and it can be assumed that virtually all adults recorded in April, May and June are immigrants. Further immigration may continue into the summer ( Emmet & Heath 1989), but there are probably two generations of adults bred in Britain ( Heath et al. 1984). Adults are seen throughout the summer until September or October, when there is some evidence of a return southwards ( Williams 1958 ). Occasional movements and congregations have been recorded along the south coast in autumn ( Emmet & Heath 1989). As recording in the BMS does not extend into October, some data for V. atalanta are lost.

In this paper we

1 present the evidence for an increase in abundance of V. atalanta during the period 1976–96;

2 examine the seasonal pattern of changes in abundance to determine whether the increase is likely to be due to increased immigration, better overwinter survival or improved breeding success within Britain.

To address the second aim, we examined trends in the early part of the recording season, when it can be assumed that the butterflies recorded were immigrants (or had overwintered) and compared them with trends later in the season, when home bred butterflies are likely to make up a substantial part of the total. To test for an effect of overwintering, we looked for a relationship between early season counts and those late in the previous year. If winter survival made a substantial contribution to the number of spring butterflies, some degree of correlation between spring and previous autumn abundance would be expected.

Records of immigrant Lepidoptera have been collected and summarized for many years in several European countries, e.g. in Britain by R.F. Bretherton and J.M. Chalmers-Hunt in the Entomologist's Record and Journal of Variation. Such schemes provide invaluable data on the species present and on patterns of migration. However, they do not permit analysis of trends in abundance because recorder effort is difficult to assess, varies from year to year, and has tended to increase over time. The systematic monitoring described here has made it possible, not only to demonstrate trends, but also to distinguish between broad factors which may be responsible for the trends. However, the ultimate causes remain unknown.

Methods
Butterfly Monitoring Scheme (BMS)

The methods used in the BMS have been described by Pollard & Yates (1993) and will be summarized only briefly here. At each site a fixed route is walked in each of 26 recording weeks from the beginning of April until the end of September. All butterflies seen within prescribed limits are recorded. Counts are made only when weather conditions meet specific criteria and are standardized according to instructions provided. The weekly counts are used to calculate an annual index of abundance; this index is the sum of the mean weekly counts, with estimates for missing weeks provided that only a few weeks are missed. These site indexes have been shown to be robust measures of changes in abundance for several species (studies reviewed by Pollard & Yates 1993) and it is reasonable to assume that this is also true of V. atalanta.

The site indexes are collated (using the method of Moss & Pollard 1993) to produce "all-sites" indexes, which can be used to show broad fluctuations and trends. The BMS sites are predominantly nature reserves or other protected areas. Thus, for many sedentary species the all-sites indexes cannot be assumed to be typical of the wider countryside. However, for wide-ranging species that breed predominantly in the farmed countryside, including V. atalanta, it is probable that national trends are closely reflected at BMS sites.

Analysis of trends

The method of estimating trends was standard regression, in which the y-values are assumed to be independent of each other. In time series data of the type examined here, data points may not be independent ( Diggle 1990); i.e. abundance in a given year may depend to some extent on abundance in the previous year. In the case of a migrant such as V. atalanta, which is not considered to form permanent populations in Britain, such 'autocorrelation' is unlikely to be a serious problem; nevertheless some checks are included in the study.

Data were log10 transformed prior to regression. Zero values were omitted in preference to the use of a log(n+ 1) transformation with a consequent bias in trend estimates.

Trends in seasonal data

The collated index, used to show overall trends in abundance, could not easily be extended for use with weekly counts that were typically small with many zeros. An alternative measure of annual changes is the use of mean index per site per year. This measure takes no account of the fact that the composition of sites changes to some extent from year to year, so the mean index might be expected to be influenced by the presence or absence, in particular years, of data from sites with typically very high (or very low) counts.

A comparison of mean index per site with the collated index for V.atalanta showed a remarkable agreement ( Fig. 1, r = 0.994). This almost perfect correlation was probably obtained because V. atalanta does not occur in local populations, with the potential for high local abundance, but at low density over the whole country. The other common migrant butterfly Cynthia cardui showed a similar close agreement between the two measures (r = 0.995).

Given the close similarity of the mean annual index and the collated index, it can be assumed that the mean weekly counts per site recorded in a given year, can be used to compare seasonal trends and fluctuations in abundance over the recording period.

To estimate seasonal trends, the data were summed over two early periods (weeks 1–7, 1 April to 19 May; weeks 8–14; 20 May to 7 July), chosen to eliminate zeros. Butterflies recorded in the first period can be regarded as immigrants (although the possibility of overwintering is also considered) and those in the second period mostly, and possibly also entirely, immigrants. Data for the weeks 15–26 contained no zeros and so were treated individually.

To test for the potential importance of overwintering, regressions were conducted using the early spring counts (weeks 1–7) as the dependent variable, with trend and mean weekly counts for the last few weeks of the previous year as independent variables.

Results

Table1:

Fig. 1

Fig. 2

Fig. 3

Fig. 4

Overall trends in abundance

Data were available from 137 sites with indexes for V. atalanta for 1 or more years. Considering the 79 sites with indexes (> 0) for 8 or more years, there were 23 significant increases (P < 0.05) and one significant decline. The increases were concentrated in southern Britain, reflecting the distribution of sites in the BMS ref ( Fig. 2). The collated index increased significantly [b = 0.036 ± 0.010 (SE), P = 0.002]. When the previous year's index was included with the current year's index in the regression as a second independent variable, the trend was larger (b = 0.046 ± 0.012, P = 0.001), suggesting that autocorrelation did not give a misleadingly high significance for trend in this analysis. The direct measure of autocorrelation, the first serial correlation after detrending the collated index, was r1 = 0.40 (P = 0.089, not significant).

Seasonal pattern of trends

The early weeks of the season had low counts ( Fig. 3), with many zero values (58% zeros in weeks 1–7). For this reason, the significance of trends in individual weeks was first estimated using the Spearman rank correlation between weekly index and year. There were significant increases (P < 0.05) in 17 of the 26 weeks; weeks 1–4, 6, 9, and 14–24.

When the indexes for the early weeks were summed, there were significant increases shown by linear regression in both of the early periods and in all but the last two individual weeks ( Table 1). The largest trend was in the earliest period (weeks 1–7), although this was not significantly larger than trends in most other weeks. The most striking feature was the similarity of trends from the start of the recording season in April until mid-September.

When trend was taken into account in multiple regression, there were significant associations between weekly abundance and abundance in weeks 1–7 for 4 weeks (including the grouped weeks 8–14) and also between the annual collated index and abundance in weeks 1–7 ( Table 1). Counts in weeks 1–7, contained only 1.4% of the total sums of mean weekly counts and so made a negligible numerical contribution to the collated index. Thus, in addition to having similar trends to that shown by the early immigrants, the later season counts tended to have similar annual variability to that of the early immigrants (Fig 4).

Testing for an overwinter effect

There was no association between spring abundance (weeks 1–7) and abundance in the previous autumn (weeks 20–26), when trend was taken into account. The partial regression coefficient for log10 weeks 20–26 was 0.196 ± 0.156 (P = 0.224).

Discussion

The overall increase in abundance of V. atalanta over the period 1976–96, as shown by the collated indexes of the BMS, closely reflects an increase in abundance in the first 7 weeks of the season (1 April to 19 May). Indeed, over much of the summer trends closely resemble those of the first few weeks, only departing from this agreement during September. The data for the first 7 weeks comprise less than 2% of the whole.

This strong association between abundance in the current year and abundance in early spring contrasts with, at most, a very weak association between spring abundance and abundance in the previous autumn. The evidence therefore indicates that increased immigration is the major factor implicated in the increased abundance of V. atalanta in Britain, with breeding success and overwintering in Britain of less importance.

Thus there was no support for the view that the increase in abundance of V. atalanta was associated with those of Polygonia c-album and Inachis io in Britain and that their common food plant Urtica dioica was involved. The results suggest a more general increase in abundance of V. atalanta farther south in Europe, but we have no information on the causes of such a trend.

The eventual break-down, which occurs in September, in the similarity of trends with those in the spring, may be associated with a return southwards. Alternatively, it may simply reflect variation in breeding success and increasing separation (in time) from the immigrant individuals.

An unexpected finding of this study, unrelated to the initial aims, was the remarkable agreement between the collated index, based on ratios of change using only sites recorded in each pair of years, and the mean index per site. Using the ratio method, collated indexes are subject to the possibility that an error (or errors) introduced early in a series continues to influence all subsequent values; such effects may result in apparent, but spurious, trends. This result suggests that for wide-ranging species of butterflies and other organisms, the mean index may provide a useful check for the presence of problems of this type.

Acknowledgements

The BMS is supported by Joint Nature Conservation Committee and the Institute of Terrestrial Ecology and we are grateful to both of these organizations. We also thank P. Rothery for comments on an earlier draft. Finally, we are grateful for the enthusiasm and commitment of the many recorders, without whom synoptic studies of this type would be impossible.

References:
1 Allen, A.A . (1996). Red admiral butterfly Vanessa atalanta (L.) (Lepidoptera: Nymphalidae) in mid February. Entomologist's Rec. J. Var., 108, 207.

2 Diggle, P.J . (1990) . Time Series, A Biostatistical Introduction. Oxford: Clarendon Press.

3 Emmet, A.M & Heath, J . (1989) . The Moths and Butterflies of Great Britain and Ireland, Vol. 7, The Butterflies. Colchester: Harley Books.

4 Frohawk, F.W . (1934) . The Complete Book of British Butterflies. London: Ward Lock.

5 Gann, P . (1996). Red admiral butterfly Vanessa atalanta (L.) (Lepidoptera: Nymphalidae) in January. Entomologist's Rec. J. Var., 108, 195.

6 Gardiner, B.O.C . (1996). A late and an early red admiral butterfly Vanessa atalanta (L.) (Lepidoptera: Nymphalidae). Entomologist's Rec. J. Var., 108, 207.

7 Heath, J , Pollard, E , Thomas, J.A . (1984) . Atlas of Butterflies in Britain and Ireland. Harmondsworth: Viking.

8 Moss, D & Pollard, E . (1993). Calculation of collated indices of abundance of butterflies based on monitored sites. Ecol. Entomol., 18, 77 83.

9 Pollard, E & Eversham, B.C . (1995) . Butterfly monitoring 2 – interpreting the changes. In Ecology and Conservation of Butterflies, ed. A.S. Pullin, pp. 23–36. London: Chapman and Hall.

10 Pollard, E , Moss, D , Yates, T.J . (1995). Population trends of common British butterflies at monitored sites. J. Appl. Ecol., 32, 9 16.

11 Pollard, E & Yates, T.J . (1993) . Monitoring Butterflies for Ecology and Conservation. London: Chapman and Hall.

12 Thomas, J.A . (1986) . RSNC Guide to the Butterflies of the British Isles. London: Newnes.

13 Tucker, M . (1996). The red admiral–overwintering in the UK in its early stages. Butterfly Conservation News , no. 62, 21 22.

14 Williams, C.B . (1958 ) . Insect Migration. London: Collins.

 

[Grammour Boy]

This is a very interesting article on why butterflies tend aggregate on hilltops. [Source: American Midland Naturalist, 120(2):441-443, 1988]

What Constitutes a Hill to a Hill topping Butterfly?
John F. Baughman; Dennis D. Murphy
Department of Biological Sciences, Stanford University, Stanford, California​

Abstract.—Butterflies observed in a nearly flat field tend to aggregate on the highest portion of the habitat, even though the elevational difference between it and other areas is very slight. This suggests that hill topping behavior in insects may be triggered by seemingly insignificant topographic relief.

Introduction​

Butterflies and other insects frequently aggregate on hilltops or in association with other distinct topographic features such as roads, gullies, rock outcrops or prominent vegetation (e.g., Shields, 1967; Bitzer and Shaw, 1979; Ehrlich and Wheye, 1986; Scott, 1986; Alcock, 1987). These aggregations are thought to facilitate mate location (Shields, 1967; Scott, 1974; Alcock, 1987), although this has not been demonstrated conclusively.

The topographic features on which aggregation behavior is most often noticeable are rather promissmanners is a goddess!nounced; for example, a prominent ridgetop in the Sonoran Desert (Alcock, 1987), a mountainside road in the Rocky Mountains (Ehrlich and Wheye, 1986) or an unscalable cliff in the Colombian rainforest (Shapiro, 1979). To test whether more subtle topographic features can play a role in determining the spatial distribution of insects, we examined the distribution of butterflies in a nearly flat habitat. Indeed, we wanted to know how big must a hill be to be recognized as a hill by a hilltopping butterfly.

Methods​

The population of Euphydryas editha Boisduval (Nymphalidae: Nymphalinae) in Area C of Stanford University's Jasper Ridge Biological Preserve occupies a grassland of remarkable homogeneity. The nearly 10-ha habitat is essentially flat, with a very small rise in elevation near the center (Baughman el al., 1988a); the vertical difference between the lowest point and the highest point, 300 m apart, is just 15 m.

In 1985 a mark-recapture program was carried out in Area C in which 63 different adult males and 64 different adult females were handled. The area was divided into 33 0.25-ha quadrants and the position of each individual within the habitat was recorded on capture and subsequent recaptures. Butterflies were handled on 14 separate days, approximately every other day, during the month-long flight season. On each sampling day, all of the butterflies observed in Area C were collected. Butterflies were marked following the procedure of Ehrlich and Davidson (1961).

Results and Discussion​

Of the 157 male "handles" (original captures plus recaptures of the same individuals), 84 (53.5%) were in the four quadrants comprising the "hill." Similarly, 46.7% (57/122) of female handles occurred in the same area. Thus, about half of all adult handles were situated in only 12.1% of the available habitat. Furthermore, no adults were ever observed in 11 of the 33 quadrants, fully one third of the habitat area. The quadrants where no adults were observed tended to be at the margins of the habitat, but were distributed throughout Area C.

Despite the small size of the hill, its elevation appears to be the only environmental factor that can explain the clumped distribution of butterflies observed in Area C during 1985. Both larval host plants and adult nectar sources of Euphydryas editha are ubiquitous in Area C (Singer, 1972; Ehrlich et al., 1975; Murphy, 1984). This widespread availability of resources suggests that they do not constrain adult distribution. Moreover, until recently, the population of Euphydryas editha in Area C has been comparatively large, often consisting of over 1000 adults, and as many as 4000 in some years (Ehrlich etal,, 1975; Murphy eta/., 1986). When the population was large in size, adults were found throughout area C. Over the last half decade, however, when several consecutive years of poor weather, particularly drought, reduced the population to the low levels observed in 1985, the effect of the "hill" became apparent. This indicates that while all of Area C may he suitable habitat, the hill is, for some reason, more attractive to adult butterflies, at least in years of low population size.

Area C is small enough that Euphydryas editha adults can and do freely fly from one end of the habitat to the other. Nonetheless, Singer (1972 and pers. comm.) has suggested that adult distributions may be strongly influenced by the distribution of adult eclosion sites. It is conceivable that more adults were found on the hilltop than elsewhere because more matured there. In the neighboring population of E. editha in Jasper Ridge Area H, however, the distributions of newly eclosed and older adults were found to be significantly different (Baughman et at., 1988a). In at least one population, therefore, adult distributions appear not to be determined by the location of pupation sites.

It may be that the hill itself is not what attracts butterflies, but rather, the hilltop may have a slightly warmer orientation, and hence be more attractive to thermoregulating butterflies than other portions of the habitat. Such an explanation seems unlikely, however, for two reasons. First, the hilltop itself is essentially flat, so that any differences in temperature between it and other flat parts of the habitat, where adults are comparatively scarce, would be slight. Second, ovipositing adult females would be expected to spend most of their time, particularly late in the flight season, in the coolest parts of the habitat where larval host plants remain green and edible the longest (Singer, 1972; Weiss et at., 1987, 1988 ). The coolest part of the Area C habitat is that with a 11° NW-facing exposure (comprising 3% of the overall habitat) connecting the hilltop to the flat area below. Sixteen handles of males (10.1%) and 16 of females (13.1%) occurred on that slope. Thus, while that slope appears to be more attractive than much of the overall habitat, it also appears to be somewhat less attractive than the hilltop itself.

Reasons why butterflies aggregate on hilltops have been considered in detail elsewhere (Shields, 1967; Ehrlich and Wheye, 1986; Alcock, 1987; Baughman et at., 1988b). Our observations, however, indicate that seemingly quite minor topographic relief, on the order of a few meters over many hectares, may play a significant role in determining the distribution of butterflies within a habitat. This has two important implications. First, behavior observed on a hilltop may not be a specific adaptation to that hilltop. Rather, it merely may be a particularly noticeable expression of an otherwise commonly, but more subtly exhibited, behavioral tendency. Second, studies which attempt to explain the distrimissmanners is a goddess!bution of adult butterflies in relation to "nontopographic" habitat features, such as the distribution of oviposition, host plants, nectar sources, or other adult butterflies, but which ignore even seemingly minor topographic relief, in fact, may be incomplete.

Literature Cited​

Alcock, J. 1987. Leks and hilltopping in insects. /. Nat. Hist., 21:319-328.
Baughman, J. F., D. D. Murphy and P. R. Ehrlich. 1988a. Emergence patterns in male checkerspot butterflies: testing theory in the field. Theor. Pup. Biol., 33:102-113.
----------,----------AND----------. 1988b. Population structure in a hilltopping butterfly. Oecologia, 75:593-600.
BrrzER, R. J. and K. C. Shaw. 1979. Territorial behavior of the Red Admiral Vanessa atalanta (Lepidoptera: Nymphalidae). /. Res. Lepid., 18:36-49.
Ehrlich, P. R. and S. E. Davidson. 1961. Techniques for capture-recapture studies of Lepidoptera populations. Lepid. Soc, 14:227-229.
---------and D. Wheye. 1986. "Non-adaptive" hilltopping in male checkerspot butterflies (Euphymissmanners is a goddess!dryas editha). Am. Nat., 127:477-483.
----------, R. R. White, M. C. Singer, S. W. McKechnie and L. E. Gilbert. 1975. Checkerspot butterflies: a historical perspective. Science, 118:221-228.
Murphy, D. D. 1984. Butterflies and their nectar plants: the role of the checkerspot butterfly Euphydryas editha as a pollen vector. Oikos, 43:113-117.
----------, M. S. Menninger, P. R. Ehrlich and B. A. Wilcox. 1986. Local population dynamics of adult butterflies and the conservation status of two closely related species. Biol. Conserv., 37:201-223.
Scott, J. A. 1974. Mate locating behavior of butterflies. Am. Midi. Nat., 91:103-117.
----------. 1986. The butterflies of North America. Stanford University Press, Stanford, Calif p. 538.
Shapiro, A. M. 1979. Notes on the behavior and ecology of Reliquia santamarta, an alpine butterfly (Lepidoptera: Pieridae) from the Sierra Nevada de Santa Marta, Colombia, with comparisons to nearctic alpine Pierini. Stud. Nevtrop. Fauna, 14:161-170.
SHIELDS, O. 1967. Hilltopping. / Res. Lepid., 6:69-178.
Singer, M. C. 1972. Complex components of habitat suitability within a butterfly colony. Science, 176:75-77.
Weiss, S. B., D. D. Murphy and R. R. White. 1988. Sun, slope, and butterflies: topographic determinants of habitat quality for Euphydryas edilha bayensis. Ecology, 69:1486-1496.
---------, R. R. White, D. D. Murphy and P. R. Ehrlich. 1987. Growth and dispersal of larvae of the checkerspot butterfly, Euphydryas edilha, Oikos, 50:161-166.

 

[bad dog]

We all need one.

 

[Grammour Boy]

"Michael Majerus's fascination for "bugs," as he calls all insects, was ignited at the tender age of 4. His mother can recall the exact moment: It was late summer, and he caught his first butterfly — a red admiral resting on a white chrysanthemum — with his bare hands. From that point on, he was hooked"

In Defense of Darwin and A Former Icon of Evolution
(Science, 304:1894-5, 2004)​

Michael Majerus's fascination for "bugs," as he calls all insects, was ignited at the tender age of 4. His mother can recall the exact moment: It was late summer, and he caught his first butterfly — a red admiral resting on a white chrysanthemum — with his bare hands. From that point on, he was hooked. At night when other children were in bed, the young Majerus roamed the English countryside tending his moth traps. Now 50 and in his 25th year of teaching evolutionary genetics at the University of Cambridge, Majerus still runs his moth traps most nights.

Majerus's research has focused on sexual selection, sex-ratio manipulation, and the evolution of melanism (the darkening of body color) in various moths, butterflies, and ladybirds. But over the past few years, half of his working life has been occupied by one controversial species, the peppered moth, Biston betularia, and an infamous study that's been attacked by both evolutionary biologists and anti-evolutionists.

Through his research, Majerus found himself embroiled in the scientific debate over the evolutionary forces behind melanism in the peppered moth. Experiments by British lepidopterist Bernard Kettlewell in the 1950s claimed to show that bird predation, coupled with pollution, was responsible for a color shift in the moth population. But problems with Kettlewell's methodology led some scientists to doubt his conclusions. Majerus was not the first to point out the flaws, but by doing so, he inadvertently set off a wave of anti-evolutionist attacks. While acknowledging that Kettlewell made mistakes, Majerus believes Kettlewell was right in his conclusions and has taken it upon himself to prove it.

As Majerus shows off some of the roughly 100,000 peppered moth pupae he'll rear for his latest experiment, it's clear that he's prepared to go to great lengths to make his case. Once the moths begin to emerge in May, Majerus begins a daily grind. He releases them at dusk and gets up at dawn to observe their fate: counting how many are plucked from their resting places by birds, and how many survive to see another night. He will continue this routine into August, as he has done for the past three summers. All he needs, he reckons, is another 2 years' worth of data — a total of some 4000 moth observations — to settle the controversy over whether bird predation is the major selective force in favoring one color form of the peppered moth over another.

Small and unobtrusive, the peppered moth doesn't look like the star in an evolutionary drama. But the rise and fall of the almost-black melanic form (carbonaria) in tandem with changing pollution levels has become the most famous example of evolution in action. Through his pioneering experiments, Kettlewell claimed to have demonstrated that melanic peppered moths were more common in industrialized areas because they escaped the attention of predatory birds when resting against soot-blackened, lichen-free bark. Because more of the darker ones survived to produce the next generation, he argued, entire populations grew darker.

But doubts emerged over Kettlewell's methodology in recent decades as researchers failed to replicate some of his results. His predation experiments were chiefly criticized for their artificiality: He placed the moths on exposed parts of trees in broad daylight, when they don't normally fly, rather than allowing them to settle naturally; he released them in large numbers, thereby inflating moth densities and possibly creating a magnet for predatory birds; and he used a mixture of lab-reared and wild-caught moths without checking to see whether they behaved the same way. Majerus summarized these criticisms in a book on the evolution of melanism in 1998 and stated that the simplified textbook story of the peppered moth was inaccurate, while asserting that Kettlewell's conclusions were qualitatively sound. Majerus had no idea at the time what a furor his book would cause.

Jerry Coyne — a highly respected evolutionary geneticist at the University of Chicago — concluded in his review of Majerus's book in the 5 November 1998 issue of Nature that "for the time being, we must discard Biston as a well-understood example of natural selection in action, although it is clearly a case of evolution." Coyne's words carried weight, and the anti-evolutionists were quick to twist them into an argument against natural selection itself. "Coyne might have just thought he was stirring things up. ... As it happened, he was the lightning conductor," observes Mark Ridley, an evolutionary biologist at the University of Oxford. Coyne thought the flaws in Kettlewell's experiments were sufficient to cast doubt on the idea that bird predation was the agent of natural selection in this case, although he says his "biological intuition is that predation is probably a major cause."

Majerus dismisses most critics of the bird predation hypothesis as being people who write about the peppered moth without having a "feel for the organism." Many biologists start with theories and then test them in the field. Majerus takes the opposite and less fashionable approach: "There's another way of doing science, which is first find your organism and look at it incredibly closely, for a long time, in great detail and see what questions it asks you," he says.

Colleagues describe Majerus as a brilliant natural historian and a great communicator. He's made numerous appearances on radio, on television, and in the popular press, and he's on the circuit as an after-dinner speaker. "He doesn't have to try very hard to get a lot of people interested in what he's saying," says former graduate student Matt Tinsley. "Mike likes having a small crowd of students around him and telling stories."

It's a talent Majerus hopes to put to good use in defending the reputation of Kettlewell and the peppered moth in a road show, which he aims to take around Britain — and possibly the United States — later this year. He is motivated by growing concern over attacks on Kettlewell's character, most notably writer Judith Hooper's scathing account of the men behind the peppered moth story in her 2002 book Of Moths and Men: The Untold Story of Science and the Peppered Moth, which helped fuel an anti-evolutionist campaign to remove Biston from school textbooks. "A lot of [the campaign] is pointed at the peppered moth as being the example that Darwinism is debunked," says Majerus, who wants to make a public stand against teaching creationism and "intelligent design" in biology classes. "To have people believe the biology of the planet is controlled by a Creator, I think that's dangerous."

After decades of moth-watching, Majerus is convinced that Kettlewell was right and that bird predation is the primary agent of natural selection on the peppered moth. "But that can never be enough," he says, "because I'm also a scientist. ... We're miles beyond reasonable doubt, but it's not scientific proof."

Majerus's experiment is designed to avoid the mistakes Kettlewell made when comparing the proportion of typical and melanic peppered moths that escape the attention of predatory birds. He's releasing a small number of moths, at night, and letting them choose their own hiding places within specially designed mesh sleeves, which he removes at dawn. Like Kettlewell, he's using a mixture of lab-reared and wild-caught moths, but his design allows him to test for potential differences between the two. Majerus is determined to get "a definite answer" on the bird predation issue.

Although Majerus expects to confirm Kettlewell's conclusions, he claims not to care which way the results go: Any findings, he thinks, would make a splash by settling the controversy. But peppered moth expert and evolutionary geneticist Bruce Grant of the College of William and Mary in Williamsburg, Virginia, doubts that Majerus will silence the critics. "To do the job the right way is going to be too labor-intensive and it's just not worth it. ... Right now, I think there are other things that need doing more."

Time is running out for studying the melanic peppered moth, which, with declining pollution levels, is expected to make up only 1% of the British peppered moth population by 2019. And for Majerus, there are other fish to fry: "I don't want to get stuck with peppered moths for the rest of my career," he says, but he doesn't see his bug obsession waning anytime soon.

–Fiona Proffitt

 

[jack]

sd

"Michael Majerus's fascination for "bugs," as he calls all insects, was ignited at the tender age of 4. His mother can recall the exact moment: It was late summer, and he caught his first butterfly — a red admiral resting on a white chrysanthemum — with his bare hands. From that point on, he was hooked"

In Defense of Darwin and A Former Icon of Evolution
(Science, 304:1894-5, 2004)​

Michael Majerus's fascination for "bugs," as he calls all insects, was ignited at the tender age of 4. His mother can recall the exact moment: It was late summer, and he caught his first butterfly — a red admiral resting on a white chrysanthemum — with his bare hands. From that point on, he was hooked. At night when other children were in bed, the young Majerus roamed the English countryside tending his moth traps. Now 50 and in his 25th year of teaching evolutionary genetics at the University of Cambridge, Majerus still runs his moth traps most nights.

Majerus's research has focused on sexual selection, sex-ratio manipulation, and the evolution of melanism (the darkening of body color) in various moths, butterflies, and ladybirds. But over the past few years, half of his working life has been occupied by one controversial species, the peppered moth, Biston betularia, and an infamous study that's been attacked by both evolutionary biologists and anti-evolutionists.

Through his research, Majerus found himself embroiled in the scientific debate over the evolutionary forces behind melanism in the peppered moth. Experiments by British lepidopterist Bernard Kettlewell in the 1950s claimed to show that bird predation, coupled with pollution, was responsible for a color shift in the moth population. But problems with Kettlewell's methodology led some scientists to doubt his conclusions. Majerus was not the first to point out the flaws, but by doing so, he inadvertently set off a wave of anti-evolutionist attacks. While acknowledging that Kettlewell made mistakes, Majerus believes Kettlewell was right in his conclusions and has taken it upon himself to prove it.

As Majerus shows off some of the roughly 100,000 peppered moth pupae he'll rear for his latest experiment, it's clear that he's prepared to go to great lengths to make his case. Once the moths begin to emerge in May, Majerus begins a daily grind. He releases them at dusk and gets up at dawn to observe their fate: counting how many are plucked from their resting places by birds, and how many survive to see another night. He will continue this routine into August, as he has done for the past three summers. All he needs, he reckons, is another 2 years' worth of data — a total of some 4000 moth observations — to settle the controversy over whether bird predation is the major selective force in favoring one color form of the peppered moth over another.

Small and unobtrusive, the peppered moth doesn't look like the star in an evolutionary drama. But the rise and fall of the almost-black melanic form (carbonaria) in tandem with changing pollution levels has become the most famous example of evolution in action. Through his pioneering experiments, Kettlewell claimed to have demonstrated that melanic peppered moths were more common in industrialized areas because they escaped the attention of predatory birds when resting against soot-blackened, lichen-free bark. Because more of the darker ones survived to produce the next generation, he argued, entire populations grew darker.

But doubts emerged over Kettlewell's methodology in recent decades as researchers failed to replicate some of his results. His predation experiments were chiefly criticized for their artificiality: He placed the moths on exposed parts of trees in broad daylight, when they don't normally fly, rather than allowing them to settle naturally; he released them in large numbers, thereby inflating moth densities and possibly creating a magnet for predatory birds; and he used a mixture of lab-reared and wild-caught moths without checking to see whether they behaved the same way. Majerus summarized these criticisms in a book on the evolution of melanism in 1998 and stated that the simplified textbook story of the peppered moth was inaccurate, while asserting that Kettlewell's conclusions were qualitatively sound. Majerus had no idea at the time what a furor his book would cause.

Jerry Coyne — a highly respected evolutionary geneticist at the University of Chicago — concluded in his review of Majerus's book in the 5 November 1998 issue of Nature that "for the time being, we must discard Biston as a well-understood example of natural selection in action, although it is clearly a case of evolution." Coyne's words carried weight, and the anti-evolutionists were quick to twist them into an argument against natural selection itself. "Coyne might have just thought he was stirring things up. ... As it happened, he was the lightning conductor," observes Mark Ridley, an evolutionary biologist at the University of Oxford. Coyne thought the flaws in Kettlewell's experiments were sufficient to cast doubt on the idea that bird predation was the agent of natural selection in this case, although he says his "biological intuition is that predation is probably a major cause."

Majerus dismisses most critics of the bird predation hypothesis as being people who write about the peppered moth without having a "feel for the organism." Many biologists start with theories and then test them in the field. Majerus takes the opposite and less fashionable approach: "There's another way of doing science, which is first find your organism and look at it incredibly closely, for a long time, in great detail and see what questions it asks you," he says.

Colleagues describe Majerus as a brilliant natural historian and a great communicator. He's made numerous appearances on radio, on television, and in the popular press, and he's on the circuit as an after-dinner speaker. "He doesn't have to try very hard to get a lot of people interested in what he's saying," says former graduate student Matt Tinsley. "Mike likes having a small crowd of students around him and telling stories."

It's a talent Majerus hopes to put to good use in defending the reputation of Kettlewell and the peppered moth in a road show, which he aims to take around Britain — and possibly the United States — later this year. He is motivated by growing concern over attacks on Kettlewell's character, most notably writer Judith Hooper's scathing account of the men behind the peppered moth story in her 2002 book Of Moths and Men: The Untold Story of Science and the Peppered Moth, which helped fuel an anti-evolutionist campaign to remove Biston from school textbooks. "A lot of [the campaign] is pointed at the peppered moth as being the example that Darwinism is debunked," says Majerus, who wants to make a public stand against teaching creationism and "intelligent design" in biology classes. "To have people believe the biology of the planet is controlled by a Creator, I think that's dangerous."

After decades of moth-watching, Majerus is convinced that Kettlewell was right and that bird predation is the primary agent of natural selection on the peppered moth. "But that can never be enough," he says, "because I'm also a scientist. ... We're miles beyond reasonable doubt, but it's not scientific proof."

Majerus's experiment is designed to avoid the mistakes Kettlewell made when comparing the proportion of typical and melanic peppered moths that escape the attention of predatory birds. He's releasing a small number of moths, at night, and letting them choose their own hiding places within specially designed mesh sleeves, which he removes at dawn. Like Kettlewell, he's using a mixture of lab-reared and wild-caught moths, but his design allows him to test for potential differences between the two. Majerus is determined to get "a definite answer" on the bird predation issue.

Although Majerus expects to confirm Kettlewell's conclusions, he claims not to care which way the results go: Any findings, he thinks, would make a splash by settling the controversy. But peppered moth expert and evolutionary geneticist Bruce Grant of the College of William and Mary in Williamsburg, Virginia, doubts that Majerus will silence the critics. "To do the job the right way is going to be too labor-intensive and it's just not worth it. ... Right now, I think there are other things that need doing more."

Time is running out for studying the melanic peppered moth, which, with declining pollution levels, is expected to make up only 1% of the British peppered moth population by 2019. And for Majerus, there are other fish to fry: "I don't want to get stuck with peppered moths for the rest of my career," he says, but he doesn't see his bug obsession waning anytime soon.

–Fiona Proffitt

 

[Grammour Boy]

FORMIDABLE!

 

[Grammour Boy]

Up close and personal with the Red Admiral at dinner time: Awesome!!

 

[Tyrant]

 

[Tyrant]