Reproductive seasonality in tropical satyrine butterflies: strategies for the dry season

Abstract.

• 1 In tropical savanna environments rainfall is often very seasonal, so that much of the year is characterized by a long and unpredictable dry season. Because the timing and availability of rain exerts a major influence on plant growth and production, many species during the dry period exhibit dramatic reduction in leaf quality. Accordingly, and kind of behaviour shown by phytophagous insects that synchronizes larval feeding with food availability will be adaptive.
• 2 The reproductive status of three Mycalesis butterflies was monitored over a 2-year period (1989–90) in north-eastern Queensland, Australia, at a lowland site (Cardwell, 18°16's, 146°02′E) which experiences a pronounced dry season. Females of these species and of five other satyrines (Ypthima, Hypocysta spp.) were also examined less intensively during the dry season in areas throughout northern and central Queensland, north of the tropic of Capricorn.
• 3 These relatively sedentary butterflies exhibit three different strategies for dealing with the unpredictable dry period and associated deterioration of larval food plants (grass). First, five species appear to breed continuously, though for most reproductive activity (mature egg number) declines markedly in the late dry season. Two of these (Hypocysta irius, H.metirius) are restricted to less seasonal and more favourable (wetter) areas but the three others (Ypthima arctous, H.adiante, H.pseudirius) occur widely in the relatively dry savanna, where they may specialize on grass in moister microenvironments. Second, two species (M.terminus, M.sirius) live in predictably moist habitats which are buffered from climatic extremes; they breed for much of the season but reproductive activity declines as the dry season progresses and may cease late in the season. Third, one species (M.perseus) is more opportunistic, breeding for only a limited interval during the favourable (wet) periods; during the long dry season adults contract to moist refugia and remain in reproductive diapause.
• 4 Spending the late dry season as an adult, either in diapause or with mature eggs, may improve the capacity to utilize new growth of grasses at the start of the favourable season, thereby enhancing population growth during good times. It may also provide additional flexibility to counter the temporal uncertainty of the dry season.
• 5 The strategy of residing in more equitable habitats or specializing on predictable foods may be the most restrictive in terms of distribution.

     

Host plant discrimination in tropical satyrine butterflies

Summary Inter and intraspecific host plant choice was examined for Mycalesis terminus and M. perseus, two tropical butterfly species of the subfamily Satyrinae. The two species preferred different suites of potential host plant species, but only M. perseus discriminated between different qualities of host. Differences in selectivity between these two species may be associated with the different lengths of time over which their larval resources may be expected to persist. The level of selectivity exhibited by these tropical species appears to be greater than their temperate counterparts. Potential factors controlling differences between oviposition strategies in temperate and tropical Satyrine butterflies and other groups are discussed.     

Habitat preference and habitat exploration in two species of satyrine butterflies

We compared the habitat preference and the mobility of different populations of primarily forest living, satyrine butterflies by observing them in a manipulated environment, containing different habitat types inside a large outdoor cage. The cage was 30 m long and divided into three parts. Each end of the cage contained a shady habitat, made to be similar to the floor of a forest with sunlit spots. In between these shady parts, there was a 14 m long open part, like a glade or a meadow. We performed two studies in the cage. In the first study, we compared a population of a species with very restricted distribution in Sweden, Lopinga achine , and a population of a widely distributed and expanding species, Pararge aegeria . We found that the two species had similar habitat preferences in the cage, but that L. achine had a lower rate of dispersal from one shady part of the cage to the other. In the second study, we compared two populations of P. aegeria , a Swedish forest living population and a Madeiran population inhabiting a fairly open habitat. We found that, although individuals of the Swedish population avoided staying in the open, middle part of the cage, they dispersed between the shady parts of the cage at a higher rate than individuals of the Madeiran population. We argue that studies of the kind we performed can serve the purpose of testing hypotheses about habitat preference and mobility that derive from other field work or from information about distribution patterns.     

Monitoring of butterflies in semi-natural grasslands: diurnal variation and weather effects

The butterfly fauna was monitored in six semi-natural grasslands in southeastern Sweden. The aim was to evaluate monitoring criteria for wind, sunshine, temperature and time of day for butterfly species richness and abundances when using the line transect method. A total of 30,111 butterflies belonging to 46 species were recorded. Data from this study suggests somewhat stricter criteria for temperature and sunshine than stated in the widely used “Pollards walk”. A sharp decline in butterfly numbers were detected at temperatures below 19°C if the proportion of sunshine of the transect walk was below 80–85%. No effect of wind speed, up to five on the Beaufort scale, on observed numbers of species or individuals were found. Several butterfly species showed well-defined diurnal rythms of flight activity, and the results indicated that transect walks can be performed between −4.5 and +4 h from the time when the sun reached its highest point. The results of this study can be used to adjust general criteria in national monitoring and also for detailed regional and local monitoring where it may be important to adjust for diurnal rhythm and weather related bias.     

Spontaneous regeneration of dry grasslands on set-aside fields

Dry grasslands are among the most threatened plant communities of Central Europe. We explore the time scale of spontaneous regeneration of dry grasslands on abandoned fields in an area of Central Europe, where also ancient grassland communities occur (Saxony-Anhalt, Germany). On three permanent plots with shallow soils we monitored during 10 years change of species composition and analysed whether spontaneous succession leads to assemblages similar to the ancient dry grassland communities in the direct surroundings. We found that dry grassland species are able to invade the permanent plots and during the 10 years of succession the number of dry grassland species increased. But even after 10 years there was a clear difference between ancient dry grassland communities and the assemblages on the permanent plots. Our findings suggest two important conclusions: First, spontaneous succession on abandoned fields is a cheap possibility for the conservation of some dry grassland species, at least on shallow soils. Second, the time scale of the regeneration process, however, is rather long. Hence, conservation of remnants of ancient grassland communities needs special attention.     

Effects of climatic change on the phenology of butterflies in the northwest Mediterranean Basin

Phenological changes in response to climatic warming have been detected across a wide range of organisms. Butterflies stand out as one of the most popular groups of indicators of climatic change, given that, firstly, they are poikilothermic and, secondly, have been the subject of thorough monitoring programmes in several countries for a number of decades. Here we provide for the first time strong evidence of phenological change as a consequence of recent climatic warming in butterflies at a Spanish site in the northwest Mediterranean Basin. By means of the widely used Butterfly Monitoring Scheme methodology, three different phenological parameters were analysed for the most common species to test for trends over time and relationships with temperature and precipitation. Between 1988 and 2002, there was a tendency for earlier first appearance dates in all 17 butterfly species tested, and significant advances in mean flight dates in 8 out of 19 species. On the other hand, the shape of the curve of adult emergence did not show any regular pattern. These changes paralleled an increase of 1–1.5°C in mean February, March and June temperatures. Likewise, a correlation analysis indicated the strong negative effect of spring temperature on phenological parameters (i.e. higher temperatures tended to produce phenological advances), and the opposite effect of precipitation in certain months. In addition, there was some evidence to indicate that phenological responses may differ between taxonomic lineages or species with similar diets. We discuss the consequences that these changes may have on species' population abundances, especially given the expected increase in aridity in the Mediterranean Basin caused by current climatic warming. We predict that varying degrees of phenological flexibility may account for differences in species' responses and, for multivoltine species, predict strong selection favouring local seasonal adaptations such as diapause phenomena or migratory behaviour.     

Canopy phenology of a dry forest in western Brazil.

Dry forests are common, although highly threatened in the Neotropics. Their ecological processes are mostly influenced by rainfall pattern, hence their cycles exhibit contrasting phases. We studied the phenology of canopy trees in a primary dry forest in Western Brazil in the foothills of the Urucum mountain chain, in order to improve our knowledge on the functioning of these poorly-known forests. Leaf shedding started in the early dry season and was massive in the latter part of this period. Most leaf loss occurred in dry hills, while wet valleys remained evergreen. Anemochorich and autochorich species predominated in dry hills, presumably due to their tolerance to dry conditions and enhanced exposition to winds, which favour diaspores removal and dispersal. Conversely, zoochorich species dominated the wet valleys. Flowering was intense in the late dry season, the driest period of the year, while fruiting was massive just after the onset of rains, as well as flushing. Therefore, most flowering was unrelated to wet conditions, although such an abiotic factor, potentially, triggered the major fruiting episode, widely comprised by zoochorich species. Anemochorich and autochorich species flowered and fruited in the course of the long dry season. The contrasting environmental conditions present in the hills and valleys determine the arrangement of a mosaic in which patches of zoochorich and evergreen trees alternate with patches of non zoochorich and highly deciduous species. Consequently, species with such syndromes exhibited marked flowering and fruiting patterns, accordingly to the pronounced seasonality.     

Comparative unpalatability of mimetic viceroy butterflies (Limenitis archippus) from four south-eastern United States populations

Viceroy butterflies (Limenitis archippus), long considered palatable mimics of distasteful danaine butterflies, have been shown in studies involving laboratoryreared specimens to be moderately unpalatable to avian predators. This implies that some viceroys are Müllerian co-mimics, rather than defenseless Batesian mimics, of danaines. Here, I further test this hypothesis by assessing the palatability of wild-caught viceroys from four genetically and ecologically diverse populations in the southeastern United States. Bioassays revealed that viceroys sampled from three sites in Florida and one in South Carolina were all moderately unpalatable to captive redwinged blackbird predators, which ate fewer than half of the viceroy abdomens presented. Red-wings commonly exhibited long manipulation times and considerable distress behavior when attempting to eat a viceroy abdomen, and they taste-rejected over one-third of viceroys after a single peck. These findings, the first based on wild-caught butterflies, support the hypothesis that the viceroy-danaine relationship in some areas represents Müllerian mimicry, prompting a reassessment of selective forces shaping the interaction. Moreover, considerable variation in palatability of individual viceroys, and in behavior of individual birds, contributes to the complexity of chemical defense and mimicry in this system.     

Dispersal of four fritillary butterflies within identical landscape

Both species-specific traits and landscape configuration, such as area and connectivity of habitat patches plus the character of uninhabitable matrix, affect animal movements in fragmented landscapes. Difficulties with disentangling species-specific and landscape effects have obscured comparisons among species, hindering the understanding of dispersal in metapopulations. To circumvent this complication, we performed a mark–recapture study of four related nymphalid butterflies within identical landscape and in single season. The studied species were three Melitaeinae checkerspots (Euphydryas aurinia, Melitaea athalia, Melitaea diamina) and one Argynnini fritillary (Brenthis ino). Applying the Virtual Migration model revealed that (1) except for mortality within habitat, model parameters differed from those found for the studied species elsewhere; (2) the three Melitaeinae species were more akin in movement parameters than the Argynnini representative (i.e., B. ino); (3) within Melitaeinae, differences between sexes were more prominent than differences among species; (4) Melitaeinae males left natal patches more readily than females, while the opposite applied to B. ino; (5) males of M. diamina and both sexes of B. ino exhibited highest values of dispersal mortality; (6) except for females of M. diamina and both sexes of B. ino, immigration and emigration scaled with area in females but not in males. Finding (1) demonstrates that geometry of habitat network affects mobility considerably and that transferring dispersal parameters across systems is unwarranted. Still, (2–6) demonstrate that within identical networks, related species follow similar dispersal patterns, suggesting that conservation scenarios suitable for a well-studied model species would suite related species as well.     

Reproductive phenology in Dendrocoelum lacteum (Turbellaria) in South Sweden

SUMMARY.

• 1 L 266 degree-days, above a threshold of 1.0°C, were required for cocoons from the triclad Dendrocoelum lacteum to hatch. The relation is described by ln D=6.12–1.17 1nr (D=number of days, T= temperature).
• 2 Laboratory and field cocoon deposition reached a maximum in late April; peaks of hatchling occurrence were 2 weeks later. The number of hatchlings per cocoon and eocoon viability declined as spring proceeded. Cocoon deposition ceased in June, coincident with regression of the reproductive organs of the adults.
• 3 Newly hatched triclads in the field are thought to avoid stones for their first weeks.
• 4 It is suggested that the earlier and more abundant offspring production in South Sweden than in Britain could be attributed to differences in spring temperature conditions.

     

Occurrence patterns of butterflies (Rhopalocera) in semi-natural pastures in southeastern Sweden

Occurrence patterns of butterflies (Rhopalocera) were analysed in relation to locality characteristics, and nested patterns were evaluated, based on a survey of 100 semi-natural pastures in the county of Östergötland, southeastern Sweden. Species richness of butterflies was positively related with locality-size, but not with the density of pastures in the surroundings. Species richness was lower in heavily grazed pastures compared with localities with a low or intermediate grazing pressure. This suggests that a uniformed, heavy grazing pressure should be avoided if butterflies are to be protected, even though such a management is favourable for many vascular plants. Out of 45 analysed species, 73% exhibited a significantly nested pattern, and species regarded to be sedentary had generally a more nested pattern. Several butterflies (such as Erynnis tages, Mellicta athalia and Pyrgus malvae) are possible to use as indicators of a relatively intact butterfly community. In a short-term perspective, the great bulk of butterflies confined to semi-natural pastures would be saved if management were adapted to the requirements of butterflies only at the localities richest in species. However, for conservation strategies to be successful over a longer time, whole landscapes harbouring the majority of the species and with a high density of semi-natural pastures should be selected and be given priority for conservation.     

Contrasting leaf phenology of woody species of dry tropical forest

We studied selected leaf traits [leaf area (LA), leaf water content (LWC), leaf fresh weight (LFW), leaf dry weight (LDW), specific leaf area (SLA) and chlorophyll content] of eight woody species (Shorea robusta, Buchanania lanzan, Diospyros melanoxylon, Lagerstroemia parviflora, Lannea coromandelica, Terminalia tomentosa, Holarrhena antidysenterica and Lantana camara) dominant at four sites in a dry tropical deciduous forest over complete two annual cycles (2008–2010). Our results showed that leaf traits varied across species (1.7–11.5 fold), months (1.2–1.5 fold) and sites (1.1–1.3 fold). However, leaf traits showed smaller variation between sites than between species. Leaf lifespan varied from 7 months (L. coromandelica) to 12 months (S. robusta). On the same sites, species differed in the length of deciduous period. The maximum LA, LDW, LFW and LWC were recorded for the semi-evergreen species, SLA for long-deciduous species and chlorophyll content for short-deciduous species, respectively. The coefficient of variation was maximum for LDW and minimum for chlorophyll content. Among the eight woody species, T. tomentosa exhibited the greatest LA, LDW, LFW and LWC. LA, LWC, LFW, LDW, SLA, LD, SD, MD & SE confirm.     

Recreating semi-natural grasslands: A comparison of four methods

Semi-natural grasslands and their species and populations are declining rapidly throughout Europe, bringing about a need for successful vegetation recreation methods. To maintain biodiversity and ecological services of semi-natural grasslands, we need more knowledge on the relative performance of different recreation methods. In a replicated experiment in western Norway, we evaluated two hay transfer methods (hard or light raking of local hay), sowing of local seeds and natural regeneration for recreating semi-natural grassland in a road verge. We compared treated trial plots with their respective donor plots (where hay and seeds were harvested) for three successive years by evaluating vegetation cover, species richness and species transfer rates, and vegetation dynamics analysed by Bray–Curtis compositional dissimilarity (BC) and GNMDS (Global Non-Metric Multidimensional Scaling) ordination. Vegetation cover at the trial site exceeded that of donor sites in three years. Transfer rates of common species were high for seed sowing and both hay transfer procedures. Species composition in trial plots for all three treatments became significantly more similar to donor plots, but was still relatively dissimilar after three years. Natural regeneration showed a different temporal pattern and also had a higher successional rate. The species composition of the other treatments followed the same trajectory toward the donor sites as revealed by GNMDS. We found relatively small differences between the two hay transfer methods and seed sowing. Transfer of local hay therefore appears to be a successful method of establishing local species when recreating semi-natural grasslands, and is generally cheaper than using commercial local seed mixtures.     

Effect of regional climate warming on the phenology of butterflies in boreal forests in Manitoba, Canada

We examined the effect of regional climate warming on the phenology of butterfly species in boreal forest ecosystems in Manitoba, Canada. For the period 1971-2004, the mean monthly temperatures in January, September, and December increased significantly, as did the mean temperatures for several concurrent monthly periods. The mean annual temperature increased ≈ 0.05°C/yr over the study period. The annual number of frost-free days and degree-day accumulations increased as well. We measured the response of 19 common butterfly species to these temperature changes with the date of first appearance, week of peak abundance, and the length of flight period over the 33-yr period of 1972-2004. Although adult butterfly response was variable for spring and summer months, 13 of 19 species showed a significant (P < 0.05) increase in flight period extending longer into the autumn. Flight period extensions increased by 31.5 ± 13.9 (SD) d over the study period for 13 butterfly species significantly affected by the warming trend. The early autumn and winter months warmed significantly, and butterflies seem to be responding to this warming trend with a change in the length of certain life stages. Two species, Junonia coenia and Euphydryas phaeton, increased their northerly ranges by ≈ 150 and 70 km, respectively. Warmer autumns and winters may be providing opportunities for range extensions of more southerly butterfly species held at bay by past climatic conditions.     

Effects of grass-encroachment on vegetation and soil in Dutch dry dune grasslands

The encroachment of tall grass species in open dune vegetation, as observed in the Dutch dry dune area, is considered unfavourable for nature conservation. The effects of grass-encroachment on the vegetation and the availability of light and its relation to nutrients were investigated through a comparative study of grass-dominated and open dune grassland plots at ten locations along the Dutch coast.Grass-dominated plots have a low species diversity and number of species, especially of mosses and lichens. This is associated with a high biomass and a low availability of light at soil surface. In addition, nutrient availability seems to be important. Root biomass and the amounts of both ectorganic and endorganic matter are significantly higher in grass-dominated plots, which may account for the higher nutrient uptake in the vegetation.It is likely that a grass-dominated system can maintain and consolidate itself because of the better competition for light and nutrients. The relevance of these results for restoration management is briefly discussed.     

The evolutionary significance of dry and wet season forms in some tropical butterflies

An explanation is developed for the adaptive significance of the forms characteristic of the dry season and wet season adult generations of some tropical butterflies. The divergent phenotypes are accounted for as adaptive responses to a shift in the optimum balance between dependence on the alternative (but interdependent) strategies of active anti-predator devices and of crypsis. The seasonal polyphenisms exhibited by the satyrines Melanitis leda (Fabricius) and Orsotrioena medus (Fabricius) and the nymphalid Junonia almana (Linné) are examined in detail. The wet season forms show prominent marginal eyespot patterns which are displayed at rest and function principally in the deflection of attacks by vertebrate predators. In contrast, the dry season forms show very small or no spots and are wholly cryptic. Wet season butterflies are more active in general than those in the dry season when aestivation behaviour is often observed. The alternative phenotypes represent responses to the differences in behaviour, environment and nature of predation. Reproductive success is optimized in each season by an interaction of phenotype and behaviour. The hypothesis must be tested in detail by an investigation of the behavioural ecology and population biology of particular species. It is argued that these seasonal polyphenisms provide examples of adaptations to a repeating pattern of changing environments where the mode of selection is, in broad terms, understood.     

Adult emergence phenology in checkerspot butterflies: the effects of macroclimate,topoclimate, and population history

The prediction of adult emergence times in insect populations can be greatly complicated by microclimatic gradients, especially in circumstances where distributions of juveniles along those gradients vary from year to year. To investigate adult emergence patterns in topographically heterogeneous habitats, we built a model of postdiapause development of the Bay checkerspot butterfly, Euphydryas editha bayensis. The model uses slope-specific insolation as the rate-controlling variable, and accounts for both solar exposure of the habitat and cloud cover. Instar-specific larval mass gains per unit of insolation were determined from mark-recapture experiments. A small correction for daily low temperatures was used to calibrate the model to five years of field data on larval mass. The model predicted mean mass of 90% of larval samples within 4 clear days over a 70–120 day growing season. The magnitude of spatial variation in emergence times across habitat slopes is greater than annual variation in emergence times due to yearly weather conditions. Historical variation (yearly shifts in larval distributions across slopes) is an important determinant of mean population emergence dates. All of these factors need to be considered in understanding adult emergence phenology in this butterfly and in other insects inhabiting heterogeneous thermal environments. Such an understanding can be useful in managing insect populations for both pest control and conservation.