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.     

Reproductive strategies of female butterflies: variation in and constraints on fecundity

ABSTRACT.

• 1 This study first examines the reproductive strategy of female Speyeria mormonia Edwards (Lepidoptera: Nymphalidae):
• 2 Egg weight and number laid per day decrease with age.
• 3 Survival and daily egg number may be affected by temperature; mean daily egg weight is not affected by temperature.
• 4 Daily egg number is not correlated with body size. In the central range of body size, egg weight is also not correlated with body size. However, exceptionally large or small females lay respectively heavier or lighter eggs than average.
• 5 A simple trade-off between offspring size and number does not occur within females on a daily basis, or among females averaged over their lifespans.
• 6 Fat body resources are depleted at a rate independent of body size.
• 7 Females are essentially monogamous.
• 8 Age-specific fecundity data reported here for S.mormonia are next compared with data for other Lepidoptera with different adult feeding habits and egg maturation patterns, and hence different possibilities for adult feeding to play a role in egg production. Based on these comparisons, I propose that the shape of the age-specific fecundity curve for each species under optimal conditions is constrained by the potential importance of adult nutrients in egg production.

     

Induction of reproductive diapause in Bruchidius atrolineatus during the dry season in a Sahelian zone

Bruchidius atrolineatus (Pic) (Bruchidae: Coleoptera) is a bruchid beetle infesting Vigna unguiculata (Walp). During the dry season, adults are in reproductive diapause. Studies of adult populations in a traditional store containing contaminated pods of V. unguiculata show that the reproductive diapause is induced during the first larval instars. The temperature decrease from November to January probably is the main climatic factor inducing this diapause. The percentage of diapausing adults depends on the duration of the larval development at low temperatures. Experiments in climatic boxes confirm these results. At 12 h 25 °T-12 h 20° C 11–13 LD, a high proportion of the adults emerging from the V. unguiculata seeds are in reproductive diapause and the duration of their growth is variable. L1 L2 L3 larvae probably are sensitive stages to the thermal variations and the processes allowing emergence of diapausing adults probably are induced during these larval instars.

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Résumé Bruchidius atrolineatus (Pic) est un Coléoptère Bruchidae se développant en zone sahélienne aux dépens d'une légumineuse Vigna unguiculata. Pendant la saison sèche, les adultes sont en diapause reproductrice. Un suivi des populations d'adultes a été réalisé dans un grenier traditionnel en terre séchée contenant des gousses récoltées dans la nature et portant des ufs de B. atrolineatus. Il a permis de constater que la diapause reproductrice était induite durant les premières phases du développement larvaire. La décroissance des températures de Novembre à Janvier avec des minima de 15 ° C et des maxima de 25 °C est probablement le principal facteur permettant l'induction de cette diapause. L'introduction dans le grenier de gousses portant des ufs venant d'être émis à différentes périodes de la saison sèche montre que le pourcentage d'adultes diapausants dépend de la durée du développement larvaire aux températures basses. Des expériences réalisées dans des étuves programmées confirment ces résultats. A 12 h 25 °T/12 h 20 °C 11–13 LD, un taux élevé d'adultes émergeant des graines est en diapause reproductrice et la durée de leur développement post-embryonnaire est très variable. En conditions plus chaudes 12 h 40 °C/12 h 25 °C 11–13 LD, le taux d'adultes diapausants est plus réduit et la durée du développement plus courte. L'analyse de la croissance larvaire montre que les premiers stades larvaires sont probablement des stades sensibles aux variations thermiques. Les modifications physiologiques permettant l'émergence d'adultes diapausants ou sexuellement actifs seraient induites durant ces stades.

     

Life history strategies and habitat templets of tropical butterflies in north-eastern Australia

Three multivoltine species of satyrine butterflies in the genus Mycalesis (Lepidoptera: Nymphalidae) are narrowly sympatric in the wet–dry tropics of north-eastern Australia. They show a range of ecological strategies and adaptations associated with contrasting habitats and varying selective pressures. Two abiotic factors, namely favorability (the reciprocal of seasonal adversity) and predictability (broadly the reciprocal of disturbance), were examined as potential environmental selective forces in shaping their life histories. Comparison of several key life history traits of the wet-season form revealed that the life histories of each species corresponded well with their habitat characteristics. M. perseus, which lives in habitats which are less favorable (i.e. adverse) and more unpredictable (i.e. temporary), shows many traits of an r-type strategy: smaller size, faster development, earlier maturation, higher fecundity, smaller egg size, and rapid population increase. By contrast, M. sirius and M. terminus, which live in more favorable and predictable (i.e. permanent) habitats, have many life history attributes and other characteristics in common which link them closer to K-type strategies. The only discrepancy is lower potential reproductive effort of M. perseus, which may be accounted for in terms of an evolutionary trade-off, such as with dispersal or dormancy. Other correlates associated with the M. perseus life history tactic include higher sex-size dimorphism, greater dispersal ability, better tolerance to adverse conditions, stronger phenotypic variation, greater degree of polyandry, and a more flexible breeding strategy. The life history patterns of these species are discussed in the context of evolutionary life history models, particularly the Southwood–Greenslade habitat templet.     

Patterns of tree growth in relation to environmental variability in the tropical dry deciduous forest at Mudumalai, southern India

Tree diameter growth is sensitive to environmental fluctuations and tropical dry forests experience high seasonal and inter-annual environmental variation. Tree growth rates in a large permanent plot at Mudumalai, southern India, were examined for the influences of rainfall and three intrinsic factors (size, species and growth form) during three 4-year intervals over the period 1988–2000. Most trees had lowest growth during the second interval when rainfall was lowest, and skewness and kurtosis of growth distributions were reduced during this interval. Tree diameter generally explained <10% of growth variation and had less influence on growth than species identity or time interval. Intraspecific variation was high, yet species identity accounted for up to 16% of growth variation in the community. There were no consistent differences between canopy and understory tree growth rates; however, a few subgroups of species may potentially represent canopy and understory growth guilds. Environmentally-induced temporal variations in growth generally did not reduce the odds of subsequent survival. Growth rates appear to be strongly influenced by species identity and environmental variability in the Mudumalai dry forest. Understanding and predicting vegetation dynamics in the dry tropics thus also requires information on temporal variability in local climate.     

Mast fruiting of large ectomycorrhizal African rain forest trees: importance of dry season intensity, and the resource-limitation hypothesis

Mast fruiting is a distinctive reproductive trait in trees. This rain forest study, at a nutrient-poor site with a seasonal climate in tropical Africa, provides new insights into the causes of this mode of phenological patterning. At Korup, Cameroon, 150 trees of the large, ectomycorrhizal caesalp, Microberlinia bisulcata, were recorded almost monthly for leafing, flowering and fruiting during 1995-2000. The series was extended to 1988-2004 with less detailed data. Individual transitions in phenology were analysed. Masting occurred when the dry season before fruiting was drier, and the one before that was wetter, than average. Intervals between events were usually 2 or 3 yr. Masting was associated with early leaf exchange, followed by mass flowering, and was highly synchronous in the population. Trees at higher elevation showed more fruiting. Output declined between 1995 and 2000. Mast fruiting in M. bisulcata appears to be driven by climate variation and is regulated by internal tree processes. The resource-limitation hypothesis was supported. An 'alternative bearing' system seems to underlie masting. That ectomycorrhizal habit facilitates masting in trees is strongly implied.     

Daylength and humidity as environmental cues for diapause termination in a tropical beetle

ABSTRACT. Photoperiod is the primary factor controlling termination of adult diapause in the fungus beetle, Stenotarsus rotundus Arrow (Endomychidae: Coleoptera), in a tropical lowland forest. at 9 N. An LD 13:11 h cycle stimulated rapid development of the gonads and flight muscles in both sexes: development was eventually initiated in an LD 12:12 h cycle but was slower. Beetles showed a graded response to intermediate photoperiods. Males consistently initiated gonad development earlier than females, but both sexes developed flight muscles at the same time.
Moisture also influences development: oocytes and flight muscles developed more rapidly at high humidity than at the naturally prevailing low humidity during the dry season. Testis development was less influenced by humidity.
In the field, male gonad development began in late January, suggesting that the beetle can perceive an extremely slight increase in daylength. Ovaries did not contain any visible oocytes before March, but oocyte development was apparent in all females by April. Egg maturation was completed after dispersal from the aggregation site. Females have larger corpora allata than males. These glands increased in size in March when oocyte and/or flight muscle development was initiated. The timing of full development of flight muscles varied slightly from year to year.
Thus, the increase in daylength occurring during the dry season appears to be the cue primarily responsible for development of the reproductive systems and flight muscles. High humidity enhances this response and further prepares the beetles for an active reproductive period at the onset of the wet season.     

The role of tropical dry forest as a long-term barrier to dispersal: a comparative phylogeographical analysis of dry forest tolerant and intolerant frogs

We used a comparative phylogeographical approach to investigate the origins of the disjunct wet forest biota of the Golfo Dulce region along the Pacific slope of Costa Rica. This region is isolated by Pacific dry forests north and south and isolated from Caribbean wet forests by mountains. We studied three sympatric lowland frog species in the Craugastor fitzingeri species group that prefer wet forest but differ in their response to dry habitats. In dry forest, C. fitzingeri can survive along streams while C. crassidigitus and C. talamancae are entirely absent. We collected samples from across the ranges of all three species, and obtained mitochondrial DNA sequence data from the COI and cytochrome b genes. We observed significant phylogeographical structure in C. crassidigitus and C. talamancae, but much less in C. fitzingeri, demonstrating that mountain barriers and dry forest habitat have reduced mitochondrial gene flow in the strictly wet-forest species. Additionally, we discovered that the Golfo Dulce and Central Panama populations of C. crassidigitus appear to have diverged in the Pliocene or earlier, suggesting that the dry forest separating these populations is old. Our phylogenetic analysis of 12 of approximately 16 species of the C. fitzingeri species group suggests that the three lowland species are each other's closest relatives. Because of this shared phylogenetic history, we attribute the striking differences in phylogeographical structure to the different ecologies of the frogs. In summary, we find that what appear to be minor differences in the natural history of these three closely related species may profoundly impact the potential for dispersal, range size, and cladogenesis.     

Pulses of soil phosphorus availability in a Mexican tropical dry forest: effects of seasonality and level of wetting

Intact cores from the upper soil profile and surface litter were collected at the peak of the dry season and during the rainy period in the tropical deciduous forest of the Chamela region, Jalisco, México, to (1) analyze upper soil phosphorus (P) movement and retention, (2) compare soil P dynamic pools (soluble, bicarbonate, and microbial) in dry and rainy seasons, and (3) determine the response of these P pools to wetting. Unperturbed litter-soil cores were treated in the laboratory with either 10 mm or 30 mm of simulated rain with carrier-free ³²P and compared to a control (no water addition) to determine the fate and retention of added P. ³¹P concentrations and pools in most litter and soil fractions were higher in the dry than in the rainy season. Soluble P was 0.306 g/m² and microbial P was 0.923 g/m² in the dry season (litter plus soil) versus 0.041 (soluble) and 0.526 (microbial) g P/m² in the rainy season. After water addition, rainy-season cores retained 99.9 and 94% of ³²P in the 10- and 30-mm treatments, respectively. Dry-season samples retained 98.9 and 80% of inputs in the same treatments. Retention after wetting occurred mostly in soil (bicarbonate and microbial fractions). Simulated rainfall on rainy-season soils increased P immobilization. On the other hand, simulated rainfall on dry-season soils released P through mineralization. The P release represents between 46 and 99% of the annual litterfall return. Our results suggest that both soluble and microbial P constitute important sources for initiation of plant growth at the onset of the rainy season in tropical dry forest. Received: 23 September 1997 / Accepted: 2 February 1998     

Adaptive strategies of overwintering adults: Reproductive diapause and mating behavior in a grasshopper,Stenocatantops splendens (Orthoptera: Catantopidae)

Abstract To understand the adaptive strategies of the overwintering adults of Stenocatantops splendens, the mechanism of maintenance and termination of the reproductive diapause, the variation in mortality between overwintering females and males, and the mating strategy of the males were investigated. The results indicated that the adult reproductive diapause in natural conditions was mainly regulated by photoperiod in the fall – long photoperiods promoted reproductive development and short photoperiods maintained reproductive diapause, and the sensitivity of the overwintering adults to photoperiod was over before the end of the winter. When transferred from natural conditions to controlled laboratory conditions on dates from September through February, pre‐oviposition became increasingly shorter with increasingly deferred transfer dates regardless of photoperiod conditions. The adults treated with low temperature for 30 days in September through November had significantly shorter pre‐oviposition, suggesting that low temperatures in winter had an important role in the termination of reproductive diapause. The female had a significantly lower supercooling point than the male, which was related to their lower mortality after winter. In addition, observations of wild populations of the species indicated that mating behavior prior to winter and the duration of pre‐mating period were not affected by photoperiod; mating and sperm transfer were mostly completed by November. Compared with females only mating before winter, females mating in the spring had shorter life span, longer pre‐oviposition, lower hatching rate and laid fewer egg pods while showing no significant difference with regard to ovipositional interval, per pod number of eggs and nymph dry weight.     

Carbon sequestration in tropical forests and water: a critical look at the basis for commonly used generalizations

Tree planting in the tropics is conducted for a number of reasons including carbon sequestration, but often competes with increasingly scarce water resources. The basics of forest and water relations are frequently said to be well understood but there is a pressing need to better understand and predict the hydrological effects of land‐use and climate change in the complex and dynamic landscapes of the tropics. This will remain elusive without the empirical data required to feed hydrological process models. It is argued that the current state of knowledge is confused by too broad a use of the terms ‘forest’ and ‘(af)forestation’, as well as by a bias towards using data generated mostly outside the tropics and for nondegraded soil conditions. Definitions of forest, afforestation and reforestation as used in the climate change community and their application by land and water managers need to be reconciled.     

Plant biodiversity of two tropical dry evergreen forests in the Pondicherry region of South India and the role of belief systems in their conservation

Natural vegetation on the south-eastern coast of Peninsular India has now been reduced to patches, some of which are preserved as sacred groves. The plant biodiversity and population structure of woody plants (>20 cm girth at breast height; gbh) in two such groves, Oorani and Olagapuram, occurring on the north-west of Pondicherry have been analyzed. A total of 169 angiosperms have been enumerated from both sites. The Oorani grove (3.2 ha) had 74 flowering plant species distributed in 71 genera and 41 families; 30 of them are woody species, 8 are lianas and 4 are parasites. The Olagapuram grove (2.8 ha) was more species-rich with 136 species in 121 genera of 58 families; woody species were fewer (21) while 9 lianas and 3 parasites occurred. The vegetation structure indicates that the Oorani grove is a relic of tropical dry evergreen forest, whereas Olagapuram is reduced to a thorny woodland. The latter is heavily degraded as it has lost the status of a sacred grove because of its conversion to Eucalyptus plantations. The Oorani grove has an Amman temple in the centre. The attendant cultural rites and religious rituals have perpetuated the status of a sacred grove which has ensured the protection of the grove.     

Individual constancy of local search strategies in the giant tropical ant,Paraponera clavata (Hymenoptera: Formicidae)

Paraponera clavata workers engage in a period of local search after encountering a small amount of artificial nectar. Giving-up times from local search are not distributed normally; there is a strong skew to longer times. There is no statistically significant relationship between the amount of time required to collect the food and the subsequent search time. Giving-up time in response to the first reward presented to an ant is positively correlated with that ant's response to a second such reward. However, giving-up times diminish when an ant is presented with a series of rewards. Local search is a function of individual strategies, which remain relatively constant in the short term.     

Feedbacks between phosphorus deposition and canopy cover: The emergence of multiple stable states in tropical dry forests

Dry forests represent a large percentage of tropical forests and are vulnerable to both anthropogenic and natural disturbances, yet important aspects of their sensitivity to disruption remain poorly understood. It is particularly unclear how changes in land-use or tropical storm patterns may affect the resiliency of phosphorus (P)-limited neotropical forests. In these systems, vegetation is sustained in the long-term by atmospheric P-inputs through rainfall, dust, or fog. Past research supports the idea that dust and fog deposition are dependent on canopy density (e.g. leaf area index). Thus, the canopy may function as a 'trap' for P, enabling a positive feedback between vegetation and P-deposition. We developed a conceptual model to investigate how Neotropical vegetation may respond to reduced P-deposition due to canopy losses. The model suggests that a canopy-deposition feedback may induce bistable vegetation dynamics; under some conditions, forests may be unable to naturally recover from relatively small disturbances.     

Fire history influences on the spatial heterogeneity of soil nitrogen transformations in three adjacent stands in a dry tropical forest in Thailand

Spatial patterns of soil nitrogen (N) transformations were examined using geostatistical analysis in three adjacent stands with different fire history (0, 10 and 35 years since the latest fire, respectively) in a dry tropical forest in Thailand. A larger pool of total inorganic N and a faster rate of N mineralization were recorded in the stand with longer fire prevention. At the spatial scale analyzed, the proportion of spatially dependent variance to the total variance of N mineralization and nitrification increased from 0.39 to 0.73, and from 0.40 to 0.77, respectively, with the time since the latest fire. The spatial autocorrelation ranges of N mineralization and nitrification decreased from 9.0 to 3.28 m, and 9.0 to 2.77 m, respectively, with the time since the latest fire. These results suggested that fire history affected not only the level of available soil N, but also the spatial heterogeneity of soil N transformations, presumably due to the difference in plant influences on soil.     

Impact of severe dry season on net ecosystem exchange in the Neotropical rainforest of French Guiana

The lack of information on the ways seasonal drought modifies the CO₂ exchange between Neotropical rainforest ecosystems and the atmosphere and the resulting carbon balance hinders our ability to precisely predict how these ecosystems will respond as global environmental changes force them to face increasingly contrasting conditions in the future. To address this issue, seasonal variations in daily net ecosystem productivity (NEP_d) and two main components of this productivity, daily total ecosystem respiration (R_Ed) and daily gross ecosystem productivity (GEP_d), were estimated over 2 years at a flux tower site in French Guiana, South America (5 °16′54″N, 52 °54′44″W). We compared seasonal variations between wet and dry periods and between dry periods of contrasting levels of intensity (i.e. mild vs. severe) during equivalent 93‐day periods. During the wet periods, the ecosystem was almost in balance with the atmosphere (storage of 9.0 g C m^?2). Seasonal dry periods, regardless of their severity, are associated with higher incident radiation and lower R_Ed combined with reduced soil respiration associated with low soil water availability. During the mild dry period, as is normally the case in this region, the amount of carbon stored in the ecosystem was 32.7 g C m^?2. Severe drought conditions resulted in even lower R_Ed, whereas the photosynthetic activity was only moderately reduced and no change in canopy structure was observed. Thus, the severe dry period was characterized by greater carbon storage (64.6 g C m^?2), emphasizing that environmental conditions, such as during a severe drought, modify the CO₂ exchange between Neotropical rainforest ecosystems and the atmosphere and potentially the resulting carbon balance.