Complex life cycles and the responses of insects to climate change

Many organisms have complex life cycles with distinct life stages that experience different environmental conditions. How does the complexity of life cycles affect the ecological and evolutionary responses of organisms to climate change? We address this question by exploring several recent case studies and synthetic analyses of insects. First, different life stages may inhabit different microhabitats, and may differ in their thermal sensitivities and other traits that are important for responses to climate. For example, the life stages of Manduca experience different patterns of thermal and hydric variability, and differ in tolerance to high temperatures. Second, life stages may differ in their mechanisms for adaptation to local climatic conditions. For example, in Colias, larvae in different geographic populations and species adapt to local climate via differences in optimal and maximal temperatures for feeding and growth, whereas adults adapt via differences in melanin of the wings and in other morphological traits. Third, we extend a recent analysis of the temperature-dependence of insect population growth to demonstrate how changes in temperature can differently impact juvenile survival and adult reproduction. In both temperate and tropical regions, high rates of adult reproduction in a given environment may not be realized if occasional, high temperatures prevent survival to maturity. This suggests that considering the differing responses of multiple life stages is essential to understand the ecological and evolutionary consequences of climate change.     

Univoltine and bivoltine life cycles in insects: A model with density-dependent selection

Selection for univoltine and bivoltine life cycles in insects under resource-limited but favourable temperature conditions is analyzed with a difference equation model including density-dependent population dynamics based on the conceptual framework of an evolutionarily stable strategy. The model predicts that the bivoltine type can spread in a univoltine population when the fraction of density-independent rate of annual increase by producing a second generation exceeds the survival rate during diapause of the univoltine type, but monopoly of the bivoltine type is not possible unless it attains an equilibrium population density exceeding that of the univoltine type. The applicability of the model prediction in explaining the occurrence of a partial bivoltine cycle in predominantly univoltine population in the temperate zones is discussed.     

Synrhabdosome life cycles

Synrhabdosomes are monospecific colonies of colonies of graptolites. Their mode of formation is unknown. We draw attention to a type of colony formation found in two genera of colonial rotifers and suggest that these represent plausible models for the life cycle of graptolite synrhabdosomes.     

Pheromones in the life of insects

Life in insect societies asks for a permanent flow of information, often carried by rather simple organic molecules. Some originate from plants as odours of blossoms or exudates from trees. Especially important are the intra- and interspecific combinations of compounds produced by the insects themselves. These are called pheromones or ecto-hormones and serve a variety of tasks. The paper deals mainly with honeybee pheromones, but takes also into consideration those of wasps and hornets. Effects of pheromones are monitored ethologically by direct observation and filming as well as in a more quantitative manner with using direct and indirect calorimetry. In all experimental set-ups alarm pheromones were used as controls. They show an up to fourfold increase of activity after a few seconds, determined for small groups of insects as well as for a whole hornet nest placed in a 25-l calorimeter. A variety of cosmetics like soaps, shampoos, lotions and perfumes are included in the investigations because of repeated reports about unwarranted insect attacks which are said to be provoked by such products. None of the applied substances provoked a significant reaction of the bees (p > 0.05). A short appendix discusses the still questionable existence of pheromones in man, which were confirmed under laboratory conditions, but not yet for daily life.     

Timing in the life histories of insects

This paper presents a heuristic model illustrating some major problems in analyzing seasonal life histories of multigeneration insects. The concept of the critical interval is introduced and defined as the age classes that survive at the end of a period of population growth. These conclusions follow from the results: The optimal age for occupying a habitat depends upon the duration of the habitat as well as the life history of the insect. Two positions of the initial age distribution may give local maxima for fitness. The critical interval should often include the youngest age classes to maximize fitness while the optimal position of the initial age distribution may be at a much older age. In this case, conflicts arise between the positions of the critical interval at the end of one growing period and the initial age distribution at the start of the next. The length of the critical interval that maximizes fitness in a particular environment may be relatively small in which case mortality at the end of a growing period may be high and timing would appear to be poor even though fitness is maximized. In this model, optimum generation lengths exist which are not the shortest attainable. Finally, the length of time that a habitat remains suitable influences all of the above results and must be taken into account in analyzing the adaptedness of life history traits.     

Assessment of two mesh sizes for interpreting life cycles, standing crop, and percentage composition of stream insects

Using Surber-type samplers and dip-net samplers, we assessed the efficiency of nets having pore sizes of 720 μm and 320 μm for determining standing crop and percentage composition of the stream fauna, and for collecting representative size-class specimens of Ephemeroptera and Plecoptera to be used in life-cycle studies. Except for one species, samples collected with either the 720-μm or 320-μm dip-net led to the same general inferences about the species' life cycle. Of fifty possible sample comparisons, there were twelve samples where the size-class frequencies of particular species collected in the 720-μm dip-net were significantly different from the size-class frequencies of the 320-μm dip-net; for five of these samples a deficit of large nymphs (> 5.0 mm) in the 320-μm net mainly contributed to the significant χ² values. On one date, we used double-bag samplers with both th e 720-μm and 320-μm nets attached to either the Surber or dip-net sampler. Approximately 50% of the insects by numbers passed through the 720 fxm mesh ofeach sampler, but only 5% by volume-biomass. Shape of the insect as well as body length was important in assessing mesh-size efficiencies. The 720-μm mesh of the double-bag dip-net sampler retained most of the Nemoura dnctipes (having stout appendages) and Epeorus longimanus (flattened) nymphs 2.0 mm in body length and larger; whereas most Baetis (streamlined) nymphs smaller than 3.0 mm and all Paraleuctra (needle-like shape) nymphs passed through the 720-μm mesh.     

Long term cycles caused by patchy predation

Simple models were developed empirically to account for possible effects of patchy annual predation upon populations of benthic species. Long-term population cycles were generated and, provided there was limited interaction among three prey species, three different cycles were obtained. With exponential or power increments in populations, the prey showed spatial aggregations, but these were less marked than those observed in nature. The failure of the models to explain certain other natural phenomena is commented upon.     

Integrating function across marine life cycles

Complex life cycles involve a set of discrete stages that candiffer dramatically in form and function. Transitions betweendifferent stages vary in nature and magnitude; likewise, thedegree of autonomy among stages enabled by these transitionscan vary as well. Because the selective value of traits is likelyto shift over ontogeny, the degree of autonomy among stagesis important for understanding how processes at one life-historystage alter the conditions for performance and selection atothers. We pose 3 questions that help to define a research focuson processes that integrate function across life cycles. First,to what extent do particular transitions between life-historystages allow those stages to function as autonomous units? Weidentify the roles that stages play in the life history, typesof transitions between stages, and 3 forces (structural, genetic/epigenetic,and experiential) that can contribute to integration among stages.Second, what are the potential implications of integration acrosslife cycles for assumptions and predictions of life-historytheory? We provide 3 examples where theory has traditionallyfocused on processes acting within stages in isolation fromothers. Third, what are the long-term consequences of carryoverof experience from one life cycle stage to the next? We distinguish3 scenarios: persistence (effects of prior experience persistthrough subsequent stages), amplification (effects persist andare magnified at subsequent stages), and compensation (effectsare compensated for and diminish at subsequent stages). We usethese scenarios to differentiate between effects of a carryoverof state and carryover into subsequent processes. The symposiumintroduced by our discussion is meant to highlight how discretestages can be functionally coupled, such that life cycle evolutionbecomes a more highly integrated response to selection thancan be deduced from the study of individual stages.     

Mathematical description of holometabolous life cycles

The deterministic continuous equations are developed for the population levels of the various life stages of holometabolous species. Special attention is paid to the larval stage in which the variable of weight, in addition to the traditional variables of age and time, is included; growth rate and the genetic variability of the growth rate are allowed for. A pair of equations is derived that permits computation of the larval population level without regard to the levels of the other life stages. Finite difference equations are developed, simple analytical forms for vital rates are adopted, and a numerical example is given.     

Morphoprocess and life cycles of organisms

In developing the ideas of V.N. Beklemishev about an organism as a form, existing in a process of determined transformation and matter/energy exchange, we consider different aspects of the term "morphoprocess" and introduce corresponding additional terms. Momentary morphoprocess characterizes an organism in the given moment of time. This term reflects a constancy of the form ("momentary form"), where the existence of an organism can be imagined as a sequence of "momentary forms". "First derivative" of this momentary characteristic is particular morphoprocess--an organism from its origin to fission/division or death. Compound particular morphoprocess is a determined and reiterating sequence of different particular morphoprocesses. And, at last, general morphoprocess--a "second derivative" of momentary morphoprocess--is rhythmical reiteration of a particular morphoprocess on the long-term scale, an ancestors/descendants lineage. To describe consecutive changes in this material system, the terms ontogenesis and life cycle are used. Ontogenesis characterizes a sequence of the morpho-functional changes of an individual organism during its life, whereas life cycle reflects a sequence of changes during one complete segment of the general morphoprocess represented by a single or several particular morphoprocesses. We also discuss morphoprocess uniformity along with the phase nature of morphoprocesses, both particular and compound particular ones.     

Long distance migration of insects to a subantarctic island

Transoceanic migration of four species of macrolepidoptera to subantarctic Macquarie Island has been detected in 7 out of 33 years during the period 1962–96 and is restricted to spring and autumn. Analyses of synoptic charts during the migration period show that autumn immigrants originated from New Zealand and comprised a single species of noctuid moth,Agrotis ipsilon (Walker). Spring immigrants originated from Australia and comprised two noctuids, Dasypodia selenophora Guenée and Persectania ewingii Westwood and a butterfly, Vanessa kershawi (McCoy). Autumn migrations were associated with depressions in the southern Tasman Sea. Spring migrations were associated with the eastward passage of prefrontal airflows ahead of cold fronts which extended from southern Australia to the west of Macquarie Island. In an analysis of one of these events, winds exceeded 30 ms^?1 at 300 m altitude and could have transported migrants from Tasmania to Macquarie Island overnight in less than 10 h. Flight activity was assisted by the presence of a nocturnal temperature inversion that maintained upper air temperatures above 5 °C. The effect of potential global warming on the migration and colonization of Macquarie Island by insects is discussed.     

Bacterial symbionts in insects: balancing life and death

Arthropods, particularly insects, form successful long-term symbioses with endosymbiotic bacteria. The associations between insects and endosymbionts are remarkably stable; many stretch back several hundred million years in evolutionary time. With the exception, perhaps, of the filarial nematodes no other group of metazoans shows such a proclivility for their intracellular symbionts. The identification and classification of bacterial symbionts and hosts has grown rapidly over the last two decades and these relationships form a continuum from classical mutualism to parasitism. Complete genomes have been sequenced for many of these bacteria and some of their hosts. Now more intractable questions regarding endosymbiosis are being addressed. Investigations on the role of the host immune system in the maintenance of symbiosis, the nature of bacteriophages and transposable elements found in the genomes of many bacterial symbionts, and the molecular mechanisms involved in establishing reproductive phenotypes such as parthenogenesis, male killing, cytoplasmic incompatibility and feminization have been recently reported. This review will focus on the impact of the secondary endosymbionts Wolbachia, Cardinium, and Spiroplasma on host fitness and immunity and will revisit the question of whether these bacteria are friend or foe from an insect’s point of view.     

The effect of delayed host self-regulation on host-pathogen population cycles in forest insects

Delayed host self-regulation using a Beverton-Holt function and delayed logistic self-regulation are included in a host-pathogen model with free-living infective stages (Anderson and May's model G) with the purpose of investigating whether adding the relatively complex self-regulations decrease the likelihood of population cycles. The main results indicate that adding delayed self-regulation to the baseline model increases the likelihood of population cycles. The dynamics display some of the key features seen in the field, such as cycle peak density exceeding the carrying capacity and a locally stable equilibrium coexisting with a stable cycle (bistability). Numerical studies show that the model with more complex forms of self-regulation can generate cycles which match most aspects of the cycles observed in nature.     

The priming of periodical cicada life cycles

Periodical cicadas in the genus Magicicada have unusually long life cycles for insects, with periodicities of either 13 or 17 years. Biologists have explained the evolution of these prime number period lengths in terms of resource limitation, enemy avoidance, hybridization and climate change. Here, I question two aspects of these explanations: that the origin of the life cycles was associated with Pleistocene ice age events, and that they evolved from shorter life cycles through the lengthening of nymphal stages in annual increments. Instead, I suggest that these life cycles evolved earlier than the Pleistocene and involved an abrupt transition from a nine-year to a 13-year life cycle, driven, in part, by interspecific competition.     

Origin and evolution of animal life cycles

The ‘origin of larvae’ has been widely discussed over the years, almost invariably with the tacit understanding that larvae are secondary specializations of early stages in a holobenthic life cycle. Considerations of the origin and early radiation of the metazoan phyla have led to the conclusion that the ancestral animal (= metazoan) was a holopelagic organism, and that pelago-benthic life cycles evolved when adult stages of holopelagic ancestors became benthic, thereby changing their life style, including their feeding biology. The literature on the larval development and phylogeny of animal phyla is reviewed in an attempt to infer the ancestral life cycles of the major animal groups. The quite detailed understanding of larval evolution in some echinoderms indicates that ciliary filter-feeding was ancestral within the phylum, and that planktotrophy has been lost in many clades. Similarly, recent studies of the developmental biology of ascidians have demonstrated that a larval structure, such as the tail of the tadpole larva, can easily be lost, viz. through a change in only one gene. Conversely, the evolution of complex structures, such as the ciliary bands of trochophore larvae, must involve numerous genes and numerous adaptations. The following steps of early metazoan evolution have been inferred from the review. The holopelagic ancestor, blastaea, probably consisted mainly of choanocytes, which were the feeding organs of the organism. Sponges may have evolved when blastaea-like organisms settled and became reorganized with the choanocytes in collar chambers. The eumetazoan ancestor was probably the gastraea, as suggested previously by Haeckel. It was holopelagic and digestion of captured particles took place in the archenteron. Cnidarians and ctenophores are living representatives of this type of organization. The cnidarians have become pelago-benthic with the addition of a sessile, adult polyp stage; the pelagic gastraea-like planula larva is retained in almost all major groups, but only anthozoans have feeding larvae. Within the Bilateria, two major lines of evolution can be recognized: Protostomia and Deuterostomia. In protostomes, trochophores or similar types are found in most spiralian phyla; trochophore-like ciliary bands are found in some rotifers, whereas all other aschelminths lack ciliated larvae. It seems probable that the trochophore was the larval type of the ancestral, pelago-benthic spiralian and possible that it was ancestral in all protostomes. Most of the non-chordate deuterostome phyla have ciliary filter-feeding larvae of the dipleurula type, and this strongly indicates that the ancestral deuterostome had this type of larva.     

Seasonal temperature alone can synchronize life cycles

In this paper we discuss the effects of yearly temperature variation on the development and seasonal occurrence of poikiliothermic organisms with multiple life stages. The study of voltinism in the mountain pine beetle (Dendroctonus ponderosae Hopkins), an important forest insect living in extreme temperature environments and exhibiting no diapause, provides a motivational example. Using a minimal model for the rates of aging it is shown that seasonal temperature variation and minimal stage-specific differences in rates of aging are sufficient to create stable uni-and multi-voltine oviposition cycles. In fact, these cycles are attracting and therefore provide an exogenous mechanism for synchronizing whole populations of organisms. Structural stability arguments are used to extend the results to more general life systems.