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.     

Phylogenetic analyses of mode of larval development

Phylogenies based on morphological or molecular characters have been used to provide an evolutionary context for analysis of larval evolution. Studies of gastropods, bivalves, tunicates, sea stars, sea urchins, and polychaetes have revealed massive parallel evolution of similar larval forms. Some of these studies were designed to test, and have rejected, the species selection hypothesis for evolutionary trends in the frequency of derived larvae or life history traits. However, the lack of well supported models of larval character evolution leave some doubt about the quality of inferences of larval evolution from phylogenies of living taxa. Better models based on maximum likelihood methods and known prior probabilities of larval character state changes will improve our understanding of the history of larval evolution.     

Deconstructing environmental predictability: seasonality,environmental colour and the biogeography of marine life histories

Environmental predictability is predicted to shape the evolution of life histories. Two key types of environmental predictability, seasonality and environmental colour, may influence life‐history evolution independently but formal considerations of both and how they relate to life history are exceedingly rare. Here, in a global biogeographical analysis of over 800 marine invertebrates, we explore the relationships between both forms of environmental predictability and three fundamental life‐history traits: location of larval development (aplanktonic vs. planktonic), larval developmental mode (feeding vs. non‐feeding) and offspring size. We found that both dispersal potential and offspring size related to environmental predictability, but the relationships depended on both the environmental factor as well as the type of predictability. Environments that were more seasonal in food availability had a higher prevalence of species with a planktonic larval stage. Future studies should consider both types of environmental predictability as each can strongly affect life‐history evolution.     

Tropical and temperate freshwater amphidromy: a comparison between life history characteristics of Sicydiinae,ayu, sculpins and galaxiids

Amphidromy is a distinctive form of diadromy, but differences in the life histories of tropical and temperate amphidromous fishes suggest that there are two types of freshwater amphidromy. The life histories of Sicydiinae gobies, ayu (Plecoglossus altivelis), Japanese sculpins (Cottus) and galaxiids (Galaxiidae), suggest that the Sicydiinae are representatives of tropical freshwater amphidromy, whereas ayu, sculpins and galaxiids are representatives of temperate freshwater amphidromy. The Sicydiine larval stage may be required to occur in the ocean for all species, but ayu, sculpins and galaxiids have landlocked or fluvial forms with larvae that do not need to enter the ocean for larval feeding and growth. This suggests that Sicydiine larvae have a high oceanic dependency whereas ayu, sculpins and galaxiid larvae have a low oceanic dependency. Freshwater amphidromous fish in tropical and temperate zones appear to have developed two different strategies in the evolution of their life histories. It is likely that the evolutionary direction of the larval stage of tropical amphidromy is to remain in the sea and that of temperate amphidromy is towards having the ability to remain in freshwater if needed. Tropical and temperate amphidromy appear to be biologically informative categories and evaluations of this hypothesis will facilitate better understanding of the various forms of amphidromy in the future.     

How larvae and life cycles evolve

Marine larvae have factored heavily in pursuits to understand the origin and evolution of animal life cycles. Recent comparisons of gene expression and chromatin state in different species of sea urchin and annelid show how evolutionary changes in embryonic gene regulation can lead to markedly different larval forms.     

Spirorbidae (Polychaeta) collected by R. V. "Vitjas" from abyssal depths of the Pacific Ocean

Pileolaria levensteinae sp. nov. was represented by juvenile and presumably adult stages, attached to ferromanganese substrata. None bore embryos, but the larger individuals had deep opercular cups and the species is closely related to certain shallow water forms which incubate in such cups. A second species was dextrally coiling but represented by a single fragment too small to describe. Spirorbids would appear to be preadapted for abyssal life, by having a brief, lecithotrophic larval stage.     

Mitochondrial DNA phylogeny and rates of larval evolution in Macrophiothrix brittlestars

Phylogenetic analysis has led to significant insights into the evolution of early life-history stages of marine invertebrates. Although echinoderms have been a major focus, developmental and phylogenetic information are relatively poor for ophiuroids, the most species-rich echinoderm class. We used DNA sequences from two mitochondrial genes to develop a phylogenetic hypothesis for 14 brittlestar species in the genus Macrophiothrix (Family Ophiotrichidae). Species are similar in adult form and ecology, but have diverse egg sizes and modes of larval development. In particular, two species have rare larval forms with characteristics that are intermediate between more common modes of feeding and non-feeding development. We use the phylogeny to address whether intermediate larval forms are rare because the evolution of a simplified morphology is rapid once food is no longer required for development. In support of this hypothesis, branch lengths for intermediate forms were short relative to those for species with highly derived non-feeding forms. The absolute rarity of such forms makes robust tests of the hypothesis difficult.     

Closing the larval loop: linking larval ecology to the population dynamics of marine benthic invertebrates

The majority of marine benthic invertebrates exhibit a complex life cycle that includes separate planktonic larval, and bottom-dwelling juvenile and adult phases. To understand and predict changes in the spatial and temporal distributions, abundances, population growth rate, and population structure of a species with such a complex life cycle, it is necessary to understand the relative importance of the physical, chemical and biological properties and processes that affect individuals within both the planktonic and benthic phases. To accomplish this goal, it is necessary to study both phases within a common, quantitative framework defined in terms of some common currency. This can be done efficiently through construction and evaluation of a population dynamics model that describes the complete life cycle.

Two forms that such a model might assume are reviewed: a stage-based, population matrix model, and a model that specifies discrete stages of the population, on the bottom and in the water column, in terms of simultaneous differential equations that may be solved in both space and time. Terms to be incorporated in each type of model can be formulated to describe the critical properties and processes that can affect populations within each stage of the life cycle. For both types of model it is shown how this might be accomplished using an idealized balanomorph barnacle as an example species. The critical properties and processes that affect the planktonic and benthic phases are reviewed. For larvae, these include benthic adult fecundity and fertilization success, growth and larval stage duration, mortality, larval behavior, dispersal by currents and turbulence, and larval settlement. It is possible to predict or estimate empirically all of the key terms that should be built into the larval and benthic components of the model. Thus, the challenge of formulating and evaluating a full life cycle model is achievable. Development and evaluation of such a model will be challenging because of the diverse processes which must be considered, and because of the disparities in the spatial and temporal scales appropriate to the benthic and planktonic larval phases. In evaluating model predictions it is critical that sampling schemes be matched to the spatial and temporal scales of model resolution.     

Utilization of larval and pupal detritus by Aedes aegypti and Aedes albopictus

The utilization of detritus sources by mosquito larvae during development may significantly affect adult life history traits and mosquito population growth. Many studies have shown invertebrate carcasses to be an important detritus source in larval habitats, but little is known regarding how invertebrate carcasses are utilized by mosquito larvae. We conducted two studies to investigate the rate of detritus consumption and its effect on larval development and life history traits. Overall, we found that Aedes aegypti and Aedes albopictus larvae rapidly consumed larval detritus, while pupal detritus was consumed at a significantly slower rate. We also found that the consumption of larval detritus significantly increased larval survivorship and decreased male development time but did not significantly influence female development time or pupal cephalothorax length for either sex. Our results suggest that the direct consumption of larval detritus can support the production of adults in larval habitats that lack allochthonous detritus inputs or where such organic inputs are insufficient. These studies indicate that different forms of invertebrate detritus are utilized in distinct ways by mosquito larvae, and therefore different forms of invertebrate detritus may have distinct effects on larval development and adult life history traits.     

Evolution and development of budding by stem cells: Ascidian coloniality as a case study

The evolution of budding in metazoans is not well understood on a mechanistic level, but is an important developmental process. We examine the evolution of coloniality in ascidians, contrasting the life histories of solitary and colonial forms with a focus on the cellular and developmental basis of the evolution of budding. Tunicates are an excellent group to study colonial transitions, as all solitary larvae develop with determinant and invariant cleavage patterns, but colonial species show robust developmental flexibility during larval development. We propose that acquiring new stem cell lineages in the larvae may be a preadaptation necessary for the evolution of budding. Brooding in colonial ascidians allows increased egg size, which in turn allows greater flexibility in the specification of cells and cell numbers in late embryonic and pre-metamorphic larval stages. We review hypotheses for changes in stem cell lineages in colonial species, describe what the current data suggest about the evolution of budding, and discuss where we believe further studies will be most fruitful.     

Who came first--larvae or adults? origins of bilaterian metazoan larvae

There is a classic controversy in zoology over whether the common ancestor of living bilaterian phyla was a benthic animal with a bilaterian body plan, or was a pelagic larva-like animal similar to what we see today in the primary larvae of indirect-developing bilaterians. We examine the current larva-like adult hypothesis, and present an alternate model for the evolution of complex life histories by intercalation of larval features into the ontogeny of an ancestral direct-developing bilaterian. This gradual accumulation of larval features results in a developmental regulatory program that produces a larva distinct in body plan from the adult. The evolution of a rapid and complete metamorphosis is made possible by the convergent evolution of set aside cells in the final stages of the emergence of indirect developing larval forms. Although convergences abound either hypothesis for the evolution of developmental pathways and life histories, the bilaterian first hypothesis is consistent with all stages of evolution of a complex life history being selectively advantageous, with the rapid evolution of larval forms, and with the frequent co-option of genes from the adult phase of the life cycle prevalent in the evolution of embryos and larvae.     

洞庭湖外睾吸虫新种及其生活史

本文报告洞庭湖区鲶鱼肠道寄生的洞庭湖外睾吸虫Exorchis dongtinghuensis sp.nov(新种)及其全程生活史,其第一中间宿主为湖北钉螺Oncomelania hupensis;第二中间宿主为鲤鱼、鲫鱼和金鱼;终宿主为鲶鱼Parasilurus asotus。作者对各期宿主作了人工感染试验和现场自然感染调查。对其发育过程作了观察比较。     

Relationship between species longevity and larval ecology in nassariid gastropods

The relationship between duration in the fossil record and larval ecology, based on the protoconch morphology, has been analysed for 40 species of Nassarius Duméril, 1806. The lifespan of species with planktotrophic larval development is significantly longer than for those with nonplanktotrophic development. Many species, representing both forms of larval ecology, cluster around certain values of longevity. This macroevolutionary tendency does not correspond to a phylogenetic pattern, nor does it depend on the ecology of the adult forms. The results are explained by means of a hypothesis on dispersion capacity of the larvae, taking into account the particular geological history of the Mediterranean and eastern Atlantic regions during the Neogene. Additional hypotheses, relating to, e.g., ecological tolerance or trophism, are rejected as being unnecessary in this case. □ Longevity, larval ecology, gastropods, Nassariidae, Neogene, Mediterranean, Atlantic.     

Testing adaptive hypotheses on the evolution of larval life history in acorn and stalked barnacles

Despite strong selective pressure to optimize larval life history in marine environments, there is a wide diversity with regard to developmental mode, size, and time larvae spend in the plankton. In the present study, we assessed if adaptive hypotheses explain the distribution of the larval life history of thoracican barnacles within a strict phylogenetic framework. We collected environmental and larval trait data for 170 species from the literature, and utilized a complete thoracican synthesis tree to account for phylogenetic nonindependence. In accordance with Thorson's rule, the fraction of species with planktonic‐feeding larvae declined with water depth and increased with water temperature, while the fraction of brooding species exhibited the reverse pattern. Species with planktonic‐nonfeeding larvae were overall rare, following no apparent trend. In agreement with the “size advantage” hypothesis proposed by Strathmann in 1977, egg and larval size were closely correlated. Settlement‐competent cypris larvae were larger in cold water, indicative of advantages for large juveniles when growth is slowed. Planktonic larval duration, on the other hand, was uncorrelated to environmental variables. We conclude that different selective pressures appear to shape the evolution of larval life history in barnacles.     

The nonfeeding auricularia of Holothuria mexicana (Echinodermata,Holothuroidea)

Among echinoderms, nonfeeding larvae usually are simplified in body shape, have uniform ciliation, and have lost the larval gut. A few species have nonfeeding larvae that express some remnant features of feeding larvae like ciliated bands and larval skeleton or larval arms, but typically their larval mouth never opens and their gut does not function. Still other species have retained the feeding larval form, a functional gut, and can feed, but they do not require food to metamorphose. The present note describes the development of a tropical holothurian, Holothuria mexicana, which hatches as a gastrula that is already generating coelomic structures. A translucent auricularia forms with a mouth that opens but becomes reduced soon thereafter. In form and ciliation this auricularia resembles a feeding larva, but it does not respond to food. A doliolaria forms on day 4 and the pentactula on day 6 post‐fertilization. Further study of this larva and that of its closely related congener, Holothuria floridana, is warranted.     

DEVELOPMENTAL CONSEQUENCES OF AN EVOLUTIONARY CHANGE IN EGG SIZE: AN EXPERIMENTAL TEST

Larvae of two species of sea urchins (Strongylocentrotus droebachiensis and S. purpuratus) differ in initial form and in the rate of development. To determine whether these differences are attributable to the large interspecific difference in egg size, we experimentally reduced egg size by isolating blastomeres from embryos. The rate of development of feeding larvae derived from isolated blastomeres was quantified using a novel morphometric method. If the differences early in the life histories of these two species are due strictly to differences in egg size, then experimental reduction of the size of S. droebachiensis eggs should yield an initial larval form and rate of development similar to that of S. purpuratus. Our experimental manipulations of egg size produced three clear results: 1) smaller eggs yielded larvae that were smaller and had simpler body forms, 2) smaller eggs resulted in slower development through the early feeding larval stages, and 3) effects of egg size were restricted to early larval stages. Larvae from experimentally reduced eggs of the larger species had rates of development similar to those of the smaller species. Thus, cytoplasmic volumes of the eggs, not genetic differences expressed during development, account for differences in larval form and the rate of form change. This is the first definitive demonstration of the causal relationship between egg size (parental investment per offspring) and life-history characteristics in marine benthic invertebrates. Because larval form influences feeding capability, the epigenetic effects of egg size on larval form are likely to have important functional consequences. Adaptive evolution of egg size may be constrained by the developmental relationships between egg size and larval form: evolutionary changes in egg size alone can result in concerted changes in larval form and function; likewise evolutionary changes in larval form and function can be achieved through changes in egg size. These findings may have broader implications for other taxa in which larval morphology and, consequently, performance may be influenced by changes in egg size.     

Foodplant effects on colour morphs of Eumorpha fasciata caterpillars (Lepidoptera: Sphingidae)

Caterpillars of the hawkmoth Eumorpha fasciata are highly polymorphic for colour, with green, pink, and pink-and-yellow forms in the second through fourth instars, and green and multicoloured forms in the fifth instar. Four years of field censuses on four foodplant species determined that all morphs were found on all plant species; morph frequencies were homogeneous on each plant species over time; and morph frequencies differed consistently among plant species. When larvae were reared from eggs on three of the hostplant species in the laboratory, differences in morph frequencies paralleled the census results. Thus foodplant quality is one factor affecting larval colour in E. fasciata. A literature survey reveals that foodplant effects on larval colouration may be widespread in the family Sphingidae, but most reports are anecdotal rather than experimental. The implications of this mechanism of colour determination are discussed.     

The ontogenetic development of refracting superposition eyes in crustaceans: Transformation of optical design

We have investigated, comparatively, the ontogenetic development of the compound eye in larvae of a mysid (Neomysis) and a euphausiid (Thysanoessa) species and found it to be close to identical in the two species. The larval eye is of apposition type with special adaptations for planktonic life. The elongated dioptric apparatus is devoid of screening pigment and instead has a proximal lens optically isolating the ommatidium. The pigmented retina is extremely compressed making the eye largely transparent and presumably suitable for a planktonic life. The presence of this specialized type of eye in the planktonic larvae of euphausiids was known before but it is intriguing to find exactly the same type in mysids, spending their entire larval life as embryos in the female marsupium. A possible explanation is offered if mysids earlier in evolution had planktonic larvae. Upon reduction of free-living larvae, the transparent type of eye may have been preserved because there is no selection pressure on the larva to change it. In late larval life, both species transform their eyes to a refracting superposition type typical for adult mysids and euphausiids. The process of transformation and the functional connection between transparent apposition and superposition is described.