Types of parasitism of acarines and insects on terrestrial vertebrates

According to the recent taxonomic revisions, over 40000 species of insects and acarines are parasites or micropredatory blood-suckers of mammals and birds. The largest fraction of them are micropredators and temporary or permanent ectoparasites, the minority being endoparasitic. Some arthropods (blood-sucking dipterans) use the host primarily as a food resource, whereas for others (many astigmatic mites) the host constitutes the entire environment. A number of life forms, or types of parasitism, have arisen in the insects and acarines in the course of their adaptive evolution to parasitism on terrestrial vertebrates. The term “type of parasitism” designates a set of convergently arising morpho-physiological and ecological adaptations (adaptive complexes), demonstrated by different arthropod taxa. A classification of the types of parasitism in arthropods is proposed based on their temporal, spatial, and trophic associations with vertebrates. The following seven types of parasitism are distinguished: micropredatory blood-suckers, nest ectoparasites (nidicoles), temporary ectoparasites with prolonged feeding, permanent ectoparasites, intracutaneous endoparasites, cavity endoparasites, and tissue endoparasites.     

The coevolution of ixodid ticks and terrestrial vertebrates

Paleontologic and zoogeographic data speak in favour of Mesozoic origin of ixodid ticks. The absence of strict restrictions for the feeding on unusual species of hosts has caused the domination of polyphagy and oligophagy over monophagy among ixodid ticks. The same peculiarities of ixodid ecology are responsible for a restricted part or absence of phylogenetic parallelism with hosts in their evolution. Primary food relations with reptiles are, apparently, preserved only in the genus Aponomma and in many species of Amblyomma while hosts for most species of other genera are mammals and, to a lesser extent, birds. The number of potential hosts in these species can be much greater than that of real ones. Restrictions in the distribution of some species are connected rather with direct effect of unfavourable environmental factors on their nonparasitic stages of the life cycle than with the absence of suitable hosts. During the evolution of natural landscapes and at a shorter stages under the influence of successions or anthropogenic factors ixodids easily adapt themselves to feeding on new species of hosts. So the differentiation of primary and secondary hosts of these parasites is rather difficult.     

寄生虫的起源与进化

介绍了寄生虫的提前适应机制在寄生虫起源与进化中的重要作用,同时还推测了寄生虫可能的进化过程,并且阐述了寄生虫对寄生生活的适应对策以及对宿主行为的改变。     

The origin of parasitism in trombiculid mites (Acariformes: Trombiculidae)]

On the basis of literary data and original investigations some phylogenetic, ecological and morphological aspects of the origin of parasitism in trombiculid mites are carefully considered for the first time. It is shown that parasitism in this group of trombidiform mites is a relatively young historical phenomenon and was formed after their ontogenesis had differentiated into active and quiescent stages. Therefore, in the life pattern of trombiculid mites the character of individual development, that defines their biotopical restriction, is much more important than the phase parasitism. Primitive organization of the digestive system and extraintestinal digestion, so characteristic of this group, are one of the main reasons of the origin of their parasitism. Under pasture conditions trombiculid mites, that initially were predators-entomophages with bite-sucking mouth parts, pass easily to parasitism on vertebrate animals and become primary lymphophages. They use the vertebrate host's organism exclusively as a source of food and by the extent of polyphagia are very close to free-living blood-sucking insects. Stylostome, that develops during feeding of trombiculid larvae and some other closely related groups of trombidiform mites, is a universal structure for achieving a large amount of food on a wide range of animals during a relatively short period of time and reflects wide host-parasite specificity of these parasitic mites. From the historical view the larvae of trombiculid mites did not pass from one group of hosts to the others, but owing to morphological preadaptation to parasitism passed in a definite historical period, not earlier than Paleogene, to parasitism on all classes of terrestrial vertebrates, especially on mammals, their primary hosts.     

Neurobiological mechanisms of the origin of terrestrial vertebrates

Some trends in the phylogenetic changes of the vertebrate neural system in the course of their transition from aquatic to terrestrial mode of life are analyzed. Wide multifunctionality of the neural centers and heterotopic substitution of the brain functions are supposed to be prerequisites of transition of archaic amphibians to the land via a stable transitional environment. The latter include underground or vegetation aquatic-air labyrinths in which conditions of gradual acquisition of adaptations to the terrestrial mode of life occurred. The principal neurological adaptations to such mode of life and less dependence on aquatic environment developed during the long-term evolution in that environment. The neurobiological model of evolutionn of the archaic amphibians in the transitional environment of aquatic-air labyrinths suggested herewith is compared to other hypotheses of the tetrapod origin.     

The evolution of the terrestrial vertebrates: environmental and physiological considerations

Physiological evidence has long been used to suggest that the gnathostomous vertebrates (those possessing jaws) were primitively fresh water. The same was also the case for the Osteichthyes (bony fish) and the Tetrapoda (Amphibia, Reptilia, Aves, Mammalia). However, the geological evidence favours a marine origin for the vertebrates as a whole, and, for the gnathostomes and the osteichthyans in particular. Some of the earliest amphibian remains may be associated with tidally influenced sediments. Furthermore, during the early part of the Devonian, fresh water chemistry may well have been different from that of today, lessening the divide between marine and non-marine environments. Urea formation via the ornithine cycle, and urea retention in the body fluids, are useful adaptations for terrestrial life. They prevent excessive water loss associated with the elimination of nitrogenous waste. These abilities may have been primitive for the gnathostomes, and were developed in the marine environment to reduce osmotic dehydration. In the aqueous medium, gaseous exchange is effected by the gills. These organs are, on the whole, useless in air. For vertebrates, air-breathing is effected by an inflatable sac, with moist linings, and an internal location. Some form of air-breathing sac was primitive for the osteichthyans, and may have been primitive for the gnathostomes. Again, this adaptation for terrestrial life developed in response to conditions experienced in the marine, aquatic environment. A new model of tetrapod evolution is proposed in the light of the basic marine origin and character of the ancestors of the tetrapods.     

Convergent evolution and locomotion through complex terrain by insects, vertebrates and robots

Arthropods are the most successful members of the animal kingdom largely because of their ability to move efficiently through a range of environments. Their agility has not been lost on engineers seeking to design agile legged robots. However, one cannot simply copy mechanical and neural control systems from insects into robotic designs. Rather one has to select the properties that are critical for specific behaviors that the engineer wants to capture in a particular robot. Convergent evolution provides an important clue to the properties of legged locomotion that are critical for success. Arthropods and vertebrates evolved legged locomotion independently. Nevertheless, many neural control properties and mechanical schemes are remarkably similar. Here we describe three aspects of legged locomotion that are found in both insects and vertebrates and that provide enhancements to legged robots. They are leg specialization, body flexion and the development of a complex head structure. Although these properties are commonly seen in legged animals, most robotic vehicles have similar legs throughout, rigid bodies and rudimentary sensors on what would be considered the head region. We describe these convergent properties in the context of robots that we developed to capture the agility of insects in moving through complex terrain.     

The urbilaterian brain: developmental insights into the evolutionary origin of the brain in insects and vertebrates

Classical phylogenetic, neuroanatomical and neuroembryological studies propose an independent evolutionary origin of the brains of insects and vertebrates. Contrasting with this, data from three sets of molecular and genetic analyses indicate that the developmental program of brains of insects and vertebrates might be highly conserved and suggest a monophyletic origin of the brain of protostomes and deuterostomes. First, recent results of molecular phylogeny imply that none of the currently living animals correspond to evolutionary intermediates between protostomes and deuterostomes, thus making it impossible to infer the morphological organization of an ancestral bilaterian brain from living specimens. Second, recent molecular genetic evidence provides support for the body axis inversion hypothesis, which implies that a dorsoventral inversion of the body axis occurred in protostomes versus deuterostomes, leading to the inverted location of neurogenic regions in these animal groups. Third, recent developmental genetic analyses are uncovering the existence of structurally and functionally homologous genes that have comparable and interchangeable functions in early brain development in insect and vertebrate model systems. Thus, development of the anteriormost part of the embryonic brain in both insects and vertebrates depends upon the otd/Otx and ems/Emx genes; development of the posterior part of the embryonic brain in both insects and vertebrates involves homologous control genes of the Hox cluster. These findings, which demonstrate the conserved expression and function of key patterning genes involved in embryonic brain development in insects and vertebrates support the hypothesis that the brains of protostomes and deuterostomes are of monophyletic, urbilaterian origin.     

The affinities of mites and ticks: a review

Mites and ticks can be divided into two well-defined clades, Anactinotrichida and Actinotrichida, for which a recent work formalized a suite of putative autapomorphies and reciprocal differences. Whether they are sister-taxa – forming a monophyletic Acari – is more controversial. Earlier supporters of two independent origins for mites largely failed to demonstrate convincing synapomorphies between either of the two lineages and other arachnid orders; although recent work on reproductive biology revealed explicit characters of this nature. Furthermore, some of the characters proposed in support of a monophyletic Acari do not stand up to detailed scrutiny when compared with Arachnida in general. Effective morphological comparisons between mites and other arachnids are hindered by incompatible nomenclature and long-standing, mite-specific characters which are difficult to score for other arachnids. Furthermore, taxon-specific characters restricted to individual mite groups have sometimes been treated erroneously as 'typical' for all Acari. Here, previous hypotheses of mite affinities are reviewed. Historically, authors have debated whether mites are basal arachnids or highly derived. Excluding weakly supported early hypotheses, mites have been resolved – in whole or in part – as sister-group of all other Arachnida (based on tagmosis), closely related to Opiliones (based mostly on genital morphology), Palpigradi (based on controversial interpretations of limb morphology), Solifugae (based mostly on the mouthparts, but now perhaps also reproductive characters) and Ricinulei (based on hexapodal larvae and perhaps mouthparts). We cannot provide a final resolution here, but we aim to highlight important character sets which should be included in subsequent phylogenetic analyses, as well as useful areas for future investigations: particularly tagmosis and the nature of the gnathosoma.     

Stability and manoeuvrability of terrestrial vertebrates

For a standing animal to be statically stable, a vertical linethrough its centre of mass must pass through the polygon ofsupport defined by its feet. Statically stable gaits are possiblefor quadrupeds but do not seem to be used. Physical and mathematicalmodels have shown that bipedal gaits can be dynamically stable.Accelerations and decelerations of animals may be limited bymuscle strength, by the coefficient of friction with the groundor by considerations of stability. Cornering ability similarlymay be limited by strength or by the coefficient of friction.It may be faster to use a longer route involving corners oflarger radius than a shorter one with sharper corners.     

Loss of larval parasitism in parasitengonine mites

Larval Parasitengona are typically parasites, yet at least 29 species of water mites and one species of Trombidiidae forgo larval feeding and any association with a host. Species with non-feeding larvae are isolated cases within species groups or genera where the remaining species have parasitic larvae. Species without larval parasitism occur in at least 14 genera, eight families and four superfamilies of water mites; the loss of larval parasitism is presumably polyphyletic, having occurred at least 21 times. Lineages of water mites with non-feeding larvae frequently exist in parallel with almost identical populations or species that have parasitic larvae. Thus, there is tremendous potential for studies comparing the relative merits of the two life history strategies. Comparisons indicate that adults from lineages with non-parasitic larvae produce smaller numbers of larger eggs; the extra nutrition included in larger eggs permits the larvae to forgo feeding. Non-feeding larvae frequently have wider dorsal plates but reduced leg length, setal length and sclerotization when compared to parasitic larvae from sister lineages. The adults of lineages with non-feeding larvae are frequently smaller in comparison to adults of sister lineages with parasitic larvae. There is no apparent pattern in relation to habitat: lineages lacking larval parasitism occur in streams, temporary ponds and the littoral and planktonic regions of permanent lakes. © Rapid Science Ltd. 1998     

The selection, testing and application of terrestrial insects as bioindicators

Although the uses and merits of terrestrial insects as indicators have been extensively discussed, there is a lack of clear definition, goal directedness and hypothesis testing in studies in the field. In an attempt to redress some of these issues and outline an approach for further studies, three categories of terrestrial insect indicators, corresponding to differences in their application, are proposed, i.e. environmental, ecological and biodiversity indicators. The procedures in terrestrial insect bioindicator studies should start with a clear definition of the study objectives and proposed use of the bioindicator, as well as with a consideration of the scale at which the study is to be carried out. Bioindication studies are conducted at a variety of spatial and temporal scales within the context of earth-system processes, but the objectives of the study will largely determine the scale at which it would be optimally conducted. There is a tendency for studies to be conducted below their space-time scaling functions, giving them apparent predictability. The selection of potential indicator taxa or groups is then based on a priori suitability criteria, the identification of predictive relationships between the indicator and environmental variables and, most importantly, the development and testing of hypotheses according to the correlative patterns found. Finally, recommendations for the use of the indicator in monitoring should be made. Although advocating rigorous, long-term protocols to identify indicators may presently be questionable in the face of the urgency with which conservation decisions have to be made, this approach is critical if bioindicators are to be used with any measurable degree of confidence.     

Olfactory receptors in aquatic and terrestrial vertebrates

In species representing different levels of vertebrate evolution, olfactory receptor genes have been identified by molecular cloning techniques. Comparing the deduced amino-acid sequences revealed that the olfactory receptor gene family of Rana esculenta resembles that of Xenopus laevis, indicating that amphibians in general may comprise two classes of olfactory receptors. Whereas teleost fish, including the goldfish Carassius auratus, possess only class I receptors, the `living fossil' Latimeria chalumnae is endowed with both receptor classes; interestingly, most of the class II genes turned out to be pseudogenes. Exploring receptor genes in aquatic mammals led to the discovery of a large array of only class II receptor genes in the dolphin Stenella Coeruleoalba; however, all of these genes were found to be non-functional pseudogenes. These results support the notion that class I receptors may be specialized for detecting water-soluble odorants and class II receptors for recognizing volatile odorants. Comparing the structural features of both receptor classes from various species revealed that they differ mainly in their extracellular loop 3, which may contribute to ligand specificity. Comparing the number and diversity of olfactory receptor genes in different species provides insight into the origin and the evolution of this unique gene family. Accepted: 29 July 1998     

Ultraviolet vision and foraging in terrestrial vertebrates

Tetrachromatic colour vision, based on four ‘main’ colours and their combinations, is probably the original colour vision in terrestrial vertebrates. In addition to human visible waveband of light (400–700 nm) and three main colours, it also includes the near ultraviolet part of light spectrum (320–400 nm). The ecological importance of ultraviolet (UV) vision in animals has mainly been studied in the context of intra‐ and inter‐sexual signalling, but recently the importance of UV vision in foraging has received more attention. Foraging animals may use either UV cues (reflectance or absorbance) of food items or UV cues of the environment. So far, all diurnal birds studied (at least 35 species), some rodents (4 species), many reptilians (11 species) and amphibians (2 species) are likely able to see near UV light. This probably allows e.g. diurnal raptors as well as frugivorous, nectarivorous and insectivorous birds to use foraging cues invisible to humans. The possible role of UV cues and existing light conditions should be taken into account when food selection of vertebrate animals is studied, particularly, in experiments with artificial food items.     

Developmental rate isomorphy in insects and mites

When the proportion of total developmental time spent in a particular developmental stage does not change with temperature, an organism shows "rate isomorphy." This is the case only if the lower developmental threshold is the same for all developmental stages. In this study, the incidence of rate isomorphy in seven species of mites and 342 species from 11 insect orders (some represented by several populations) was determined. Whether a species shows rate isomorphy or not was determined over a range of temperatures where the relationship between the rate of development and temperature is linear. Proportion of total developmental time spent in a particular stage was plotted against temperature and the existence of rate isomorphy inferred from a zero change in proportion. Rate isomorphy was detected in 243 (57%) of 426 populations. In the rest of the cases, rate isomorphy was violated by deviations in the proportion of time spent in a stage by an average of 0.2% (range 4.5E-06% to 2.8%) at the mean of the range of temperatures of all the data sets (11 degrees C). The violations occurred most frequently at the extremes of the linear phase, which is attributed to methodical biases, mortality at low temperatures, or too coarse an estimate of developmental time at high temperatures. Similarly, a meta-analysis also revealed an overall prevalence of rate isomorphy. Consequently, in insect and mite species, all the developmental stages appear to have the same population-specific lower developmental threshold. The existence of rate isomorphy could be of great practical importance, for example, in the timing of life-history events and in determining preadult thermal requirements. There are also indications that it may act as a phylogenetic constraint.     

文昌鱼—研究脊柱动物起源和进化的模式动物

长久以来,文昌鱼一直被认为和生活在约5亿年前的脊椎动物的直接祖先相似。由于文昌鱼在进化上的重要性,它在动物学研究史上发挥着关键作用,近100多年来,文昌鱼作为研究对象曾数次受到动物学界青睐或冷落,大约10年前,随着分子生物学技术应用于文昌鱼研究,又激发了动物学家对文昌鱼的研究兴趣,又一次出现在文昌鱼研究的高潮,并且一直持续至今,分子生物学研究结果表明,文昌鱼样生物可能是环节动物样动物和最早的脊椎动物之间的进化中间体,因此,文昌鱼在动物学研究史上好像绕了个大圈又回到了原处,在被忽视一段时间之后,又重新占据脊椎动物起源和进化研究中心舞台的位置,成为研究脊椎动物起源和进化的模式动物。     

Uricoteley:its nature and origin during the evolution of tetrapod vertebrates

The hepatic mechanism for detoxication of ammonia formed during amino acid gluconeogenesis in uricotelic vertebrates requires the intramitochondrial synthesis of glutamine by glutamine synthetase. This glutamine then serves as a precursor of uric acid in the cytosol. The evolutionary development of uricoteley thus required the localization of glutamine synthetase in liver mitochondria. The mechanism for the mitochondrial import of glutamine synthetase in uricotelic vertebrate liver is not yet known. Tortoises, extant relatives of the stem reptiles, possess both the ureotelic and uricotelic hepatic systems. It therefore seems likely that the genetic events allowing the mitochondrial localization of glutamine synthetase in liver occurred in the amniote amphibian ancestors of the stem reptiles. The selection of ureoteley by the theropsids and of uricoteley by the sauropsids were major events in the divergence and subsequent evolution of these two lines. Once established in the sauropsid line, uricoteley has persisted through to the higher reptiles, crocodilians, and birds. Uricoteley was in part responsible for the radiation of the archosaurs during the Triassic as a water-conserving mechanism in the adult, thereby allowing them to invade the arid environments of that period. Contrary to dogma, uricoteley was probably of minor significance in the development of the cleidoic egg. Neither mammalian nor avian embryonic liver tissues catabolize amino acids to any great extent, so it is inappropriate to attribute to them a kind of "waste" nitrogen metabolism.     

Controversial aspects of photoperiodism in insects and mites

This review examines several controversial aspects of photoperiodism in insects and mites including the role of the circadian system in night length measurement, the nature of apparent hourglass-like responses, and whether or not the circadian component in photoperiodism is the same as that in overt behavioural rhythms. These aspects of the phenomenon are discussed in terms of the entrainment of circadian oscillations by cycles of light and temperature. There is considerable variety of photoperiodic response within the insects (and other arthropods) to show, inter alia, circannual rhythms, internal and external coincidence night length timers, and in some species, non-circadian hourglass-like devices. Many apparent hourglass-like responses, however, could be circadian ‘clocks’ of the external coincidence type involving oscillations that dampen below threshold in extended periods of darkness. The review also concludes that there is little evidence in favour of the “Hourglass clock-oscillator counter” model proposed for the mite Tetranychus urticae by Vaz Nunes and Veerman (1982a). The responses of this species to complex light and temperature cycles may also be interpreted in terms of a damped oscillator version of external coincidence.