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Habitat, food and population dynamics of the field vole Microtus agrestis (L.) in south Sweden
(1971)
The habitat, food and population dynamics of the field vole (Microtus agrestis) were studied mainly in southernmost Sweden (Scania) with a few comparisons from a northern locality. The habitats of Microtus agrestis consist mainly of successional stages, arising from abandoned cultivated fields, drained mires and felled mature forests. They were characterized botanically and classified according to moisture and time since abandonment. Capture distributions were significantly clumped, being most contagious at the lowest densities. Microdistribution was affected more by shelter than by food conditions. The weight of the stomach contents showed various skewed distributions, giving indications of the feeding behaviour of various population categories. The staple food in southernmost Sweden was various grasses, while herbs, gnss seeds, vegetative storage organs and moss were primary food items during summer, summer-autumn, winter and early summer respectively. In a northern locality herbs played a more important role in summer. Among plant species e.g. Agrostis spp., Festuca rubra, Achillea millefolium and Ranunculus spp. were often eaten, while the common Deschampsia caesjJiiosa and Charnaenerion angustifoliurn were disliked. There were clear differences between habitats but plant species belonging to the same group replaced each other in the diet. There were few differences between populatipn categories. Experiments disclosed the same preferences as found at stomach analysis. Digestibility was higher during spring than winter, that of the preferred species being highest. The digestibility was most affected by the proportions of crude protein but the changes in weight by the amounts consumed. The animals showed a decided interest in certain carbohydrates and fatty substances with differences between seasons and years. The interest in sugars was not correlated with the similar interest in bark. The probability of capture was especially low in juveniles and during winter. Trap lines and grids showed the same trappability, but, the intervals between checkings influenced the removal rate. The density changes on two distant, large, dry fields in southernmost Sweden were correlated, while a nearby isolated population in a spruce plantation fluctuated out of phase. The increases during the reproductive season were significantly lower than those expected at exponential increase. The density in various habitats did not depend on the moisture conditions. Differences in population structure and body weight in various areas were partly related to the phase of population development. However, there were also differences in the same area between habitats, which might differ in plant nutrient supply. The animals had a low energy reserve in their depot fat, which would only sustain the animals for 5-14 hours in an emergency. It was highest during midwinter but with significant depressions in April and December. Drier habitats could sustain populations of little more than 200 animals/ha over :winter and these densities were not realized in southernmost Sweden. The spring-summer density variations in a northern locality were correlated with the spruce seed supply during the following autumn-winter. A relation was assumed with the quantity or quality of previous storage tissues. In southernmost Sweden, with irregular and low-amplitude fluctuations, there was no clear relation to primary production.
Arthropods inhabiting the sporophores of Fomes fomentarius (Polyporaceae) in Gatineau Park, Quebec
(1971)
The fauna of the sporophores of the perennial bracket fungus Fomes fomentarius (L. ex Fr.) Kickx were examined in a 3-year study. One species of molluscs and more than 152 species of arthropods excluding mites, representing 13 orders, 70 families, and 5400 individuals, emerged from or were found on or in, 1448 sporophores detached from dead birch trees; the sporophores were collected each year in Gatineau Park, Que., kept individually in screen-topped glass jars in a laboratory, and examined for several months. Mites, which were recorded quantitatively only in the final year, added 4 orders, 19 families, and 30 species to the preceding totals. Mites were the most frequently occurring and probably the most numerous arthropods, followed by Coleoptera, Psocoptera, Collembola, Hymenoptera, and Diptera. The key organisms of the fauna were five species of beetles that were primarily responsible for tunnelling and destruction of the sporophores and would therefore exert great influence on the composition of the community. Their tunnels provided shelter and food for many smaller arthropods or facilitated their feeding. Some beetle species tunnelled the sporophores for one season and others, for several, but many living sporophores and most dead ones tunnelled by beetles were tunnelled in the same season by more than one species of beetles. There was considerable latitude in types of sporophores inhabited by various arthropods but some species were particularly attracted to living or dead, to younger or older, and to smaller or larger, sporophores. Some species were also attracted to certain regions of the sporophore more than were other species. Possible economic implications of observations made in the study are discussed.
Application of vibrational spectroscopy to the problem of structure determination of molecules of biological interest goes back to the early uses of raman and infrared spectroscopy in the study of organic molecules. For reviews of earlier work the reader is referred to compilations by Kohlrausch (1943) and by Jones and Sandorfy (1956), whereas more recently a comprehensive discussion has been presented by Bellamy (1975). These compilations accentuate the correlation of vibrational spectra with molecular structure from an essentially empirical point of view and culminate in the establishment of empirical correlation charts. For typical examples the reader is referred to Weast (1974) and Bellamy (1975). There have been many treatments of the theoretical basis of molecular vibrational spectroscopy. Among them the classical work by Herzberg (1945) and by Wilson et al. (1955) should be mentioned. Applications of infrared spectroscopy (IR) to structure problems of biological interest have been summarized by Susi (1969), Fraser and MacRae (1973), and Wallach and Winzler (1974). It was remarked quite eraly that relevant structural information about biological systems often requires study in aqueous solution, which forms the natural environment for most biologically important systems. Besides critical control of experimental conditions and samples the conventional methods of raman spectroscopy may be applied to aqueous solutions in a quite straightforward manner, cf. the contribution by Lord and Mendelson, Chapter 8. The condition of biological environment, i.e., the study in aqueous solutions, by IR spectroscopy is difficult to achieve by conventional absorption technique, since the high absorption coefficient of water in wide regions of the mid and far infrared implies use of thin layers and high concentrations. As a consequence the application of special techniques for measurement of IR spectra of biological material has been a necessity in many cases. This contribution covers the following topics: (1) specific spectroscopic techniques used in this field, in particular for membrane spectroscopy, (2) discussion of typical results derived from application of IR techniques to model and natural membrane systems and to important constituent molecules of such systems.
Hatching asynchrony and the onset of incubation in birds revisited : when is the critical period?
(1995)
1. Birds are unique among animals in being able to influence the birthing intervals of their young through the timing of the onset incubation. However, many species hatch their young asynchronously, frequently resulting in reduced survivorship for later-hatched young. This is the Paradox of Hatching Asynchrony. 2. The Brood Reduction Hypothesis provided a resolution to the paradox by suggesting an adaptive function to the offspring mortality that results from asynchrony. Experimental tests have provided little support, and 16 alternative hypotheses have been proposed, but few have been tested. Most experimental tests have not measured important parameters such as parental effort and postfledging survival. Many have lacked adequate controls or sufficient statistical power. 3. We divide the hypotheses for hatching asynchrony into four categories based on the effects of intrinsic or extrinsic factors during a critical period of the nesting cycle which constrains reproductive success. Hatching asynchrony could be simply the consequence of the early onset of incubation during egg-laying, either as a result of physiological constraints on incubation or because parents derive fitness benefits from the protective function of early incubation. During the nestling period, hatching asynchrony could be adaptive if it allowed parents to eliminate one or more nestlings selectively, or increased parental efficiency. Alternatively, parents could manipulate the duration of the different periods of the nesting cycle to maximize benefits. 4. Because the onset of incubation generally determines hatching patterns, we encourage refocusing attention from the search for adaptive hatching patterns during the nestling period to the events surrounding the onset of incubation during egg-laying. Many factors can affect when incubation is begun, including physiology, and interactions with the environment, predators, competitors, and mates. 5. Patterns of the onset of incubation are difficult to determine and to quantify, in part because many birds begin incubating gradually, or at night. In some species, the onset of incubation varies with clutch size, but not in others. 6. The onset of incubation is the principle proximate control of hatching patterns, but other factors, such as egg size, embryonic vocalizations, and time of year may also affect hatching patterns. 7. Synchronous hatching is the primitive condition in birds, and is widespread in the lower, primarily precocial taxa. Most altricial species hatch their eggs asynchronously, although some exhibit synchrony as a secondarily derived trait. Hatching patterns show wide variation within some orders and families. 8. Patterns of the onset of incubation and hatching in a species may reflect the influence of multiplefactors. The relative importance of those factors may depend on the trade-offs associated with the potential benefits of early incubation to the survival of eggs and the potential costs to the survivor of later-hatching young associatedwith nestling size hierarchies. 9. The relative effects of multiple factors can be examined by integrating the results of empirical tests of single factors through modeling. 10. We demonstrated the use of a stochastic model by using empirical data from the House Sparrow. Results revealed the trade-offs inherent in the onset of incubation from differences in egg viability and nestling survivorship. An intermediate onset of incubation produced the greatest fledging success. 11. Other factors may be integrated into such models if they can be measured in terms of their effects on fledging success. Different factors, represented by different hypotheses, vary in how readily they may be modeled.
Fly larvae of the Anthomyiidae, Muscidae, Calliphoridae, and Sarcophagidae are scavengers and breed in filthy material such as animal dung, human feces, dead animals and decaying vegetables. Therefore, the presence of these larvae in our environment indicates defective sanitary conditions, and offers many problems from the standpoint of public health. Clinically, the larvae of some species cause myiasis in man and animals. On the other hand, in the field of legal medicine postmortem time of dead bodies is estimated in some cases by the species and age of larvae collected from them. Thus, the fly larvae are closely associated with our lives. Morphologic studies on larval stages of flies were started in the beginning of this century by Portchinsky (1910), Banks (1912), MacGregor (1914), and Keilin (1915 and 1917). Portchinsky first described chiefly the habits and metamorphosis, and later MacGregor emphasized structure of posterior spiracles in the identification of the larvae. Keilin (1917) discovered and emphasized the importance of the characteristic structures such as the cephalopharyngeal sclerite, anterior and posterior spiracles and other external and internal characters. Following them, many authors made comparative and systematic studies of fly larvae and there are many reports of such research. However, morphology, taxonomy and ecology of these fly larvae have not been studied systematically in Japan. During the period from 1915 to 1936, 10 or more accounts were made about fly larvae that caused intestinal myiasis in man. In 1937, Kodama and Yasuda reported Ophyra nigra Wied. asa cause of intestinal myiasis. Yasuda (1939) reported 12 species of fly larvae in Seoul, Korea with detailed drawings and descriptions. Lopes (1943 and 1946) reported sarcophagid larvae from Neotropical region. Hall (1947) described calliphorid larvae from North America in his publication "The blow flies of North America". In 1951, Roback used the characters of the larvae such as pharyngeal sclerites and posterior spiracles for the classification of the Calypterate, Diptera. In the same year, Zimin reported Russian muscoicllarvae in his publication "Fauna USSR, Muscidae" using the external characters such as spines, papillae, and anal plate on the body surface as well as anterior and posterior spiracles and cepahlopharyngeal sclerite. Fan (1957) reported some filth fly larvae from China. In Japan, Kana et a!. (1950-1958) described larvae of 39 species belonging to 4 families. In the present paper, the author describes 3rd stage larvae belonging to 70 species in 33 genera of 4 families. In addition, photographs of internal and external structures and keys to families and species of 3rd stage larvae are given. Moreover, 6 species belonging to Muscidae and 1 species ofSarcophagidae for which the larvae were not seen are cited from the descriptions and figures published by Keilin (1917), Zimin (1951), Kano and Sato (1951), and Fan (1957). Those species are also included in the keys of this paper.
Fauna of Isopoda and Tanaidacea in the coastal zones of the Antarctic and Subantarctic waters
(1968)
This work presents results of the processing of the material collected by the Soviet Antarctic Expedition (1955 - 1963), chiefly on the "Ob" and to a lesser extent by the geographer E. S. Korotkevich and the ichthyologist V. M. Makushek., during their wintering at the Mirnyi station, as well as by the hydrobiologist G. A. Solyanik on board the whalers "Slava" and "Ivan Nosenko."
Notes on some primates, Carnivora and the babirusa from the Indo-Malayan and Indo-Australian regions
(1949)
The Great Spotted Woodpecker is the most common and best-known woodpecker species in the W Palearctic. The sections Habitat, Distribution, Population, Movements, Food, Social pattern and behaviour, Voice, Breeding, Plumages, Bare parts, Moults, Measurements, Weights, and Geographical variation have been updated or completely rewritten, and a new section has been added on Conservation. High flexibility enables the Great Spotted Woodpecker to utilize a great variety of habitats, from Arctic taiga through boreal and temperate to Mediterranean (N Africa and Canary Islands) and Alpine forest zones, wherever there are mature trees of any sort with sufficient growth to accommodate nest-holes and with a supply of available food. The Distribution and Population sections include new data of population tendencies, with stable or positive trends in most European countries and fluctuating populations in N Europe. Recent negative range trends with small decreases only in S Europe, apparently due to loss of wooded habitats. The Conservation section presents literature on the Great Spotted Woodpecker as the key or umbrella species for secondary cavity-nesting species. This section also demonstrates the importance of woodpeckers as indicators for naturally dynamic forests with tree species diversity, forest management, and sustainable forestry. Woodpeckers can be part of a monitoring system of e.g. sustainable forestry, but species from other organism groups are also required. The section on Movements has been updated with new data on dispersal and summer/autumn and spring migration. The Great Spotted Woodpecker uses a very wide and varied diet and is characterized as a 'universalist' in food-gathering; pecking and hammering are most important in autumn and winter, gleaning and probing are important at times of high food availability at the tree surface in spring and summer. It takes arthropods and insect larvae, coniferous seeds and various nuts mainly in autumn and winter, and drills holes for sap-sucking in spring, takes surface dwelling arthropods and caterpillars, bird eggs and nestlings, and fruits and berries in spring and summer. The Social pattern and behaviour section presents new data on the mating system, parental effort, pair-bond, divorce rate, survival rate, and mortality. Great Spotted Woodpeckers are socially and genetically monogamous with a potential to polyandry, which was recorded in Japan. Changes of partner between seasons common. Males usually invest more in nesthole construction and guarding than the females and contribute the same amount or more to brood care. Males usually incubate and brood at night, as with all studied woodpeckers, and defend territories, which seem to be important for female choice. Females compete intensely for access to males and perform male-like courtship behaviours such as drumming. Great Spotted Woodpeckers are intelligent and currently doing damage to house facades. The Voice section presents mainly new data on calls and instrumental signals of the young. The Breeding section has been updated with new information about nest-sites, breeding behaviour, and breeding success. New data on age determination are shown in the Bare parts and Moults sections. Additional data are provided on size and weight.
The Japanese micropterigid moths are revised. Seventeen species in five genera are recognized from Japan, described or redescribed with the male and female genital figures. Of these, two genera, Issikiomartyria HASHIMOTO and Kurolkopteryx HASHIMOTO, and seven species, Issikiomartyria akemiae HASHIMOTO, Issikiomartylia plicata HASHIMOTO, Issihiomartyria distincta HASHIMOTO, Issihiomartyria bisegmentata HASHIMOTO, Kurokopteryx dolichocerata HASHIMOTO, Neomicropteryx hiwana HASHIMOTO, and Neomicropteryx redacta HASHIMOTO, are new to science. A new combination is given: Issikiomartyria nudata (Issuu). Biology and immature structures of the Japanese species are also described together with the keys to genera and to species provided on the basis of the adult characters. Phylogenetic relationships among the Northern Hemisphere genera are analyzed by the cladistic analysis using PAUP* (SWOFFORD, 2002) based on the morphological characters of adults. A monophyly of the Northern Hemisphere genera except for Micropterix is supported by nine apomorphies, but their immediate sister taxon remains unresolved.
Revision of the millipede family Andrognathidae in the Nearctic region (Diplopoda, Platydesmida)
(1975)
Homology of virtually all major components of facial anatomy is assessed in Archosauria in order to address the function of the antorbital cavity, an enigmatic structure that is diagnostic for the group. Proposed functions center on its being a housing for a gland, a muscle, or a paranasal air sinus. Homology is approached in the context of the Extant Phylogenetic Bracket method of reconstructing unpreserved aspects of extinct organisms. Facial anatomy and its ontogeny was studied in extant archosaurs (birds and crocodilians) to determine the osteological correlates of each soft-tissue component; resemblances between birds and crocodilians comprised the similarity test of homology. The congruence test of homology involved surveying phyiogenetically relevant fossil archosaurs for these bony signatures. The facial anatomy of extant birds and crocodilians is examined in detail to provide background and to discover those apomorphic aspects that contribute to the divergent specialization of these two groups and thus obscure homologies. Birds apomorphically show enlarged eyeballs, expanded nasal vestibules, and reduced maxillae, whereas crocodilian faces are dorsoventrally flattened (due to nasal rotation) and elongated. Most facial attributes of archosaurs are demonstrably homologous and in fact characterize much more inclusive groups. Special emphasis has been placed on the nasal conchae and paranasal air sinuses. Within Amniota, the following conchal structures are homologous, and all others are neomorphs: avian caudal concha, crocodilian concha + preconcha, Sphenodon caudal concha, squamate concha, and probably the mammalian crista semicircularis. The avian antorhital paranasal air sinus is homologous with the crocodilian caviconchal sinus; the maxillary sinus of placental mammals is not homologous with the archosaurian paranasal sinus. With regard to the function of the antorbital cavity, archosaurs possess homologous nasal glands, dorsal pterygoideus muscles, and paranasal air sinuses, but the osteological correlates of only the paranasal sinus involve the antorbital fenestrae and fossae. Thus, the antorbital cavity is best interpreted as principally a pneumatic structure.
An annotated checklist and bibliography of 197 species (representing 78 genera and 26 families) of non-marine polychaetes of the world is presented, including synonymies, information on ecology, distribution, habitat, and references to the taxonomic and biological literature. Over half (57%) of the checklist species are represented by just three families as follows: Nereididae (61 species including Namanereis, Namalycastis, Neanthes and Hediste), Aeolosomatidae (27 species, mostly Aeolosoma) and Sabellidae (24 species including Caobangia and Manayunkia). Other well-represented taxa are the epizoic histriobdel1id Stratiodrilus (11 species), the inland-sea-specialist ampharetid Hypania and related genera (5 species), and the freshwater-tolerant spionid Marenzelleria (5 species). One new combination is proposed for the nereidid Nereis tenuipalpa Pflugfelder, 1933, viz. Paraleonnates tenuipalpa n. comb.
Birds are characterized by pneumatization of their skeletons by epithelial diverticula from larger, air-filled cavities. The diverticula-or 'air sacs'-that invade the postcranium result from outgrowths of the lungs; poslcranial pneumaticity has been very well studied. Much more poorly understood are the air sacs that pneumatize the skull. Study or craniofacial pneumaticity in modern birds (Neornithes) indicates the presence of two separate systems: nasal pneumaticity and tympanic pneumaticity, The lacrimal and maxillary bones arc pneumatized by diverticula of the main paranasal cavity, the antorbital sinus. There are five tympanic diverticula in neornithines that pneumatize the quadrate, articulare and the bones of the braincase. The pneumatic features of the following six genera of Mesozoic birds are examined: Archaeopteryx, Ellaliornis, Baplomis, Parahesperornis, Hesperornis and lchthyornis. Despite the 'archaic' aspect of most of these birds, many of the pneumatic features of neornithines are found in .Mesozoic birds and are considered primitive for Aves. The phylogenetic levels at which most of the avian pneumatic features arose within Archosauria are uncertain. Until the phylogenetic levels at which homologous pneumatic features arose are determined, it is unwise to use most pneumatic characters in the discussion of avian origins. Within avian phylogeny, Ornithurae and Neornithes are well-supported by pneumatic synapomorphies. There is a trend towards reduction of craniofacial pneumaticity within Hesperornithiformes. Witthin Neornithes, four derived pneumatic characters suggest that the Palaeognathae (ratites and tinamous) is monophyletic.
After giving a brief historical account of the use and precise definitions of the various measurements and their indices in termites, the need for bringing together all such known measurements and indices in the fonn a monograph is explained. Precisely defined measurements obviate the necessity of using vague expressions for the comparison of allied taxa. Of the 88 measurements and 53 indices thus listed, 66 and 34 respectively have been used already in the published literature, and 22 and 19 new ones are added here.
Carnian (Upper Triassic) fishes from Polzberg bei Lunz have been known since 1886 but no comprehensive account has been published. Eleven species are described nine of which, Saurichthys calcaratus, Polzbergia brochatus, Peltoplellrus dinlmptus, Habroichthys gregarius, Nannolepis elegans, Phaidrosoma lunzensis, Elpistoichthys pectinatus, E. striolatus and Pholidophoretes salvus are new, and two others, Thoracopterus niederristi Bronn and Gigantopterus telleri Abel, previously little-known. New supraspecific taxa defined are: the order Polzbergiiformes, the family Thoracopteridae and the genera Polzbergia, Nannolepis, Phaidrosoma, Elpistoichthys and Pholidophoretes. Habroichthys. Thoracopterus, Gigantopterus and Nannolepis show an unusual skull-roof pattern and are included in the re-defined order Luganoiiformes. Two new ichthyokentemids considerably extend the known time-range of this family. The genus Pholidophoretes is intermediate between the Archaeomenidae Goodrich 1909, and the Pholidophoridae sensu stricto Nybelin 1966. The Polzberg assemblage was probably mainly marine with a small freshwater contribution; it shows less similarity to the Besano and Raibl assemblages than these do to each other. The Luganoiiformes are probably, but not certainly, monophyletic; relationships within the order are analyzed and a cladogram constructed. The Platysiagiformes, Peltopleuriformes, Luganoiiformes and Cephaloxeniformes could all have been derived from a common ancestor at the Perleidus level and are probably offshoots of the perleidid radiation.
The Opisthobranchia comprise highly specialized marine gastropods and have therefore been subject to diverse investigations covering various biological disciplines. However, a robust phylogeny of these gastropods is still lacking and several subclades have only been rarely studied. Furthermore, crucial aspects for the evolution of Opisthobranchia have not been comparatively analysed. Therefore, the aim of the present thesis is to gain new insights into the phylogeny of the Opisthobranchia with special focus on certain critical groups (Pleurobranchomorpha, Acteonoidea) and to assess several crucial features of the evolution of the investigated clades. The combination of four different gene markers (18S rDNA, 28S rDNA, 16S rDNA and CO1) and modern molecular systematic analysis tools were used to construct phylogenetic hypotheses focussing on Opisthobranchia as a whole as well as Pleurobranchomorpha and Acteonoidea in more detail. Intriguing new aspects of phylogeny and evolution of Opisthobranchia were revealed. First of all, monophyly of Opisthobranchia is definitely rejected based on the present data, while monophyly of Euthyneura (comprising Opisthobranchia and Pulmonata) is supported. Monophyly of opisthobranch subclades is confirmed for Nudipleura (as well as its constituting groups Nudibranchia and Pleurobranchomorpha), Umbraculida, Pteropoda (as well as subclades Thecosomata and Gymnosomata) and Acochlidiacea, for Cephalaspidea (if Runcinacea is regarded as a separate clade) and for Sacoglossa (if Cylindrobulla is accepted as an Oxynoacea). Aplysiomorpha are rendered paraphyletic due to the position of Akera bullata, but this result needs further investigation and should be considered with caution. The Nudipleura are found as the first single offshoot of the Euthyneura implying an early evolutionary separation of the last common ancestor of this clade. The remaining taxa form two main clades, one comprising the opisthobranch subgroups Umbraculida, Cephalaspidea, Aplysiomorpha and Pteropoda, while the other contains the pulmonate taxa and the opisthobranch Sacoglossa and Acochlidiacea. The interrelationships within these clades remain largely unresolved due to low statistical support values. However, a possible sister group relationship of Acochlidiacea and Eupulmonata receives statistical support. Opisthobranchia display various highly specific adaptations to diverse food sources. However, evolution of these specialized traits has never been assessed at an analytical level. The current thesis reconstructs the evolution of dietary preferences with novel methodologies based on the newly proposed phylogenetic hypothesis. Reconstruction of dietary evolution revealed herbivory as the ancestral condition in Euthyneura implying that carnivory evolved at least five times independently in the diverse lineages. The first comprehensive molecular phylogenetic hypothesis of the Pleurobranchomorpha could not reveal monophyly of the two main subclades Pleurobranchaeidae and Pleurobranchidae. This is due to the position of a single taxon (Euselenops luniceps) which is assigned to the Pleurobranchaeidae based on morphology but clusters within Pleurobranchidae in the current hypothesis. Furthermore, the tribe Berthellini and the genus Berthella are rendered paraphyletic by the current analyses. The results of molecular systematic analyses were used to reconstruct historical biogeography of Pleurobranchomorpha. Four different methodological approaches were applied yielding ambiguous results for Pleurobranchomorpha. However, the Pleurobranchidae comprising about 80% of the extant Pleurobranchomorpha most probably derived from an Antarctic origin. Dating of the phylogenetic tree via molecular clock methods yielded divergence of Pleurobranchidae into the Antarctic Tomthompsonia antarctica and the remaining species in Early Oligocene. Afterwards the latter underwent rapid radiation during Oligocene and Early Miocene. This divergence event coincides with two major geological events in the Antarctic region. On the one hand, the onset of glaciation and on the other hand the opening of the Drake Passage with concurrent formation of an Antarctic circumpolar current (ACC). I suppose that these sudden and dramatic changes in climate and palaeogeography probably accounted for migration of the last common ancestor of Pleurobranchidae (besides Tomthompsonia) into warmer regions via the Drake Passage to the Western Atlantic and Eastern Pacific and via the South Tasman Rise to the Indo-West Pacific. Furthermore, the ACC may have triggered larval dispersal to the Eastern Atlantic. The phylogenetic position of Acteonoidea has been a matter of debate for decades and they have long been considered as basal opisthobranchs. Results of the present thesis rather support placement in “Lower Heterobranchia” as sister group of Rissoelloidea. The current division of Acteonoidea into three families has never been investigated by means of phylogenetic methods. Thus, this thesis provides the first comprehensive investigation of this clade challenging present division into three families. The results rather support division into two main clades with the monogeneric Bullinidae clustering within Aplustridae doubting its separate status. Additionally, Rictaxis punctocaelatus which has been assigned to Acteonidae clusters basal to Aplustridae rendering Acteonidae paraphyletic. Since information on morphology of R. punctocaelatus was lacking until now, I conducted the first detailed investigation on morphology and histology of this species in order to reassess the unexpected molecular systematic placement. Character tracing analyses revealed similarities with both acteonoidean families implying an intermediate position of this species which might be assigned to a separate family in the future. Furthermore, the common features of Acteonidae and Rictaxis (massive shell, small foot, anterior mantle cavity opening, and absence of oral gland) are possibly plesiomorphic for the whole Acteonoidea. In summary, the results of the present thesis provide valuable novel insights into the phylogeny and evolution of the Opisthobranchia by employing state-of-the-art approaches of molecular systematics and evolutionary reconstruction. Thus, diverse hypotheses on opisthobranch phylogeny and evolution were either supported or rejected as well as novel hypotheses proposed which offer the basis for further research on these extraordinary gastropods.
Recently, the first part of the morphological revision of the Southeast Asian water monitor lizards of the Varanus salvator (Laurenti, 1768) species group provided a taxonomic overview over the members of this successful and widespread species complex (Koch et al. 2007). There, the Philippine taxa marmoratus, nuchalis and cumingi were reelevated to species status due to diagnostic morphological characteristics, e.g. significantly enlarged scales on the neck region. In this second part of the ongoing revision, these three species are re-investigated using additional voucher specimens and advanced statistical techniques including canonical variates analysis and principal component analysis. Our new investigations indicate that V. marmoratus represents a composite species, comprising at least three distinct taxa. Hence, the populations of the Sulu Archipelago (Tawi-Tawi Island) and those of the Palawan region are described as new species, viz. Varanus rasmusseni sp. nov. and V. palawanensis sp. nov., respectively. The allopatric island populations of V. cumingi inhabiting Samar, Leyte, and Bohol (the East Visayan subregion) show characteristic and geographically correlated colour patterns distinct from the type locality Mindanao (the second subregion of Greater Mindanao), warranting subspecific partition of this species. The new subspecies is named V. cumingi samarensis ssp. nov. In contrast, the taxonomic status of V. nuchalis remained unchanged, although this species shows some considerable variation in colour pattern. The systematic chapters are supplemented with notes about biology and conservation status. The hitherto underestimated diversity and zoogeography of Philippine water monitors is discussed in the light of Pleistocene sea level fluctuations. Finally, we introduce a scenario for the evolution and spread of Southeast Asian water monitor lizards and provide an identification key for the Philippine members of the V. salvatoI' complex.
1. Fomes applanatus, a perennial polypore of wide distribution and of common occurrence in North America and in Europe is responsible for the decay of very large quantities of wood annually. It attacks practically all deciduous species and several coniferous species, both dead wood and living trees. This fungus, heretofore commonly regarded as a pure saprophyte, has now for the first time been comprehensively studied; the investigations recorded in this paper have followed along three main lines of inquiry; (1) a study of the morphology and the ecology of F. applanatus and of the action on its host; (2) a determination of its etiological relationships by culture methods, and a testing of the applicability of such methods to a study of the timber destroying fungi; (3) an investigation of the possibility of finding criteria by which we may distinguish parasitic action on wood from saprophytic. 2. Fomes applanatus produces basidiospores only; conidia are not produced by the mycelium, nor, as is commonly affirmed, on the upper surface of the sporophore. The basidiospores are not of the ordinary type; each consists of a yellow, papillate, thick-walled chlamydospore within a thin hyaline wall. The so-called" truncate" base is in reality the apical end of the spore. Spore discharge is enormous and continues for by far the longest period recorded for fungi. It is continuous day and night for about six months-visible from vigorous fruiting bodies as spore clouds. Discharge is not affected by variations in light, humidity of the air, or temperature within very wide limits; frost causes an instant cessation and thereafter there is no further spore fall until a new set of pores is organized. The spores were not found to retain their viability for more than 6 1/2 months. They germinate in water and various other media within 48 hours after sowing, but the percentage of viable spores is very low and their behaviour with respect to germination is erratic. 3. No difficulty was experienced in culturing F. applanatus on artificial media or on wood. Three parallel series of cultures on wood were carried through to sporulation with mycelium obtained from (a) spores, (b) pieces of sporophore, (c) pieces of diseased wood. 4. Wood rotted by F. applanatus exhibits a mottled appearance sufficiently characteristic to permit of its recognition. when compared with other rots. In the case of living wood the area of attack is delimited by a wide dark band. Histologically, the decay is characterized by a perforation and corrosion of the elements culminating in disappearance of the tissues in localized pockets into which the mycelium gathers forming strands. Chemically, the change is one of delignification followed by a progressive solution of the resultant cellulose wall except for a few resistant fragments. In the later uniform white stage of decay, the threads of F. applanatus were always found to be accompanied by other destructive agentsbacteria or fungi. 5. The particular type of decay in timber due to Fomes applanatus has now been linked with its causal agent by actual culture. 6. Fomes applanatus has been proved to be a wound parasite, and in southern Ontario at least is one of the commonest and most destructive of this type. The proof rests on three grounds: (a) the conventional test applied to other such fungi-the mycelium works upward most readily by the way of the heartwood causing a characteristic decay and outward into the sapwood, eventual1y reaching the cambium, and is apparently the cause of the death of the tissues traversed by it; (b) a broad brown band is present in the wood of living trees along the advance line of the invading mycelium of this fungus. Within this band there is a copious production of brown wound gum and an excessive multiplication of tyloses; This band steadily moves fonvard with the advancing hyphae, the tyloses and wound gum being destroyed by the mycelium along its posterior margin as rapidly as they are formed along its antedor edge. The tyloses (and possibly the wound gum alsoJ certify to the living condition of the invaded tissues; their production can be ascribed only to the influence of the fungus, and the invasion of these tissues and their fate demonstrates directly its ability to act as a parasite; (c) inoculations with the spores and mycelium of F. applanatus into living trees resulted in an extensive browning of the inoculated wood with a multiplication of tyloses-both far in excess of similar phenomena due to traumatic stimulation. 7. The occurrence of wound gum and the multiplication of tyloses in a band marking the advance of a wood~destroying fungus in a living tree would appear to furnish an unerring criterion for the recognition of the fungus as a parasite.
Over 6260 fungi have been isolated from samples of feathers, nests, pellets, droppings, cloaca contents and visceral organs of 92 species of free-living birds in Czechoslovakia and Yugoslavia. Several species have been obtained of fungi pathogenic to homeotherms, poikilotherms and plants, and many fungi recorded belong to the toxinogenic species.Cellulolytic fungi have participated with 44.3 % in the total number of fungi, and keratinolytic fungi with 18.3 %. The thesis has been confirmed that habits (specific bionomics) of the birds influence,. to a certain degree, the distribution of the fungi among them. According to the results of this study, the fungi may conveniently be divided into three categories with respect to the character of their distribution and dispersal by free-living birds: (1) euryornithochous fungi , the occurrence of which in birds is undetermined by the birds' habits (ubiqitous species: Aspergillus flavus, Gliocladium roseum, Rhizopus. nigricans, Trichothecium roseum); (2) mesoornitohochorous fungi, the occurrence of which in birds is determined to a certain degree by the birds' habits (e. g., Aphanoascus fulvescens, Arthrodermcr curreyi, Chrysosporium keratinophilum, C. tropicum, Mucor hiemalis); (3) stenoornithochoruss fungi, which are associated with certain habit groups of the birds only (e. g., Aphanoascus terreus, Arthroderma ciferrii, A. tuberculatum,.Aspergillus fumigatus, Chrysosporium pannorum, Ctenomyces serratus). The importance of free-living birds in the spreading of pathogenic organisms has also been discussed from a more general point of view, particularly with regard to several epidemiologically important aspects of bird ecology such as synanthropism, migration (making possible a long-range carriage of pathogens or infected vectors), colonial breeding or mass roosting.
In 1080 samples (feathers, nesls, pellets, droppings, cloaca contents and visceral organs) collected from 92 species of free-living birds in many localities and habitats, a total of 6266 fungi was found; we identified 232 species and 112 genera of fungi (73.4 % Ascomycetes, 12.4 % Fungi imperfecti, 8.0 % Zygomycetes, 5.7 % Endomycetes, 0.3 % Oomycetes; and 0.2 % Basidiomycetes). The most common fungal genera were, in plumage, Alternaria. Cladosporium, Arthroderma, Chaetominum, Aspergillus, Penicillium and Chrllsosporium; in nests, Alternaria, Arthroderma, Penicillium, Aphanoascus and Scopulariopsis; on pellets, Penicillium, Mucor, Aspergillus, Chrysosporium and Rhizopus; in droppings, Scopulariopsis, Mucor, Aspergillus, Penicillium and Candida; from the cloaca contents, we isolated Penicillium, Candida, Aspergillus, Scopulariopsis, a. o., and from the visceral organs, Aspergillus, Penicillium, Rhodotorula, and some others. Of the fungi with a potential pathogenicity for homeotherms, we obtained, e. g., Absidia corymbifera, Aspergillus fumigatus, Candida albicans, Cryptococcus neoforrnans and Microsporum gypseum. We confirmed also the presence of several species of toxinogenic fungi (e. g., Aspergillus flavus), of the fungi with a potential pathogenicity for poikilotherms (e. g., Beauveria bassiana. Paecilomyces jarinosus, P. fumoso-roseus), and of plant parasites (Cylindrocarpon radicicola, Fusarium oxysponom and Verticillium albo-atrum). Aphanoascus reticulisporum has been described as comb. nov. (basionym: Anixiopsis: reticulispom ROUTIEN). 66 species of the fungi have not been recorded earlier in Czechoslovakia. Similarly, the occurrence of 111 species had, apparently, not been confirmed in free-living birds on a world-wide scale.
Compounds containing a carboxylic acid group may be metabolized along several routes. However, the most common pathways involve conjugation reactions with amino acids or glucuronic acid. Also, chain shortening due to beta-oxidation occurs when the carboxylic acid group is attached to a suitable aliphatic moiety. The various metabolic reactions of compounds containing the carboxylic acid group were reviewed by Caldwell. I Additional reactions of the carboxy group have been discovered which reveal that numerous I ipophilic conjugates may be formed. This subject was reviewed by CaldweW3 and Quistad and Hutson4 and the findings demonstrate that the acids may undergo chain extension or incorporation into triglycerides, cholesterol esters, and fatty acid derivatives. In addition to these reactions which occur in the tissues, the metabolism (e.g., decarboxylation or reduction) of some carboxylic acids may be carried out by the intestinal microflora.
Colonies of up to 30 cats, Felis catus, which were partially dependent upon man for direct provisioning with food, were recorded on 82% of 775 English farms. They lived at a mean density of 6.3 per km2. Members of one such colony were observed intermittently from 1978-81. These cats' ranging, foraging and scent marking behaviours are described briefly as a background to observations of their social interactions. The adult male's home range was 83 ha, whereas those of three adult females averaged 13.1 (SD 7.2) ha. The cats visited the observation barn independently of each other. When in the barn each cat differed in how it positioned itself with respect to the others. Furthermore, the cats' social relationships were structured by differences in the numbers, rates and types of interactions with one another. Some individuals were classed as net initiators of interactions, whereas others were net recipients. The tendency to rub the perioral and cheek regions of the face on another cat was the clearest single indicator of initiator status. Kittens were initiators to adults, females to the adult male, and some adult females were initiators to others. On average, each adult female in the colony rubbed on another once every 25.3 h. Behaviour within the colony was generally amicable, whereas towards outsiders it was aggressive. All adult females in the colony gave birth to kittens each year, and used communal nests. Females tended, groomed and nursed kittens other than their own, and cooperated with each other during parturition. Although a female might nurse certain kittens preferentially, these preferences were not necessarily for her own kitten. The frequency with which a female nursed a kitten and the frequency with which it rubbed on her were positively correlated. A case of infanticide, when an unrelated adult male killed kittens, is described, together with circumstantial reports indicating that this incident was not unique. Farm cat society appears to be structured centripetally, with interactions flowing predominantly from socially (and, sometimes, spatially) peripheral individuals to socially central ones.
The 70 Frullania species (+ 1 subspecies, 9 varieties, and 3 formae) belonging to 9 subgenera are confirmed in China, among which 1 subgenus and 4 species are new to science, and 12 species are newly reported from China. A new name (nom. nov.) and 12 combinations are proposed and several species are reduced to synonyms of other species. This study reveals the Frullania flora of China, particularly Yunnan and the neighboring provinces, to be most highly differentiated, containing many phytogeographic ally interesting taxa, and may be regarded as a center of the Paleotropic Frullania flora.
As a preliminary step towards a more intensive research on the diversity of macromycetes in Greece, an updated check-list of the Greek mycoflora is presented together with information on the host-substrates and geographic occurrence. The data originated from a thorough literature search and the authors' field observations. In total, 58 families, 214 genera and 811 species of fungi are recorded belonging to Basidiomycetes. The systematics and nomenclature of the relative bibliography have been updated and suitably revised. The large gaps in our knowledge on the existence and distribution of higher fungi in Greece are emphasized.
Echinococcosis/hydatidosis, caused by Echinococcus granulosus, is a chronic and debilitating zoonotic larval cestode infection in humans, which is principally transmitted between dogs and domestic livestock, particularly sheep. Human hydatid disease occurs in almost all pastoral communities and rangeland areas of the underdeveloped and developed world. Control programmes against hydatidosis have been implemented in several endemic countries, states, provinces, districts or regions to reduce or eliminate cystic echinococcosis (CE) as a public health problem. This review assesses the impact of 13 of the hydatid control programmes implemented, since the first was introduced in Iceland in 1863. Five island-based control programmes (Iceland, New Zealand, Tasmania, Falklands and Cyprus) resulted, over various intervention periods (from < 15 to > 50 years), in successful control of transmission as evidenced by major reduction in incidence rates of human CE, and prevalence levels in sheep and dogs. By 2002, two countries, Iceland and New Zealand, and one island-state, Tasmania, had already declared that hydatid disease had been eliminated from their territories. Other hydatid programmes implemented in South America (Argentina, Chile, Uruguay), in Europe (mid-Wales, Sardinia) and in East Africa (northwest Kenya), showed varying degrees of success, but some were considered as having failed. Reasons for the eventual success of certain hydatid control programmes and the problems encountered in others are analysed and discussed, and recommendations for likely optimal approaches considered. The application of new control tools, including use of a hydatid vaccine, are also considered.
A brief account of the present state of weevil taxonomy is followed by a detailed study of certain structures used in their classification, namely the venter, abdominal tergites, sternite 8 of the male, apex of the hind tibia and deciduous mandibular processes. A key to some 50 families and subfamilies of Curculionoidea is followed by a list of family-group taxa. The following changes are made: Brachyceridae, Erirhinidae. Cryptolnryngidae und Raymondionymidae are promoted to family rank from Curculiollidne; Antliarhininae is demoted to a subfamily of Brentidae, and Allocoryninae to a subfamily of Oxycorynidne; Coptonotini is demoted to a tribe of Curculionidue Scolytinae; Carinae, sufam. n. is erected for Car Blackburn (genus incertae sedis) in Belidae; Dinomor'phini is demoted to a tribe of Molytinae and Brachyccropsidinae is revived from synonymy with Dinomorphinae (Curclliionidae); Urachyderini, Eremnini, Otiorhynchini and Sitonini are demoted to tribes of Entiminue; Desmidophorinae is transferred from Brentidae to Brachyccridae, Ocladiini is promoted to a tribe of Desmidophorinae (from Curculionidae-Cryptorhynchinae); Campyloseelini (including Phaenomerina) is transferred from Rhynchophoridae to Curculionidae-Zygopinae; Carphodicticinae is promoted to subfamily rank and transferred from Curculionidae-Scolytinae to Platypodidae; Perieges; Schönherr is transferred from Curculionidae-Thecesterninae to Cryptoiaryngidae and Agriochaeta Pascoe from Cryptorhynchinae to Hyperinae (Curculionidae); Schadlarius Wood and Mecopelmus Blackman are transferred from Coptonotidae to Platypodidae.
The primary subdivisions of the brain (telencephalon, diencephalon, mesencephalon, metencephalon, and myelencephalon) have similar relations and comparable functions in all vertebrates. Accordingly, the landmarksthat define their boundaries can be regarded as reliable for following their development. On the basis of a more complete series of well preserved embryos than has been available hitherto, we present evidence that the subdivisions of the adult brain can be traced back to neural-fold stages in which a series of growth centers can be recognized, differing from one another in form, size, and relations. The possibility of following the constrictions between the various subdivisions throughout development has been doubted by some, notably Hochstetter (1919). At present we are convinced that they can be distinguished if certain criteria are followed. These are: (a) constrictions involve the neural tube as a whole; (b) constrictions do not give rise primarily to any neural centers; (c) constrictions change in relative length and width, and in certain stages they become inconspicuous in models. The anatomical descriptions of progressive stages of development have important practical implications. It is known, for example, that congenital malformations of the central nervous system in man are common and that they are responsible for a substantial portion of fetal wastage as well as infant mortality and morbidity. In certain patients comprehensive clinical studies may indicate the underlying abnormality, such as dysraphism, arhinencephaly, hypoplasia of the cerebellum. or absence of the corpus callosum. In addition, anatomical examination of the affected brains may reveal in detail such abnormalities as lyssencephaly, polymicrogyria, or other cortical dysgeneses. These very complex cerebral malformations can only be understood and unraveled in the light of normal development. An investigation of early development of the brain must necessarily begin with a stage in which the major landmarks of the adult brain can be readily identified.As progressively younger stages are analyzed certain landmarks can no longer be recognized, although others persist at least to the third week of gestation. We believe that the evidence on which this study is based can be followed more satisfactorily in this inverted sequence, and the detailed description is so presented. It is followed by a summary of the sequence of events written in the conventional manner, as far as the eighth week of gestation.
Delthyridoid spiriferids are characterized by a global abundance and fast evolution during Silurian and Devonian, and, therefore, are used as important biostratigraphical and palaeobiogeographical tools. In this work, delthyridoid brachiopod faunas from different regions of today’s world, resp., of different palaeobiogeographical units, are compared side-by-side to investigate their phylogenetic relationships and to improve, in a second step, the palaeobiogeography from Late Silurian to Early Eifelian time. A new systematics of Delthyridoidae is established which is more complicated than hitherto assumed. The results of this study are mainly based on direct comparison of articulated and isolated brachiopod shells, external and internal moulds, as well as latex casts and serial sections. The computer supported cladistic analyses have turned out not to be useful due to different kinds of preservation resulting in an incomplete matrix which is insufficient for reliable cladograms. A further problem in terms of cladistical analyses are various convergences during the evolution of spiriferids. Many characters evolved independently from each other at different times in each lineage so that autapomorphies are hardly or not at all recognizable. As a result, families and genera are only definable by a combination of characters rather than by a single or a few autapomorphies. As a new method, 3D reconstruction from serial sections is introduced which made it possible for the first time to compare directly mouldic and shelly material. Preliminary results are presented herein. Statistical analyses of measurements taken from new taxa are made but regarded as a descriptive argument rather than a deciding factor for taxonmy due to incomplete preservation and/or tectonic deformation. Brachiopods, especially type material, from collections of different institutions and museums are studied as well as personal material, whenever possible collected from topotype outcrops. Emended diagnoses, if necessary, from family to species level are given. During this work several new taxa have been erected: 7 new families: Australospiriferidae, Murchisonispiriferidae, Orientospiriferidae, Otospiriferidae, Patriaspiriferidae, Rostrospiriferidae, and Trigonospiriferidae; 6 new genera, 1 of these in open nomenclature: Cyclopterospirifer, Hallispirifer, Parlinispirifer, Murchisonispirifer, Shujiapingensispirifer, and gen. nov. B; and 3 new species: Patriaspirifer merriami, Patriaspirifer johnsoni, and Murchisonispirifer feldmani; 1 taxon is defined as nomen novum: Orientospirifer nakaolingensis wani. In the framework of this project, 2 families: Filispiriferidae and Multispiriferidae; 1 subfamily: Multiplicatispiriferinae, 6 genera, 1 of them in open nomenclature: Frequentispirifer, Leonispirifer, Multiplicatispirifer, Ovetensispirifer, Turcispirifer, and Gen. A; and 9 new species, 3 of them in open nomenclature: Filispirifer hamadae, Leonispirifer leonensis, Multiplicatispirifer foumzguidensis, Oventensispirifer novascotianus, Quiringites arensentiae, Turcispirifer turciae, Multiplicatispirifer cf. foumzguidensis, Quiringites cf. arensentiae, and ?Turcispirifer sp. A which have already been established are also described in this work. The brachiopod faunas studied consist of externally very similar spiriferids which have been identified as same genera, species, or even subspecies in earlier times. These forms are considered as 6 distinct morphotypes Howellella-, Arduspirifer-, Acrospirifer-, Euryspirifer-, Paraspirifer-, and Multiplicatispirifer-like morphotypes, which are briefly introduced. The new systematics is characterized by different clades, the European/North African delthyridoid spiriferid clade, the North American delthyridoid spiriferid clade, the Asian delthyridoid spiriferid clade, the Malvinokaffric delthyridoid spiriferid clade, and the delthyridoid multiplicated spiriferid clade. Each of them is described in a cladistic and in a phylogenetic way. Their phylogenetic relationship sheds new light on palaeobiogeographical interpretations for the different stages from Late Silurian to early Middle Devonian time. A tendency for increasing endemicity is seen until the end of the Early Emsian, which is interrupted by short term regional faunal exchange within a province or within a realm, followed by a loss of endemicity resulting in global distribution of brachiopod genera until the end of Givetian time. The Old World Realm is re-defined due to the lack of phylogenetic relationship between its faunas and subdivided into the European Realm, consisting of the Gondwanan and Avalonian provinces, and the Asian Realm, consisting of the Siberian, Sino, and Mongolian provinces. A reconstruction of Lower Devonian palaeobiographical map is introduced.
TeaABC from the halophilic bacterium Halomonas elongata belongs to the family of tripartite ATP-independent periplasmic (TRAP) transporters. It facilitates the uptake of the compatible solutes ectoine and hydroxyectoine which protect the cell from dehydration by accumulating in the cytoplasm during hyperosmotic stress. It is the only known TRAP transporter activated by osmotic stress. Ectoine and hydroxyectoine accumulation in H. elongata is regulated by the cytoplasmic universal stress protein TeaD. The gene encoding TeaD is located in the same operon as the TeaABC gene. TeaD regulates the cellular homeostasis of ectoine possibly by interacting directly or indirectly with TeaABC. All subunits of TeaABC and TeaD were expressed in E. coli and purified. With TeaD and the solute binding protein (SBP) TeaA high levels of expression suitable for crystallization could be obtained and their 3D structures solved. The small transmembrane protein TeaB and the transporter TeaC showed only moderate and low levels of expression respectively. Functional analysis on TeaA was performed using Isothermal Titration Calorimetry. The measurements demonstrate that TeaA is a high affinity ectoine-binding protein (Kd = 0.19 _M) that also has a significant affinity for hydroxyectoine (Kd = 3.8 _M). The structure of TeaA was solved using ab initio phase determination by MAD (multiple anomalous dispersion). TeaA structures were determined in three conformations: TeaA alone, TeaA in complex with ectoine and TeaA in complex with hydroxyectoine. The resolutions of the structures were 2.2, 1.55 and 1.80 Å, respectively. These represent the first structures of an osmolyte SBP associated to a TRAP transporter. The structures reveal similar ligand binding compared to osmolyte SBPs of ABC transporter pointing to coevolution of the ligand binding modes. Moreover, unique features such as the solvent-mediated specific binding of the ligands ectoine and hydroxyectoine could be observed for TeaA. The structure of TeaD in complex with its cofactor ATP was solved by molecular replacement at a resolution of 1.9 Å. Comparison with other structures of universal stress proteins shows striking oligomerization and ATP binding in TeaD. In conclusion, this work presents the first detailed analysis of the molecular mechanisms underlying ligand recognition of an osmoregulated transporter from the TRAP-transporter family.
Mitochondria are dynamic organelles indispensible for viability of eukaryotic cells. Diffusion of proteins in mitochondrial membranes is a prerequisite for the correct functionality of the organelles. However, its study is made complicated due to the nontrivial geometry, small size and positional instability of the organelle, restricting the usability of regular experimental methods and theoretical understanding of acquired data. Therefore, here the molecular transport along the main mitochondrial axis was investigated using highly accurate computational methods combining them with traditional experimental approaches. Using recently reported electron microscopic tomography data concerning the constitution of mitochondria [Fre02], a lattice model of the inner mitochondrial membrane (IM) reproducing its structure in great details was built up. With Monte Carlo (MC) simulations of particle dynamics on this model, it was found that the membrane geometry induces nonlinear effects in the motion of molecules along the mitochondrial axis, which in turn lead to a transient violation of the 2nd Fick?s equation. We show that mere curvature of the IM resulting from the presence of cristae is sufficient for the emergence of transient anomalous diffusion (TAD) in the membrane. The MC calculations have enabled an accurate estimation of regularities in the extent of deviations from the normal regime, therefore allowing us to propose non-homogenous power law as a suitable generalization of the current approach to the analysis of experimental data for the transient dynamics. The general cause of TAD resulting from the membrane curvature alone, without any involvement of specific inter-particle interactions prompted us to predict the similar dynamical effect also for other curved cellular membranes, be it diffusion in endoplasmic reticulum or in plasma membrane of cells possessing dense microvilli. The data indicate that the geometry-induced anomalous diffusion should be easily detectable with current experimental methods, but only in the restricted range of time scales corresponding to high temporal resolution. Until now, experimental measurements of molecular diffusion in biological membranes indiscriminately assumed either pure normal or pure anomalous diffusion schemes for the analysis of data acquired in very wide range of temporal resolutions, which often lead to ambiguities in the interpretation of diffusion parameters. The MC calculations have clearly illustrated the necessity for a more subtle treatment of experimental conditions: the assumption of pure Gaussian diffusion model is justified only if the applied temporal resolution is sufficiently low (as is often the case when using scanning techniques exemplified further); otherwise, the transient regime should be tested for by means of the non-homogenous power function. In the second part of the study the Fluorescence Recovery after Photobleaching (FRAP) with the laser scanning microscope is introduced as a method of choice for studying protein mobility within mitochondrial membranes. The conventional FRAP methodology [Axe76] was extended to enable its application for the determination of confined diffusion with conventional laser scanning microscopes which allowed us to communicate for the first time the direct measurement of protein diffusion in mitochondrial membranes of living cells. This is achieved through adaptation of FRAP data analysis to account for the spatial dimensions of the organelle and the spatiotemporal pattern of light pulses induced by the microscope. The experimental circumstances existing during the particular measurement session are computationally recreated and this way the best suited values of diffusion parameters are found. The method is validated experimentally for four FP-tagged mitochondrial membrane proteins: the IM OxPhos complexes F1F0 ATPase and cytochrome c oxidase and for Tom7 and hFis1 - components of the mitochondrial protein import and fission machineries respectively localized in the outer membrane. We find that for all proteins simple normal diffusion is not a sufficient description. In the inner membrane, diffusion coefficient of F1F0 ATPase expressed in HeLa cell line is found to be 0.2 ?m2/s, with more than 1/3 of the protein molecules being immobilized, while cytochrome c oxidase (in CEF primary cells) demonstrated a similar diffusivity pattern (0.4 ?m2/s, 30% immobile). In the outer membrane, the D (0.7 ?m2/s) and immobile fraction (7-8%) of GFP-Tom7 and GFP-hFis1 (both in HeLa cells) are identical, which designates a substantial difference in comparison to the IM protein mobility. Diffusion coefficients of mitochondrial membrane proteins studied here lay in the intermediate region between those measured in artificial bilayers and in plasma membranes. Protein crowding and intermolecular interactions will be among the major causes responsible for the detected slowdown of diffusion.
A recently completed field survey of springs throughout the Great Basin yielded collections of hydrobiid snails from more than 500 sites, and revealed a wealth of undescribed diversity of these small gastropods. In this, the first or a two-part taxonomic series treating this material, 58 new species of Pyrgulopsis Call & Pilsbry, 1886, are described; and new records are provided for 10 previously described members of this genus. Assignment of these novelties to Pyrgulopsis is done with the acknowledgement that this large genus, as currently constituted, is probably not monophyletic, but a more refined classiffcation of these snails reflecting evolutionary relationships must await preparation of a phylogenetic analysis, which is beyond the scope of this work. Pyrgulopsis occur in a variety of spring-fed water bodies in the Great Basin, including brackish and/or thermal habitats. Although a few species are widespread in the region, local endemism is prevalent lind 22 of the new species are known only from single localities. Several areas contain concentrations of locally endemic snails which may represent species flocks, notably Duckwater Valley (seven species) and southern Steptoe Valley (five species). This fauna is hugely distributed in an allopatric fashion, although a few springs harbor two or three species. Most of the springs inhabited by hydlrobiids in the region are small, fishless, and have been ignored by state and federal land management agencies. However, many of these sites are degraded by livestock grazing, water withdrawal, anti other activities and will require protection in order to conserve snails and other native aquatic biota. Two of the novellies described herein have become extinct during the past two decades.
This second and final part of a taxonomic treatment of hydrobiid snails of the Great Basin region in the western United States (based principally on material collected during a recently completed field survey) focuses on fauna other than the genus Pyrgulopsis. A new genus of small amnieoline snails, Colligyrlls, is proposed for Hydroia greggi Pilsbry, 1935, together with a new species from the Harney Lake basin of Oregon. This group is strongly differentiated from other amnicolines by a unique female genitalic groundplan. New records are provided for three species of Fluminicola, and two new congeners are described from the northwest Great Basin, both of which had previously been confused with F. turbiniformis (Tryon, 1865). A new genus of cochliopine snails, Eremopyrgus, is erected for a new species from Steptoe Valley, Nevada. Eremopyrgus is distinguished from other cochliopines by unique aspects of its glandular penial lobes and other genitalic features. New records are provided for two species of Tyronia, and a new congener is described from thermal springs in central Nevada. Several new records of Pristincola hemphilli (Pilsbry, 1890) from the extreme northwest Great Basin are provided
The genus Maculinea van Eecke, 1915 (Lepidoptera: Lycaenidae) from the East Palaearctic Region
(1994)
We revise the classification of taxa belonging to the genus Maculinea from the East Palaearctic Region. In this region, in addition to the well-known three species: M. arion (Linnaeus, 1758), M. ationides (Staudinger, 1887) and M. teleius (Bergstriisser, [1779] 1778-1780), two additional species occur: M. alcon ([Denis & Schiffermiiller], 1775) (upper and middle Amur River, Primor'e, China Northeast/Manchuria and North Korea) and M. kurentzovi sp. nov. (upper and middle Amur River, Primor'e, China Northeast and North Korea). Lycaena kondakovi (Kurentzov, 1970) described from Primor'e is a composite species: the lectotype if' designated here represents an East-Asian subspecies of M. alcon, but its single paralectotype is a female to be assigned to M. kurentzovi sp. nov. Only limited numbers of specimens have been known with M. alcon kondakovi from lowlands of "Far-Eastern" Russia and China Northeast, but in North Korea we found a conspicuous allied taxon arirang nov. (female unknown), which we treat here as a highland subspecies of M. alcon but which may actually represent a good species. Of kurentzovi, we have found a series of specimens which have so far been mostly confused with M. teleius in various collections. We treat Glaucopsyche xiaheana Murayama, 1991 from western Gansu as a subspecies of M. arion along with other subspecies from the central and western parts of China: M. adon philidor (Fruhstorfer, 1915) from the east end of the Qilian Range as well as Mongolia, the type locality, and M. arion inferna nom. nov., a replacement name for Lycaena talsienluica (OberthUr, 1910) (praeoccupied) from Tibet, Sichuan and Qinghai. Because of the similarity of male genitalia and existence of intermediate forms, we regard M. sinalcon Murayama, 1992 described from Qinghai as a subspecies of M. teleius despite a few significant characteristics of the holotype. East continental Asia may be regarded as the headquarter of the genus Maculinea.
Glyptostrobus Endlicher is well represented in early Early Cretaceous to Pleistocene deposits in the middle to high latitudes of North America and Eurasia. Although the taxonomy and nomenclature of the genus is complicated, the fossil record indicates Glyptostrobus was represented by a small number of species. The genus first appears in Aptian age deposits from western Canada and Greenland, and achieved a wide distribution early in its evolutionary history. Exchange of Glyptostrobus between Asia and North America occurred across the Spitsbergen and Beringian corridors, which were functional about 110 and 100 million years ago, respectively The Late Cretaceous fossil record of Glyptostrobus shows that the genus had spread into Russia, China and the shores of the Turgai Strait. By the early Tertiary, Glyptostrobus was a prominent constituent of the polar broad-leaved deciduous forests. Paleocene age deposits across western Canada and the United States indicate the genus was present in great abundance in the lowland warm temperate and subtropical forests east of the Rocky Mountains. The broad distribution in North America and Russia during the Paleocene and Eocene indicates that Glyptostrobus grew and reproduced under a diverse range of climatic and environmental conditions, including the cold and unique lighting conditions of the polar latitudes. The presence of Glyptostrobus in Europe indicates the North Atlantic land bridges that extended between North America and Eurasia (Fennoscandia) and Europe during the early Tertiary were used. In Europe, extensive Glyptostrobus dominated swan1ps occupied the Central European Depression during the late Tertiary. Increasing global aridity and cooling, as well as landscape stabilization together with increasing competition for resources and habitat by representatives of the Pinaceae, seem to have forced the genus out of North America, Europe and most of Asia during the Miocene and Pliocene. In Japan, Glyptostrobus persisted until the early Pleistocene. After the early Pleistocene extinction in Japan, Glyptostrobus reappeared in southeastern China. Details of the taxonomic and biogeographic history of Glyptostrobus are examined.
The regular or obligate aphytophagy of certain lycaenid butterflies (Lepidoptera) is discussed within the framework of the most recent general classification of the family. A summary survey of all Lycaenidae known to be aphytophagous is presented, together with a brief account of cannibalism and other opportunistic aphytophagy exhibited by normally phytophagous butterflies. The range of food sources (plants, animals, excretions and regurgitations) exploited by lycaenids is reviewed with emphasis falling on the ecology of myrmecophilous early stages and the significance of their ant-related adaptations. Adult feeding and oviposition behaviour reveal further associations with ants. Specificity oflycaenid/ant relationships and the possible biological effects ofaphytophagy on the Lycaenidae are discussed. Finally, speculations concerning the evolution of aphytophagy by these bulterflies are critically presented.
Epilabidocera amphitrites is one of the most common copepods in the deep waters adjacent to Friday Harbor and shows characteristic swarming behavior in the surface film of the water from later spring through early summer. That the swarms are composed mainly, up to 99 %, of adult males appears to be due to difference in phototaxis to a weak light. This species, at least in copepodid stages, is omnivorous, but seems to prefer an animal diet rather than diatoms. Reproduction takes place continuously from early spring through autumn. The external anatomy of both the female and male has been described in detail. The cuticle forming the arthrodial membrane and the lining of the esophagus, hindgut, and hypostomal and labral troughs appears to be of the same nature throughout, consisting of a single stratum. The cuticle on the general body surface, however, consists of two main strata. The endoskeletal structures consist of two categories, the endoskeleton proper and the endoskeletal tendons. The former involves apodemes and apophyses. Of these the major ones are described in detail. The latter consist of two median tendinous endosternites in the « head », four pairs of ventral intersegmental thoracic tendons, and a pair of dorsal longitudinal tendons in the metasome. The endosternites are well developed, serving as origins for dilators to the atrium oris and esophagus and also for a number of extrinsic muscles to the head appendages. The skeletomusculature may be divided into longitudinal trunk and limb muscles. The paired dorsal and ventral longitudinal trunk muscles in the metasome extend, respectively, from the levels of the cervical groove and the post-maxillulary apodeme to the end of the metasome. The longitudinal trunk muscles in the urosome origate at the anterior end and run most of its length. They are arranged as paired dorsal and ventral groups and a pair of lateral muscles. The extrinsic limb muscles are described in detail. They originate either from the lateral to dorsal exoskeleton or from the endosternites. The digestive tract starts with the atrium oris in the oral cone, followed by the mouth proper, esophagus, midgut, and finally by the hindgut which opens as the anus at the end of the urosome. The oral cone consisting of the three lobed labrum and the paired paragnaths has a longitudinal groove, the oral groove, which is covered ventrally by the spinulose setae of the maxillae and laterally by the gnathobasal endites of the maxillules, these together forming an effective feeding apparatus. The midgut is produced anteriorly into a diverticulum which is higly secretory. In the middle portion of the midgut the epithelial cells are highly vacuolated. As they pass through this vacuolated region the gut contents are cemented into fecal pellets by a mucous secretion and they acquire a peritrophic membrane. There is a strong valve between the midgut and the hindgut. Peristalsis in the midgut is irregular but powerful and primarily in the reverse direction. The circulatory system involves a single heart, enclosed in a large pericardial space, and an anteriorly directed aorta terminating in an anterodorsal aortic sines. The latter communicates through three paires of openings with the sinuses in the head, which are in turn continuous with the perivisceral cavity, from which blood is returned to the pericardium. The heart has the form of a flask with an aortic valve at the tapered anterior end and a posterior ostium. The aortic wall is continued posteriorly over the heart and wraps around the anterior three-fifths as an outer membrane. This outer membrane is extended dorsally at three places to attach the heart to the dorsal exoskeleton; and it is also drawn out ventrally to form the anterior and lateral walls of the pericardium. These walls are continuous with the pericardial floor which seals the pericardia! cavity from the perivisceral cavity. The heart-beat and the blood flow through the system have been discussed. The excretory system consists of a pair of maxillary glands, each comprising a coelomic end-sac, a coelomic secretory tubule and an ectodermal excretory duct. The end-sac communicates with the tubule through a valvular opening. Antennary glands are not gound either in the nauplius stage or in the adult. The male reproductive system consists of a single testis and a single genital duct which is divided into four differentiated sections, the vas deferens, the seminal vesicle, the spermatophore sac, and the ductus ejaculatorius. The vas deferens is a thick-walled glandular tube secreting the various constituents of the spermatophore. The seminal vesicle serves mainly as a reservoir for the various components of a definitive spermatophore, and it is here that these take up their final positions. The spermatophore sac is highly glandular and is mainly responsible for formation of the coupling apparatus of the spermatophore. The spermatophore is not open directly to the outside but is connected with a canal system in the coupling apparatus. When transferred to the female genital segment at copulation, the central secretion of the spermatophore is discharged through the canal system of the coupling apparatus to glue down the spermatophore. A duct through which the spermatozoa can pass from the spermatophore to the spermathecae of the female appears to be formed later by an action of the female, possibly secretion of an enzyme or lysin. The discharge of the contents of the spermatophore is effected by swelling of Q-spermatozoa in the distal region of the spermatophore. The functional spermatozoa are spherical or polygonal and nonmotile. The female reproductive system consists of a single ovary, two oviducts, each with several diverticula, leading to the paired opnenings into the vaginal vacity, a pair of spermathecae and a pair of glands which open into the oviducts. In the mature female the oviducts are wide and sac-like, expanded by growing oocytes. However, the last portion of the oviduct is usually empty of eggs and is highly secretory. The oldest oocytes in the oviducts are usually at the metaphase of the first maturation division. The evidence points to the conclusion that the eggs are laid in this stage, and they are fertilized when they pass through the vaginal cavity. Oogenesis has been studied in detail. There are two periods of yolk formation: the first immediately after the dispersion of the mitochondrial bodies and the second in the last phase of the oocyte growth when the vacuoles in the cytoplasm are gradually replaced by yolk. Two dorsal ocelli, in the copepodid stages, are placed dorsolaterally against the exoskeleton and highly developed, each with a perfectly spherical, cuticular lens, while a single ventral ocellus remains unspecialized through the copepodid stages. Each dorsal ocellus proper is suspended in the head sinus by several connective tissue stands in addition to an aye muscle and consists of a large, syncytial pigmented cup occupied by a cellular sphere which is composed of 9 retinular and 4 crystalline cells. Each of the 9 retinular cells gives off an axon which leaves the ocellar cup at one of three places to proceed to the nauplius eye center in the protocerebrum. The ventral ocellus consists of two multinucleated pigmented cells, a cup-shaped tapetum, 6 retinular cells and about 8 conjunctival cells. Each of the 6 retinular cells sends an axon which loops over the posterior rim of the ocellar cup in common with the others to course to the nauplius eye center in the protocerebrum. The ventral ocellus is innervated by two afferent nerve fibers. There is also found a pair of conspicuous nerve fibers, possibly afferent, associated with the dorsal and ventral ocelli. A pair of accessory retinular groups, each consisting of three retinular cells, is found posterior to the dorsal ocelli. Three efferent aXOl1S from each group form a nerve running to the nauplius eye center in the protocerebrum. A pair of frontal organs, each innervated by a frontal nerve, lies in the anterior end of the head. The frontal nerves can be traced up to a pair of neuropiles immerdiately ventral to the nauplius eye center in the proto cerebrum. A pair of suprafrontal nerves branched off from the frontal nerves is found to innervate a pair of sensory filaments, the suprafrontal sensiIla, at the lower anterior end of the head. The central nervous system, consisting of a well developed brain connected by massive circumesophageaI connectives to the ventral nerve cord, has been described in detail. The ganglion cells are found throughout the nerve cord, and they are arranged into ganglia in the thoracic segments bearing the swimming legs. The stomatogastric nervous system has two pairs of labral and a single gastric ganglia. The medial pair of the labral ganglia forms anteriorly a single ganglion which is connected to the brain by three small nerves. The giant fiber system, consisting of giant motor fibers and giant interneurons, has been studied in detail, and it appears to constitute the effector portion of an escape reflex. The cutaneous glands opening through small pores in the cuticle of the metasome, urosome, and the appendages have been described. Chromatophores, unicellular or syncytial with several nuclei, are scattered deep in the body and are responsible for the metachrosis.
During the last decade, three new acidophilous forests associations were detected in the Mecsek Mts (SW Hungary), and described as acidophilous beech wood (Sorbo torminalis-Fagetum (A. O. Horvat 1963a) Borhidi et Kevey in Kevey 2001), acido-mesophilous oak wood (Luzulo forsteri-Quercetum petraeae (A. O. Horvat 1963a) Borhidi et Kevey 1996) and acido-xerophilous oak shrubland (Genisto pilosae-Quercetum polycarpae (A. O. Horvat 1967) Borhidi et Kevey 1996). In this article two further new associations are described: the acidophilous oakwood of the Mecsek (Viscario-Quercetum polycarpae Kevey, ass. nova) and the acido-mesophilous oakwood of western Hungary (Campanulo rotundifoliae-Quercetum petraeae (Csapody 1964) Kevey, ass. nova). These associations are related to the acidophilous forests of the Balkan Peninsula based on the infrequent presence of sub-Mediterranean species. A detailed comparative study of these new associations with the earlier known ones permitted to develop a reshaped classification of the syntaxonomy of these units, creating four new suballiances: within the frame of Quercion farnetto I. Horvat 1938 the suballiances Luzulo forsteri-Quercenion polycarpae Kevey, suball. nova and the typical Quercenion farnetto Kevey, suball. nova, in the frame of Quercion petraeae Zolyomi et Jakucs 1957 the suballiances Luzulo multiflorae-Quercenion petraeae Kevey, suball. nova and the Quercenion petraeae Kevey, suball. nova.
5-lipoxygenase (5-LO) is the key enzyme in the formation of inflammatory leukotrienes, which are mediators of inflammation and allergy. The 5-LO catalyses the oxidation of arachidonic acid to 5-HPETE and subsequently to LTA4. The leukotrienes are involved in the development and maintenance of inflammatory diseases, like asthma and allergic rhinitis. Additionally, 5-LO is overexpressed in some cancer types, although its relevance is still not fully understood. 5-LO expressing cells are B- lymphocytes and cells of myeloid origin like monocytes, macrophages and granulocytes. The 5-LO promoter lacks a TATA or CCAT box and covers two CpG islands. These are characteristics of a housekeeping gene, but as the 5-LO is not expressed ubiquitiously, the expression of the 5-LO is tightly regulated. Epigenetic mechanisms were known to be involved in the control of the 5-LO expression. The HDAC inhibitor TsA significantly induced the transcriptional activity of the 5-LO promoter in reporter gene assays as well as on 5-LO mRNA transcript level in MM6 cells. The GC-boxes GC4 and GC5 in the proximal 5-LO promoter were identified to be essential for the TsA effect, as deletion of these element led to an attenuated TsA effect in reporter gene assay. Recruitment of the transcription factors Sp1 and Sp3 and the RNA polymerase II to the 5-LO promoter was detectable after TsA treatment in MM6 cells by chromatin immunoprecipitation assays (ChIP), while the acetylation status of histone H4 remained unchanged. Likewise it is known that DNA methylation leads to silencing of 5-LO expression in-vitro and in-vivo. The 5-LO promoter is densely methylated in the cell line U937, but unmethylated in HL-60 cells and - elucidated in this study - also in MM6 cells. Reporter gene assays with in-vitro methylated 5-LO promoter containing plasmids revealed that the frequency of methylated CpGs is directly proportional to reduction of 5-LO promoter activity. Incubation of U937 cells with 5-AdC, an inhibitor of DNA methyltransferases, was able to reactivate 5-LO transcription and to demethylate CpG dinucleotides. In the first part of this study the mechanism of TsA induced promoter activation was further investigated. I elucidated the mechanism of Sp1 and Sp3 recruitment to the 5-LO promoter after TsA treatment. Immnoprecipitation assay was used to detect a transcription factor complex containing Sp1 or Sp3 interacting with HDAC proteins, which might change its composition after TsA treatment. Besides the posttranslational modifications of the transcription factors Sp1 and Sp3 after TsA treatment were investigated, potentially causing an increased interaction of the proteins with the 5-LO promoter. Both aspects and their response in HDAC inhibition have been described. TsA did not affect the composition of the Sp1/HDAC1/HDAC2 complex. Sp3 was not located in a complex with the HDAC enzymes. Acetylation of Sp1 and Sp3 was detectable, but no change occurred after TsA treatment. Since neither release of the transcription factors off a complex, nor alterations in posttranslational modifications of Sp1 and Sp3 are the reason for the increased Sp1 and Sp3 binding to the 5-LO promoter, I elucidated alterations in the chromatin structure. The acetylation status of the histone proteins H3 and H4, as well as the chromatin marks H3K4me3, representing active chromatin, and H3K9me, representative for repressive state, were investigated. Additionally, the time course of the TsA effect was determined on 5-LO mRNA level using real-time PCR. The acetylation status of the histone proteins on the 5-LO core promoter correlated with the basal 5-LO mRNA transcript expression in MM6, HL-60 and U937 cells. The highest 5-LO mRNA level was detectable in MM6 cells, followed by HL-60 cells. The lowest 5-LO mRNA level was detected in 5-LO promoter methylated U937 cells. The order of the basal 5-LO mRNA expression of the three cell lines correlates with the basal acetylation status of histone proteins H3 and H4. In MM6 cells the highest basal levels in acH3 and acH4 were detected, followed by HL-60 and U937 cells. Moreover, the data obtained in U937 cells revealed that the correlation between DNA methylation and histone hypoacetylation is alike on the 5-LO promoter. TsA treatment induced the 5-LO mRNA level in the three cell lines with different intensity: 5-LO mRNA level in MM6 cells was induced 11-fold, in HL-60 cells 6- fold and in U937 cells 4- fold. The histone acetylation and methylation levels on the 5-LO promoter after TsA incubation were investigated. No increase in acH3 and acH4, but in H3K4me3 was detectable in MM6 cells by ChIP assay. HL-60 cells showed an increase in acH3 and acH4 as well as in H3K4me3. H3K9me was only detectable in untreated U937 cells, but disappeared after TsA treatment, while acH3, acH4 and H3K4me3 increased constantly after TsA treatme nt. A strong correlation between the histone modifications and the time course of the mRNA expression was detectable in all three cell lines. The combination of the posttranslational modifications acH3, acH4 and H3K4me3 led to a fast effect in transcriptional activation and the maxima of acH3 and acH4 were usually associated with the maximum in 5-LO mRNA transcript level. An increase in H3K4me3 alone, as detected in MM6 cells, led to continuous increase in the 5-LO mRNA expression with a late maximum. Additionally, we detected a slight overall decrease in 5-LO promoter methylation in U937 cells after TsA treatment. This fact taken together with the observed histone modifications could explain the 4- fold response in 5-LO mRNA level to TsA treatment of the methylated cell line U937. Another aim of the present study was to identify the specific HDAC enzymes involved in the 5-LO promoter regulation. Reporter gene assays and real-time PCR with selective HDAC inhibitors revealed that HDACs of class I are involved in 5-LO promoter regulation, namely HDAC 1, 2 and 3. The influence of each of the enzymes seemed to depend on the cell type, as inhibition of HDACs 2, 3 strongly induced 5-LO promoter activity in reporter gene assay in HeLa cells, whereas in MM6 cells HDACs 1 and 2, 3 seemed to be responsible for the 5-LO promoter regulation, measured as 5-LO mRNA level. The HDACs of class IIa and class III are not involved in the regulation of 5-LO mRNA expression. The second part of this study investigated the influence of MBD proteins on the methylated 5-LO promoter and the 5-LO mRNA expression. ChIP assays revealed MBD1, 2 and MeCP2 protein binding to the proximal 5-LO promoter in U937 cells. MBD1 was detectable on the 5-LO promoter in unmethylated HL-60 cells, while no MBD protein was located on the 5-LO promoter in MM6 cells. To elucidate the functional role of the MBD proteins, stable knocked down of MBD proteins was established in U937 cells. 5-LO mRNA transcript level was determined in the knock down clones by real-time PCR. The 5-LO transcript level was increased in all knock down samples. MBD2 knock down clones showed the highest effect in activating 5-LO with a 3- and 4.4-fold increase in the 5-LO mRNA level, followed by MBD1 (3.5- fold) and MeCP2 (2.5-fold) knock down clones. A combined participation of these three enzymes in the corepression of the methylated 5-LO promoter is indicated. Taken together, the data reveal that epigenetic mechanisms are strongly involved in the regulation of 5-LO transcription and might function as a crucial control mechanism of 5-LO expression.
Westwoodilla Bate. 1862 is revised and 12 species are recognized as distinct, 2 of which, W. tone and W. helle. are described as new. New characters of taxonomic importance are described and illustrated. The many variable characters that have led to taxonomic confusion in the pas! are converted to morphometries and the intra- and interspecific variation are illustrated. This has led to a new understanding of the genus and a clarification of the "W. caecula complex". The type species Westwoodia caecula Bate. 1857 is redescribed and a neotype is designated. W. megalops (Sars, 1883) is resurrected. The junior synonymy of W. longidactyla Carausu, 1949 with W. caecula is supported by the find of Carausu's original unpublished notes and drawings. Notes on undescribed species are given. New drawings are provided for W. manta Griffiths, 1974, W. longimana Shoemaker, 1934, W. brevicalcar (Goes, 1865) and W. megalops. All species are diagnosed; new distributions are given and plotted on a map; and a key to the named species is included.
Amphibians of Malawi : an analysis of their richness and community diversity in a changing landscape
(2009)
This study summarizes the state of the knowledge of the amphibian diversity in Malawi highlighting the possible threats impending on this fauna correlated with human encroachment and land use change. New data about diversity, distribution and ecology have been gathered, whereas the old ones have been summarised, reviewed and commented. In order to put in context the responses of the amphibian communities to land use change, the main environmental characteristics of the country at a broad space and time scale have been explored. Furthermore, the original habitats and vegetation have been described, and their status in the present day Malawi discussed. In the same way, an overview of the actual state of the knowledge about the Malawian amphibians has been provided, and their ability to act as surrogate of environmental integrity in Sub-Saharan Africa commented on the basis of the available studies. Afterwards, the results of the study of the selected areas and samples have been analysed within this newly generated context. Different field and laboratory methods were applied for the quantitative analysis of the richness and diversity of the communities. Opportunistic search was used to detect species richness, whereas the visual encounter survey was applied to detect the relative abundance of species. Several indices of diversity and similarity, and extrapolations by means of true richness estimators were used for the analysis of the alpha and beta diversities. Additional information were gathered by means of pitfall traps with drift fence, and by the recording of the advertisement calls. Supplementary methods were applied for the analysis of the taxonomic composition of the collected material. In Malawi 84 amphibian species are recorded, two of which still undescribed (Leptopelis sp. and Phrynobatrachus sp.). Three further species need to be confirmed and might be possibly present too: Amietia viridireticulata, Hemisus guineensis, and Hyperolius minutissimus. Additionally, other unrecognised cryptic species — at least one — are present within the Hyperolius nasutus complex. Most of the species belong to the order Anura (82 species; 97.6%), whereas only two species belong to the Gymnophiona (2.4%). Anurans are divided into 12 families and 23 genera, whereas the two caecilians species into one family (Caecilidae) and two genera. The more diverse family is the Hyperoliidae (21 species, 25%) followed by the families Ptychadenidae (13 species, 15%), Arthroleptidae (11 species, 13%), Phrynobatrachidae (10 species, 12%), and Bufonidae and Pyxicephalidae (9 species, 11% respectively). The remaining high family diversity (seven families, Caecilidae included) is contrasted by a low number of species (11 species in total, 14%). Based on the available distribution data, the value of species richness of the anuran communities in Malawi is comprised between 5‒45 species. In average 16.8 ± 9.0 species (N=80) are to be found, 75% of the sites have less than 21 species, and only two sites have more than 25 species. Four hot spots of amphibian diversity were identified: the Nyika Plateau (24 species), Mangochi-Malombe (25 species), Zomba Plateau (32 species) and the Mulanje Massif (45 species). In the studied areas a mean of 14.7 ± 1.6 species was observed and extrapolations by means of the true richness estimators were in good agreement with this result. Among the studied areas the richest was Palm Forest Reserve (17 species), followed by Kaningina Forest Reserve (16 species) and Vinthukutu F. R., and Vwaza W. R (15 species). The poorest area was the Misuku Mountains with 12 species only and a slightly different ranking was generated by the true richness estimators. The mean of the species present in the samples was 4.8 ± 2.1 species, considerably less than the true species richness detected in the respective areas. Basing on the ranking generated by the K-dominance plot the most diverse samples were Palm F. R. and Misuku, whereas the less diverse were Kaningina F. R. and Fort Lister, confirmed by the values of the diversity indices. The main finding of this study was the observation of the lack of a clear match between environmental degradation and amphibian diversity, and the crucial importance of temporary water bodies for the preservation of the amphibian diversity. In fact, despite most of the original habitat formerly present in Malawi have been destroyed and replaced by cultivations, the amphibian communities of different areas showed a comparable diversity at both family and species richness level, and no evident match between environmental degradation and amphibian diversity was recognisable. Differences in species richness could mostly be explained by natural factors such the elevation gradient and the presence of temporary water bodies. However, it was not possible to exclude that the communities have changed during historical time and the shift in species composition already occurred together with the modification of their relative frequencies. Most of the species showed a remarkable ecological plasticity and several species were found in a variety of both natural and altered habitats. The classification of the Malawian amphibians on the basis of ecological guilds based on the available natural history data showed the preponderance (76%) of generalist pond breeders. As a consequence, most of these amphibians possessed a scarce capacity to act as surrogates of habitat integrity. Based on the result of this study the farm bush landscape with traditional agriculture practices bears a great potential to support amphibian diversity in terms of species richness, representing a compromise between local economic development and conservation. Furthermore, the results of this study indicate the outstanding importance of the southern-east region of Malawi for the conservation of the country’s amphibians.