Study of the late embryogenesis of Daphnia (Anomopoda, ‘Cladocera’, Branchiopoda) and a comparison of development in Anomopoda and Ctenopoda

The embryonic development of Daphnia galeata and D. hyalina (`Cladocera', Anomopoda, Daphniidae) has been investigated by observing living embryos removed from female brood pouches. The sequence of morphological changes was analysed, as was the time at which the activity of certain organs began. The timing of these events at 22 °C is documented for both species.These data were compared with similar information, previously obtained for two representatives of the Ctenopoda (Kotov & Boikova, 1998). The sequence of events is basically similar in the two groups during early and late phases of their development, but the time of shedding of the embryonic membranes is different in the Anomopoda and Ctenopoda. The ctenopod embryo hatching from the second egg membrane is covered by the third membrane, which will be cast some hours later. The anomopod embryo hatches from the second egg membrane approximately simultaneously with the shedding of the third membrane, and it is covered already by the fourth membrane after the shedding of the second egg membrane.Earlier (Kotov & Boikova, 1998), we determined four embryonic instars in the course of the development of the Ctenopoda. Two of them are passed within the egg membranes, the next two instars occur after the shedding the egg membranes within the mother's brood pouch. However, in anomopods, one of the latter (the third) occurs within the second egg membrane, one is incorporated into the egg. Thus, the development of the Anomopoda is more embryonized in comparison with that of the Ctenopoda.     

Phylogeny and adaptive radiation within the Anomopoda: a preliminary exploration

The distinctness of the Anomopoda and the polyphyletic nature of the so-called Cladocera are emphasized.An attempt is made to reconstruct the ancestral anomopod, which probably lived in Palaeozoic times. This task is facilitated by the availability of detailed information on extant forms, which includes functional as well as purely morphological considerations and enables us to understand the means whereby complex mechanisms were transformed during evolution. Comparative studies on the ecology and habits of extant forms also throw light on the probable way of life of the ancestral anomopod.Adaptive radiation within the Anomopoda is briefly surveyed and an outline of the suggested phylogeny of the order is indicated.Institute of Environmental and Biological Sciences, University of Lancaster     

Evolutionary morphological development of the cladocera of the superfamily sidoidea and life strategies of crustaceans of continental waters

Comparative-morphological analysis of the structure of the Cladocera of the superfamily Sidoidea ( = Ctenopoda) comprising the families Sididae and Holopediidae, compared with representatives of other groups: Chydoroidea (=Anomopoda), Limnadiiformii (= Conchostraca), Polyphemiformii (Polyphemiformes (= Onychopoda) and Leptodoriformes (= Haplopoda)) and Branchipodiones (Branchipodiformes (= Anostraca) and Triopiformes (= Notostraca)) is made. The main parameters and structures important in functional-morphological and ecological aspects are considered: including dimensions of the body, its form and proportions, the complex eye, swimming antennae, body covers, thoracic limbs, the postabdomen. In distinction from all crustaceans under consideration in the Sididae and the Polyphemiformii organs contributing to rapid manoeuvrable swimming, soaring and good orientation in the environment received the prevailing development in evolution: antennae, stabilizing appendages of the posterior part of the body, the eye, etc. All this predetermined elaboration of the life strategy in these forms mainly aimed, especially in the Sididae, at active avoidance of predation combined with elements of passive protection (imperceptibility attained by various methods). Representatives of the Sididae, including planktonic Diaphanosoma, well survive in water bodies with the intensive press of predation. All others, including the Holopediidae, clearly possess the passive protection (development in localities inaccessible to predators, imperceptibility, protective structures on the body surface, etc.).     

On the ontogeny of Leptodora kindtii (Crustacea,Branchiopoda, Cladocera), with notes on the phylogeny of the Cladocera

Leptodora kindtii, a large predaceous cladoceran, is among the most deviant species of the Cladocera. Therefore, its phylogenetic position has traditionally proven difficult to determine. Its many peculiar features include, among others, long, stenopodous, forwardly directed trunk limbs, a posteriorly placed dorsal brood pouch, a tri-lobed lower lip, and a long, segmented abdomen. This study describes the ontogeny of L. kindtii (Haplopoda), including general body proportions, appendages, the carapace, and other external structures in an attempt to facilitate the comparison of its aberrant morphology to that of other branchiopods. In general, the early embryos are similar to the early embryos of other cladoceran taxa with respect to body shape and size and position and orientation of the early limb buds. Many of the unusual features of L. kindtii appear late in ontogeny. The carapace appears at an early stage as a pair of dorsolateral swellings in a position corresponding to the gap between the mandibles and the first pair of trunk limbs; it later becomes posteriorly transposed by a gradual fusion of its more anterior parts to the dorsal side of the thorax. The tri-lobed "lower lip," under the labrum of the late embryo and the adult, develops as a fusion of the first maxillae (lateral lobes) to an elevated sternal region behind the mouth (median lobe). The stenopodous, segmented trunk limbs in the adult develop from embryonic, elongate, subdivided limb buds, similar to those seen in early stages of other branchiopods. Two conflicting possibilities for the phylogeny of the Cladocera, involving two different positions of L. kindtii (Haplopoda), are discussed. Several characters support a sister-group relationship between the Haplopoda and Onychopoda. However, some characters support the Anomopoda and Onychopoda as sister groups, leaving the Haplopoda outside this clade. In contrast to recent suggestions, we prefer to retain the term "Cladocera" in its original sense as comprising the Haplopoda, Ctenopoda, Anomopoda, and Onychopoda.     

The phylogenetic relationships of "predatory water-fleas" (Cladocera: Onychopoda, Haplopoda) inferred from 12S rDNA

Within the Cladocera, the water-fleas, four major taxa can be distinguished: Anomopoda, Ctenopoda, Haplopoda, and Onychopoda. Haplopoda and Onychopoda are called "predatory water-fleas." The Haplopoda is monotypic; its only representative, Leptodora kindtii, is common in palearctic and nearctic freshwater bodies. The Onychopoda show a remarkable geographic distribution. Most of the described species are restricted to the Caspian Sea, the Aral Sea, and peripheral areas of the Black Sea, including the Sea of Azov--all remnants of the Eastern Paratethys. The remaining onychopods are either freshwater inhabitants or marine animals, widespread in the world oceans. We present molecular evidence for a sister group relationship between Haplopoda and Onychopoda within the Cladocera. The Onychopoda and its three families are monophyletic. We suggest an independent invasion into the Ponto-Caspian basin at least three times, twice originating in the palearctic freshwater bodies and once starting from the world oceans.     

The Branchiopoda (Crustacea: Anomopoda, Ctenopoda and Cyclestherida) of the rain forests of Cameroon, West Africa: low abundances, few endemics and a boreal–tropical disjunction

Aim We provide the first in‐depth study of the Branchiopoda of the rain forests of Cameroon and also of the African continent. Location Surface water environments, Cameroon. Methods Qualitative plankton samples were collected in all types of surface water environments present, ranging from big lakes to water collected in rock crevices or fallen fruit cavities. A tow or hand‐held plankton net of mesh size 100 μm was used, and water volumes filtered were at least several m³ in large water bodies, or half to whole water volume in small water bodies. Results We recorded 61 species (53 first records for the country), based on 700+ samples collected between September 1998 and March 2002. Anomopoda (92%) was the dominant order, followed by Ctenopoda (6.5%) and Cyclestherida (1.5%). Chydoridae (67%) was the most speciose family followed by Macrothricidae (6.5%) and Daphniidae (5%). Alona (11%) was the dominant genus followed by Chydorus (10%) and Pleuroxus (8%). Several species of Chydorinae, especially of the genus Pleuroxus, are shared with continental Eurasia–North America, but are absent from the Mediterranean and desert–steppe–savanna zones of Africa (boreal–tropical disjunction). Daphnia was absent, as in most tropical lowlands. No single species was really abundant, and a majority were rare to very rare, and of restricted occurrence within the rain forest patches. Comparing Africa, South America and Southeast Asia, we found a current total of 196 species for the combined rain forest areas, out of a world total of 500+ species. Systematic trends in richness at three taxonomic levels were the same for all continents: Anomopoda–Ctenopoda–Cyclestherida at ordinal level, Chydoridae–Daphniidae–Macrothricidae–Sididae at family level and Alona–Chydorus–Macrothrix–Diaphanosoma at genus level. Southeast Asia was richest (111 species, 14 endemics) with South America a close second (110 species, 27 endemics). Africa was the most species‐poor (95 species, of which only 5 are endemics). Main conclusions We hypothesize that the post‐Miocene cooling and aridization of the world climate hit the freshwater biota of Africa particularly hard, with more extinction here than elsewhere, and little recolonization. Most extinction occurred in the savanna‐desert belt, and eight disjunct boreal species (four Pleuroxus, Picripleuroxus laevis, Kurzia latissima, Alonella exigua, and Monospilus dispar) survive morphologically unchanged since pre‐Pleistocene times in the Cameroon rain forest. Slow evolution thus appears typical of these cyclic parthenogenetic branchiopods in which sexual recombination occurs only at intervals. Illustrative of the same slow evolution is the fact that the two endemic cladocerans of Cameroon (Nicsmirnovius camerounensis and Bryospilus africanus) belong to tropicopolitan genera of Gondwanan age.     

Notothrix halsei gen. n., sp. n., representative of a new family of freshwater cladocerans (Branchiopoda, Anomopoda) from SW Australia, with a discussion of ancestral traits and a preliminary molecular phylogeny of the order

The parthenogenetic female of a new anomopod crustacean, Notothrix halsei gen. n. et sp. n., is described from Lake Angove and found in five other sites in SW Australia. Although macrotrichid-like at first glance, a detailed morphological study including thoracic appendages revealed a set of primitive characters and similarities with primitive families, in particular with the Eurycercidae and Acantholeberidae. Ancestral traits of Notothrix gen. n. comprise: (i) a single large head pore; (ii) second maxilla; (iii) incomplete dorsal merger of valves; (iv) second antenna with a three- and a four-segmented branch; (v) six trunk limbs; (vi) two setae on a sixth endite in the first limb, vs. complete row of posterior setae on the second limb; (vii) large number of setae on exopodites III–V; (viii) well-developed gnathobase V; and (ix) massive postabdomen with wide, telson-like terminus between end-claws that show complex spinulation but no basal spine. Analysis of the complete 18S (SSU) rRNA gene sequence including taxa from all but four anomopod families and using Ctenopoda as an outgroup, confirms our separation of the new taxon, the Nototrichidae fam. n. Although different alignments resulted in different trees, the analyses confirm an isolated position of Notothrix , which has the shortest SSU rRNA gene length known for anomopods. However, the phylogenies derived from a comparison of 18S rRNA gene sequences between families of the order Anomopoda, does not resolve the descent of the various family groups.     

A reclassification of the anomopod families Macrothricidae and Chydoridae,with the creation of a new suborder,the Radopoda (Crustacea: Branchiopoda)

An investigation, using optical microscopy and SEM, of the trunk limbs of the Anomopoda has revealed a large number of characters, previously underused or unused in taxonomy and comparative morphology. All these characters, which are nicely paralleled by some more conventional traits (head shield and pores, postabdomen, antennae ...), show one clear tendency across all groups studied: a state of complexity at one extreme, and a state of often incisive simplification at the other extreme, with a number of transitional stages in between. The complex character state, which itself is a simplification of the leg structure of the Ctenopoda and other, large Branchiopoda, is here considered to represent a primitive condition. The simplified state is considered advanced. Based on this assumption, we list a number of unifying characters (mainly structural aspects of P1 and P2, but also the gnathobase of P3 and P4) for all macrothricid and chydorid-like anomopods, which we unite in the new suborder Radopoda. Non-radopod Anomopoda are not reclassified. We then derive a cascade of (mainly trunk-limb based) characters to work out a hypothesis on the evolution of the Radopoda. The chydorid line (basically the former family Chydoridae) is classified as a superfamily (the Eurycercoidea), with three families; the macrothricid line is capped by the superfamily Macrothricoidea, with four families. Of these seven families, four are upgraded from subfamily status, the Chydoridae are left status quo, the Macrothricidae are redefined, and the Neothricidae are a new family. The Macrothricidae are further subdivided in two subfamilies, of which the Macrothricinae appear reasonably homogeneous (monophyletic), while the non-Macrothricinae require further study. Some of these (e.g. Guernella) have almost completely lost their P5, a situation parallel to that of the P6 in the Eurycercidae, Acantholeberidae, and Ophryoxidae.     

A daphniid ephippium (Branchiopoda: Anomopoda) of Cretaceous age

An anomopod ephippium from the Lower Cretaceous of Australia, remarkably similar to that of Simocephalus , shows that daphniids whose ephippia at least were very similar to those of extant species, existed about 120 million years ago. The family clearly originated in pre-Cretaceous times, and the Anomopoda considerably early than this.     

The Interaction of Acridine Dyes with the Densely Packed DNA of Bacteriophage

The interactions of acridine dyes with intact phage DNA differ from those with extracted DNA in the following respects. Strong binding (intercalation) is greatly reduced in intact phage but probably not eliminated. The cooperative, weak binding is stronger and the stacking tendency is increased. In gels of DNA the stacking tendency is seen to increase with decreasing hydration. These influences of the dense packing of DNA must be taken into account when using basic dyes to study chromosome structure.     

The origin and evolution of variable-region helices in V4 and V7 of the small-subunit ribosomal RNA of branchiopod crustaceans

We sequenced the V4 and V7 regions of the small-subunit ribosomal RNA (SSU rRNA) from 38 species of branchiopod crustaceans (e.g., Artemia, Daphnia, Triops) representing all eight extant orders. Ancestral large-bodied taxa in the orders Anostraca, Notostraca, Laevicaudata, and Spinicaudata (limnadiids and cyzicids) possess the typical secondary structure in these regions, whereas the spinicaudatan Cyclestheria and all of the cladocerans (Anomopoda, Ctenopoda, Onychopoda, and Haplopoda) possess three unique helices. Although the lengths and primary sequences of the distal ends of these helices are extremely variable, their locations, secondary structures, and primary sequences at the proximal end are conserved, indicating that they are homologous. This evidence supports the classical view that Cladocera is a monophyletic group and that the cyclestheriids are transitional between spinicaudatans and cladocerans. The single origin and persistence since the Permian of the unique cladoceran helices suggests that births and deaths of variable region helices have been rare. The broad range of sequence divergences observed among the cladoceran helices permitted us to make inferences about their evolution. Although their proximal ends are very GC-biased, there is a significant negative correlation between length and GC content due to an increasing proportion of U at their distal ends. Slippage-like processes occurring at unpaired nucleotides or bulges, which are very U-biased, are associated with both helix origin and runaway length expansion. The overall GC contents and lengths of V4 and V7 are highly correlated. More surprisingly, the lengths of these SSU rRNA variable regions are also highly correlated with the length of the large-subunit rRNA expansion segment, D2, indicating that mechanisms affecting length variation do so both across single genes and across genes in the rRNA gene family.     

Characteristics of the sperm tail of the velvet worm, Euperipatoides leuckarti (Onychophora)

The onychophoran sperm tail contains several kinds of microtubulcs; probably more than that of any other animal group. There are thus a peripheral manchette consisting of many tightly spaced microtubules, a ring of nine 'peripheral singlets' and a central axoneme of the classical 9 + 2 type (nine doublets and two central singlets). The protofilament organization of these various microtubules was examined and compared to the structure and mode of formation of the peripheral singlets with that of its analogues in other animal groups. The onychophoran peripheral singlets were found to differ in two respects from those in insects: they are formed from the manchette rather than from the axonemal doublets and their transient connection to the axoneme is to the A-subtubules of the doublet rather than to the B-subtubules. The manchette microtubules as well as the peripheral singlets consist of 13 protoh'laments. The manchette may serve a mechanical function (to strengthen the unusually thick sperm tail) hut the role of the peripheral singlets remains unknown.     

Stationary phase deletions in Escherichia coli. I--Evidence for a new deletion pathway

Deletions in the plasmid pMC874 join the promoter of the km(r) (kanamycin resistance) gene coding for the enzyme aminoglycoside 3'-phosphotransferase to a promoterless lac operon downstream giving a phenotypic change from Lac(-)-->Lac(+). They differ from most deletions studied in Escherichia coli, which occur in actively dividing cells, in several important respects, as follows. (1) They occur in "resting" cells incubating on McConkey's or minimal lactose agar and increase in number gradually over a period of 1-2 weeks. Thus, like "adaptive" mutations, they are time rather than generation dependent. (2) They are extremely rare events (frequency 1x10(-11)-5x10(-11)) in wild type cells, but their frequency is increased between 1 and 2 orders of magnitude by null recC(-) mutations. In these respects they differ from "adaptive" mutations which are equally frequent in recC(+) and recC(-) cells. (3) Their frequency is not increased by mutations which stimulate log phase deletions. (4) Based on a computer search for homologies and sequencing of one deletion, it appears that they differ from log phase deletions in that they can occur in the absence of major terminal homologies (direct repeats) or intervening homologies (inverted repeats) which could stabilize a transient secondary structure and determine the deletion endpoints. Thus, they are not explained by the misaligned mutagenesis model. In conclusion, resting phase deletions occur through a totally different pathway from deletions in actively dividing cells and probably originate from unrepaired double strand breaks.     

Phylogenetic relationships within the Phyllopoda (Crustacea,Branchiopoda) based on mitochondrial and nuclear markers

For several decades the relationships within the Branchiopoda (Anostraca + Phyllopoda) have been a matter of controversy. Interpretations of plesiomorphic or apomorphic character states are a difficult venture, in particular in the Phyllopoda. We explore the relationships within the Phyllopoda at the level of nucleotid comparisons of the two genes 12S rDNA (mitochondrial) and EF1alpha (nuclear), and at a higher molecular level based on introns found in the gene EF1alpha. Within the Phyllopoda our explorations show further evidence for a non-monophyletic Conchostraca (Spinicaudata + Cyclestherida + Laevicaudata). The monotypic Cyclestherida is more closely related to the Cladocera, both together forming the Cladoceromorpha. The Spinicaudata (Leptestheriidae, Limnadiidae, and Cyzicidae) is well supported. Spinicaudata and Cladoceromorpha form a monophylum. The position of the Laevicaudata remains unclear but we find neither support for a sister group relationship to the Spinicaudata nor for a close relationship of Laevicaudata and Cladocera. Within the Cladocera, we favour the Gymnomera concept with the monotypic Haplopoda being the sister group to the monophyletic Onychopoda. The Ctenopoda seems to be the sister group to the Gymnomera, which contradicts the common view of a more basal position of the Ctenopoda.     

Dumontia oregonensis n. fam., n. gen., n. sp., a cladoceran representing a new family of `Water-fleas' (Crustacea,Anomopoda) from U.S.A., with notes on the classification of the Order Anomopoda

Dumontia oregonensis, a cladoceran representing a new family in the Order Anomopoda is described from rain pools in the Agate Desert, Oregon, U.S.A. The proposed family, Dumontiidae, is the newest family within Anomopoda that is not just a reshuffling of already-known species. The general appearance of this novel cladoceran is similar to that of members of the family Macrothricidae. However, a detailed examination of the trunk limbs, particularly of the second pair of limbs, showed that the new species lacks the scraper-setae typically observed in all members of the recently erected suborder Radopoda, to which macrothricids belong. Instead, limb morphology suggests a closer relation of Dumontiidae to the family Daphniidae. Dumontiidae appears to be a `missing' link between the suborder Radopoda and the `non-radopodid' anomopods. The families Daphniidae, Ilyocryptidae, Bosminidae, Moinidae and the new Dumontiidae are similar in that they lack of typical radopodid setae on the second pair of trunk limbs. Further studies on the limb morphology of non-radopodid cladocerans are required to solve the phylogenetic relationships among the members of the order Anomopoda.     

Adaptations of Anomopoda crustaceans (Cladocera) to the benthic mode of life

Crustaceans of the order Anomopoda inhabit both pelagic and littoral zones of fresh water bodies. Different adaptations to the benthic mode of life in representatives of different families and genera of this order are described. They include incomplete molting, the presence of a mucous film on the body surface, protection of the head from frontal damage, large lateral projections on the valves and additional sculpture on the head and valves, well-developed lateral armature of the postabdomen, the presence of projections on the postabdomen preanal margin, specialization of setae and spines of the second antenna, especially large exopodites on limbs IV and V, a reduced compound eye, and a large, well-developed ocellus. Similar adaptations to the benthic mode of life in the Anomopoda and Ostracoda are discussed.     

Experiences and attitudes towards dependence among students in Vukovar, Croatia

The sample consisted of 104 students (male and female) from Croatian town of Vukovar, which was heavily destroyed during the Homeland war in 1991. Through self-report, author researches their experiences and attitudes towards different dependences, from so-called light to so-called heavy dependences. The conclusions of the survey are as following: a) experiences of Vukovar' students differ a lot depending on potential danger of particular dependence, so students prefer so-called light dependences than so-called heavy dependences b) Vukovar' students attitudes towards noxiousness/ usefulness also differ according the potential danger of particular dependence and, again, so-called light dependences are more acceptable for students. c) Students have the same attitudes and experiences towards dependences, apart from gender or educational program. Suggesting the practical preventive usefulness, author stresses that basic conclusion of the survey is that students have positive attitude towards so-called light dependences, which should be changed by better information and education of students.     

Morpho-functional grounds of mode of life of Cladocera. V. Morphology and adaptive modifications of trunk limbs of anomopoda

Summary The structure of trunk limbs of Cladocera is comparatively reviewed. The structure of thoracic limbs of the primitive Chydorids Saycia cooki and Eurycercus glacialis is described and figured. The structure of leg I is described and figured in detail. The component parts of leg I of Chydoridae are homologized with those of other Anomopoda. The main phylogenetic trends are pointed out for leg I formation: involution of the posterior lobe of the endite, oligomerization of the setae in the homologous groups, especially those of the posterior lobes of the endite, specialization of the external branch of the endite for creeping in littoral forms and its general involution in planktonic forms. In connection with creeping the setae of the external branch of the endite differentiate in length and form, some of setules may enlarge or disappear. Legs II–V are built according to the same pattern but considerably differ in form and function in Cladocera, which are related to Conchostraca, possessing little differing appendages of different segments. The structure and the chaetal system of the homologous parts of legs II–VI of Anomopoda are analysed. Phylogenetic trends towards specialization and oligomerization are pointed out. The setae tend to form functional groups (e.g. in legs I, III). Thoracic legs II–VI present an example of a plasticity of a metameric organ built by the same pattern. These legs lost a locomotory function and perform other tasks. Setae of Cladocera have rather high formative possibilities. The examples of extreme changes of form are pointed out.     

Effect of calcium on adrenocortical secretion

The authors investigated the effect of calcium on the basal cortisol secretion and on the adrenocortical secretory reserve, the indicator of which is the so-called ACTH stimulation test. They revealed that during acute hypercalcaemia there is a significant increase of the basal cortisol secretion. The adrenocortical secretory reserve is significantly reduced during hypercalcaemia, this being manifested by a smaller rise of cortisolaemia after ACTH. The Synacthen test repeated after a 48-hour interval is fully reproducible under normocalcaemia. The basal values and the response to Synacthen do not differ from the control examination. The stimulatory action of calcium can be explained indirectly via other humoral factors, the production of which is enhanced by calcium but also the function of calcium which acts as the so-called "second messenger" in steroidogenesis. Calcium has, however, probably also a direct corticotropic effect, resembling that of ACTH.     

The taxonomic position of Asian Holopedium (Crustacea: Cladocera) confirmed by morphological and genetic analyses

Limnology - The cladoceran Holopedium gibberum Zaddach, 1855 (Ctenopoda: Holopediidae) was once thought to occur broadly in the northern hemisphere, but its cryptic sister species was recently...