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     

A special moult after the release of the embryo from the brood pouch of Anomopoda (Branchiopoda,Crustacea): a return to an old question

In the past there were doubts about the existence of a especialmoult after the release of an anomopod embryo from the maternalbrood pouch. This moult occurs in all studied anomopods and marksthe beginning of the first juvenile instar. The term neonata (orneanate) must be applied to the unexpanded animal (the terminalphase of embryogenesis) before this moult, and not to the earlyphase of the first postembryonic instar. The latter is the nextstage of development. Using knowledge of the presence of thismoult, it is possible to understand and explain the contradictionsin the periodization of postembryonic development as described bydifferent authors. Under normal conditions males of bosminids andevolutionarily advanced chydorids have only two juvenile instars(with specific morphology) in the course of development. This is anadditional argument to support the idea of close relationship ofthe families Chydoridae and Bosminidae.     

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.     

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.     

Assessment of the importance of fish predation versus copepod predation on life history traits of Daphnia hyalina

Daphnia hyalina is a cladoceran present throughthe whole year except for late summer in Maranhão,a meso-eutrophic reservoir in central Portugal. Apartfrom the influence of food, both vertebrate andinvertebrate predation pressures seem to have aneffect on D. hyalina population dynamics.Enclosure experiments were designed to assess therelative importance of both types of predation. Afterthe summer crash, D. hyalina reached highernumbers in the fishless enclosures than in the lakedespite of high predation pressure upon juveniles byAcanthocyclops robustus. Fish predation upon thelargest individuals, especially large egg bearingfemales, was responsible for the lower fertility ofthe open water population when compared with theenclosure population. In the enclosures an increase intail spine length was observed. The longer tail spineprobably offered protection from copepod predation,allowing at least some of the juveniles to coexistwith their potential predator and reach the adultstage, less susceptible to copepod predation.     

A phylogenetic analysis of the Conchostraca and Cladocera (Crustacea, Branchiopoda, Diplostraca)

A computer-assisted cladistic analysis on morphological characters of the Diplostraca (Conchostraca and Cladocera) has been undertaken for the first time. The morphological information has been obtained from literature and transformed into 56 suitable characters. The analysis included 47 ingroup taxa, comprising five conchostracan taxa (four families of the Spinicaudata and the Laevicaudata) and 42 genera of the Cladocera. A detailed character discussion is presented which will be a useful working base for future phylogenetic studies on the group. A number of systematic groups were, with differing degrees of certainty, supported in all 218 equally short trees. These are the Diplostraca, Cladocera, Gymnomera (Onychopoda and Haplopoda), Onychopoda, Podonidae, Cercopagididae, Anomopoda, Daphniidae, Moininae, Scapholeberinae, Chydoridae, Chydorinae and Sididae. The Spinicaudata were only supported on some of the 218 equally short trees while no support was found for the Conchostraca. Two taxa—the Macrothricidae and Aloninae—were relatively strongly indicated to be paraphyletic. A suggested classificatory hierarchy, without indication of absolute rank, is presented.     

Corrections and additions to the Dumontiidae Santos-Flores &; Dodson, 2003 (Crustacea: Branchiopoda: Anomopoda), and implications for anomopod phylogeny

The presumed sixth limb of Dumontia Santos-Flores & Dodson, 2003 is shown to correspond to the inner part of the fifth limb. Comparison of the latter structure within the order increases the similarity between the Dumontiidae and the Daphniidae (plus Moinidae), but important similarities with the Radopoda remain. The relationship between a ‘‘daphniid line’’ and radopods needs further attention. Handling editor: K. Martens     

Genetic variation in Simocephalus (Anomopoda: Daphniidae) in North America: patterns and consequences

The presumption of intercontinental distributions of many species of Anomopoda has been reinforced by their considerable morphological uniformity and ease of passive dispersal via ephippial eggs. To test the validity of this dogma, genetic variation among taxa in the cladoceran genus Simocephalus was examined on a continental scale. Genetic variability (percentage polymorphic loci = 8.8%, individual heterozygosity = 3.4%) was comparable to that determined on a local scale in Simocephalus and somewhat less than for other zooplankton groups. Four species complexes were distinguished allozymically with unique allelic substitutions found for at least two loci between these taxa. Eight species in North America were differentiated within these complexes (Nei s genetic distance, D>0.30), all of which were clearly separated from two European species. Both S. cf. vetulus and S. cf. serrulatus are broadly distributed in North America, and intraspecific divergence is relatively low. In contrast, S. cf. congener and S. cf. exspinosus represent species complexes, both consisting of several species, differentiated allozymically and morphologically. The concept of generalist species with moderate genetic variation throughout a broad range and specialist species, genetically depauperate, with restricted distributions, is explored.     

Mating behaviour in Moina brachiata (Jurine, 1820) (Crustacea,Anomopoda)

Information on mating behaviour in Anomopoda isavailable for very few species only, though matelocation and recognition certainly play an importantrole in maintaining reproductive isolation betweenspecies. Ephippial females and males of Moinabrachiata were observed in a drop of water under amicroscope for 10–20 minutes. Different combinationsof specimens were used, but copulation was onlyobserved when two males and two ephippial females wereplaced together. Males were very active, and oftentried to grasp a female, whereas females usually madeattempts to escape during the entire period of mating.Three phases were recognized: capture, positioning andcopulation. The male captured the female from thedorsal side, then moved to the ventral side and tooka position with its length axis being perpendicular tothat of the female, forming a sort of cross.Thereafter the pair started rotating around the lengthaxis of the female, while the male pushed thepostabdomen into the females brood pouch. Copulationlasted from 16 to 25 seconds. When different kinds offemales were used, males showed preference forephippial females with an empty ephippium and enlargedovaries. Our results indicate that not only visual andtactile cues may be important in identifying speciesidentity and receptivity of the female, but alsochemical signals.