Introgression as a Factor in the Evolution of Polytypical Plankton Cladocera

The genera Bosmina and Daphnia offer considerable taxonomical difficulties due to the existence of a very great number of different races. An attempt has been made to understand this phenomenon mainly as the result of introgressive processes.     

Naturally occurring interspecific hybridization inSimocephalus (Cladocera,Daphniidae): its potential significance

Among the Cladocera, the occurrence of interspecific hybrids in nature has been reported frequently but rarely adequately documented. During the course of a survey of the genetic variation and population structure of several species ofSimocephalus in ponds in southern Ontario, Canada, readily distinguishable congeners often co-occurred in a pond. Occasionally, clones were recovered that were intermediate in phenotype (as determined from both morphological and electrophoretic analyses) between two species. Two different hybrid clones have been examined in detail to determine the parentals involved in the crosses. Evidence from field and laboratory studies is presented to document the hybrid nature of these clones. Factors favouring formation and long-term survival of hybrids in littoral zone and pelagic species are discussed.     

Population dynamics and production of Daphnia hyalina Leydig and Daphnia cucullata Sars in Tjeukemeer

As part of a research programme on the food chains in Tjeukemeer, the Daphnia hyalina and Daphnia cucullata populations were studied for three successive years. To analyse the factors regulating the production of these two species, their population parameters (density, size distribution, fecundity) and population dynamics (birth rate, mortality rate) were studied and related to environmental factors. Since Daphnia in Tjeukemeer shows continuous recruitment, the population dynamics model INSTAR was developed and used to integrate field data with laboratory data on development rates and length-weight relationships. The dynamics of the Daphnia species are mainly regulated by temperature and fish predation, the latter affects both birth rate and mortality. Total annual Daphnia production was 3.1–6.9 g org. dry wt M^–2, and annual P/B ratio ranged from 25 to 40 for D. cucullata and from 45 to 49 for D. hyalina.     

Interspecific hybridization betweenDaphnia hyalina,D. galeata,andD. cucullata and seasonal abundances of these species and their hybrids

The three cladoceran speciesDaphnia hyalina, D. galeata, andD. cucullata frequently coexist in the lakes of northern Germany. Although there are some problems in distinguishing them morphologically, they are easily determined by gelelectrophoresis: each species carries a different allele at the glutamate oxaloacetate transaminase (GOT) locus. Animals morphologically intermediate between two species are heterozygous for the alleles carried by the species they resemble. This pattern is in agreement with the findings at other loci, where also diagnostic alleles exist. These findings are most easily explained by interspecific hybridization between the three species. No evidence is found for backcrosses involving hybrids ofD. cucullata, whereas some backcrosses betweenD. hyalina, D. galeata, and their hybrids are found in some lakes. In four lakes the seasonal abundances of the three species and their hybrids are determined.     

The Bosmina kessleri-like morphotype of Eubosmina in Lake Muskoka,Ontario, Canada,as putative interspecific hybrids

Like Bythotrephes, Bosmina (Eubosmina) coregoni coregoni Baird appears to have invaded North America from Eurasia, since there are no records from before 1965. All previously known North American populations have a well-rounded ventrocaudal corner of the carapace, suggesting that the founding race is one of the rotunda type, the most common eubosminid in European lakes. In Lake Muskoka, immediately to the east of Georgian Bay, Ontario, eubosminids were found that shared features of B. (E.) longicornis kessleri Ulianine. In contrast to other North America populations of B.(E. )c. coregoni, these specimens have a short, broadly based mucro. Morphological features suggest that these are interspecific F₁-hybrids between B. (E.) longispina Leydig and B. (E.) coregoni, without any signs of introgression.This paper is dedicated to the memory of Dr. John C.H. Carter.     

Notes on Cladocera and Copepoda from high altitude lakes in the Mount Everest Region (Nepal)

The paper deals with Cladocera and Copepoda species from eight high altitude lakes in the Khumbu area. In all lakes, an endemic known diaptomid of the palearctic genus Arctodiaptomus is found. The dark and large Daphnia tibetana occurs in five of the lakes sampled, characterized by the persistence of the spine in adults (parthenogenetic and ephippial females, males). Apparently, this is an unknown feature of this species. The literature on the diagnostic traits of the different morphs described is reviewed. In addition, a transparent and smaller-sized Daphnia species occurs in two lakes. This is a D. longispina characterized by the absence in adults of the carapace spine (var. aspina Weretschagin, 1911). The presence of these two species is discussed in relation to water transparency, colour, and vertical distribution. Two hypotheses on the evolution of cuticular pigmentation in Daphnia are examined. In addition to these mostly dominant species, a macrothricid also typical of high altitude lakes in the Alps was found (Macrothrix hirsuticornis) together with two cosmopolitan Chydoridae.     

Experimental interspecific hybridization in Daphnia

Hybridization is a common phenomenon in Daphnia (Cladocera; Anomopoda); interspecific hybrids have been found between several species and hybrids are found in many European lakes. Although much information on the morphology, ecology and genetics of hybrids is available, little is known about the level of reproductive isolation among species or about the relative fitness of hybrids and parental species. In order to facilitate studies on differentiation and speciation processes and comparative experimental studies on hybrids and recombinant genotypes, we present the first successful laboratory crossing experiments of two different Daphnia species, D. galeata and D. cucullata. Males and sexual females from two D. galeata and two D. cucullata clones were reciprocally crossed, juveniles hatched from resting eggs and reared until maturity. Hatching and juvenile survival rates of hybrids were relatively low (12.1% and 24%, respectively). D. galeata and D. cucullata clones vary in their level of successful interspecific matings and in the number of subsequent offspring. In general, hybrid crosses between D. cucullata females and D. galeata males were more successful than reciprocal crosses.     

The systematics of Australian Daphnia (Cladocera: Daphniidae). Multivariate morphometrics

Forty nine characters were measured in each of 76 male and 200 female Daphnia including specimens from all eleven taxa described from the genus in Australia, except D. jollyi. Separate multivariate analyses (cluster, principal components and discriminant) of males and females each revealed only five morphological groups in the genus corresponding to the species D. occidentalis, D. lumholtzi, D. cephalata, D. nivalis, and a conglomerate of all other taxa described from the D. carinata complex, D. carinata (sensu lato). Interpretation of the male results was straightforward, but that for females was not because of the occurrence of an environmentally induced seasonal change in morphology (cyclomorphosis) in females. Log transformations of the female data successfully isolated the effects of cyclomorphosis. The seasonal morphs formed two groups which were further differentiated into taxonomic groups.     

The systematics of Australian Daphnia (Cladocera: Daphniidae). Electrophoretic analyses of the Daphnia carinata complex

Genetic variation at eleven allozyme loci was scored in approximately 1600 individual Daphnia. The samples included representatives of all nine taxa described from the D. carinata complex in Australia. Principal coordinates and cluster analyses revealed only three groups of genotypes within the complex, corresponding to the taxa D. cephalata, D. nivalis, and a conglomerate of all other taxa described from the complex, D. carinata (sensu lato). These results are consistent with recent multivariate morphological analyses of the complex.     

Egg predation by copepods in Daphnia brood cavities

Both field observations and enclosure experiments show that juvenile copepods enter Daphnia brood cavities to feed on Daphnia eggs and embryos. The ability to perform such in vivo exploitation is reported for both cyclopoid and calanoid copepods. Copepodites of Acanthocyclops robustus (G.O. Sars) were found to eat eggs in brood cavities of D. magna, D. pulex and D. pulicaria in experimental enclosures rich in algae. Copepodites of Eudiaptomus gracilus (G.O. Sars) were found in brood cavities of D. hyalina in a mesotrophic lake. The copepods' intrusions into brood cavities caused dramatic declines in the clutch size of infested Daphnia, and this predation effect could easily be confused with the effect of severe food limitation.     

A note about Daphnia in Venezuelan waterbodies

After examining numerous water samples of 19 lakes and reservoirs in Venezuela, only two species of Daphnia were found: D. laevis (in colder, high altitude lakes) and D. gessneri. This observation supports the theory of the scarcity of Daphnia species in tropical inland waters.     

Introgressive hybridization between Galaxias depressiceps and Galaxias sp D (Teleostei: Galaxiidae) in Otago, New Zealand: Secondary contact mediated by water races

Central Otago (New Zealand) was the focus of an intense period ofgold mining in the late 19th century. Mining relied on high waterpressure for sluicing, often requiring extensive systems of water raceslinking headwaters over large distances. On Rough Ridge, races connectedheadwaters of Otago's two major river systems, the Taieri and theClutha. We employed genetic analysis of freshwater fish(Galaxias) populations to assess the biological implications ofrace-development. The two galaxiid taxa found on Rough Ridge (G. spD and G. depressiceps) are reciprocally monophyletic formtDNA and have three fixed allozyme differences. Rough Ridge aside,G. sp D is apparently absent from the Taieri system and G.depressiceps from the Clutha system. Distributions of mitochondrialDNA, allozyme and microsatellite alleles imply that artificialconnections have facilitated bidirectional introgression: movementof G. sp D alleles into the Taieri system and G.depressiceps alleles into the Clutha. The finding of some identicalcontrol region haplotypes (of both lineages) on either side of thecatchment boundary confirms the recency of genetic exchange. Near theinferred region of contact, samples from Linn Burn (Taieri) are pureG. depressiceps for three diagnostic allozymes and mtDNA.Samples from upper Pool Burn (Clutha) are pure G. sp D forthese same four markers. However, other collections from around thecontact zone represent genetic mixtures forming a cline. We fear thatsimilar human-induced mixing may have happened elsewhere in uplandOtago.     

The utility of nuclear introns for investigating hybridization and genetic introgression: a case study involving Brachyramphus murrelets

Interspecific hybridization can have importantconsequences for conservation, but can bedifficult to detect using morphologicalmarkers. To test the utility of nuclear intronsfor investigating hybridization and geneticintrogression, we analyzed variation in fivenuclear introns and the mitochondrialcytochrome b gene in two species ofseabirds that are declining and may behybridizing: marbled murrelets(Brachyramphus marmoratus) and Kittlitz'smurrelets (B. brevirostris). No alleleswere shared between samples of the two species,and intron alleles formed reciprocallymonophyletic groups in gene trees. Our resultssuggest that few murrelets in Alaska areF ₁, F ₂or back-cross hybrids,and that gene pools of these species have beenindependent for 1.8 to 5.7 million years. Weconclude that introns are a potentially richsource of markers for analyzing hybridizationand introgression in endangered or decliningspecies.     

Low fish predation pressure in London reservoirs: II. Consequences to zooplankton community structure

The 1992 survey of zooplankton structure in fourteen London supply reservoirs showed the overall dominance of large-bodied zooplankton, mainly species of Daphnia. These reservoirs can be considered as ‘anti-fish’ by virtue of their steeply sloping concrete or brick sides. The average biomass of large Daphnia spp (retained on a 710 μm sieve) in the total zooplankton biomass was higher than 20% for twelve out of fourteen reservoirs. The cladoceran-copepod ratio was inversely correlated with both dominance of large-bodied Daphnia magna and cladoceran body-size structure. Parallelly, there were tendency of more efficient utilization of lowered algal crops in reservoirs dominated by large-bodied Daphnia spp. A graphical model is presented which relates daphnid species composition and zooplankton size structure to a presumed gradient of fish biomass in these reservoirs.     

The relationship between cladoceran body size and the growth of underyearling roach (Rutilus rutilus) (L.) in two shallow lowland lakes: a mechanism for density-dependent reductions in growth

Large planktonic Cladocera are typically the most important components of the diet of underyearling roach. Selection for large species and individuals by fish can result in a shift in the species composition of the cladoceran community as well as a reduction in the mean size of the individuals of large species and in the assemblage as a whole. Laboratory experiments demonstrated that underyearlings feeding on Daphnia hyalina smaller than 1 mm in length has a significantly lower intake of prey volume per unit time than when feeding on prey greater than 1.5 mm. A decrease in the nutritional quality of zooplankton prey, brought about by increasing predation pressure is suggested as the mechanism for density-dependent reductions in the growth of underyearling roach in eutrophic water bodies.     

Sexual and somatic hybridization in the genusLycopersicon

In recent years, a large number of reports have been published on the recovery of somatic hybrids in the genusLycopersicon and their potential use as a tool in plant breeding programs. Somatic hybridization as a way of enabling the incompatibility barriers which exist within the genusLycopersicon to be bypassed has attracted great interest. WildLycopersicon species harbor numerous interesting agronomic characteristics, which could be transferred to tomato by somatic hybridization. In particular, the production of asymmetric hybrids is explored as an approach to obtain the transfer of only a part of the nuclear genome of wildLycopersicon species. Considerable information is available on the fate of chloroplasts and mitochondria in fusion products inLycopersicon, and unfortunately, cybridization (transfer of chloroplasts and/or mitochondria) seems often difficult to achieve.     

Morphological and molecular evidence for hybridization and introgression in Central European Arctium (Asteraceae)

Three species of burdock (Arctium lappa, A. tomentosum, and A. minus; Asteraceae), which commonly occur in sympatry in Central Europe, are thought to hybridize frequently, hence raising questions about their integrity. We surveyed mixed, monotypic, and/or single plant stands of these species in Central Germany to assess their extent of hybridization and introgression, and also the likelihood of insect host shifts between them. Individual plants were characterized by multivariate analyses of morphological and RAPD variation in combination with RAPD-based character-index-scores. All analyses recognized the three species as distinct units, but also identified a limited number of first (or early) generation hybrids between sympatric A. lappa and either A. tomentosum or A. minus, respectively, as well as one introgressant of A. tomentosum into A. lappa. While pre-zygotic isolation may partly account for the rarity of first generation crosses (F₁'s), the near absence of later generation hybrid segregants or introgressants likely reflects post-zygotic isolation due to the disruption of co-adapted gene complexes and/or habitat-mediated superiority shared by the ruderal parentals. As the tephritid fly Tephritis bardanae oviposits on both A. tomentosum (T) and A. minus (M), but not on A. lappa (L), the occurrence of L × M and L × T hybrids and/or introgressants may have important evolutionary consequences regarding the acquisition of a new host, i.e. A. lappa, by these insects.     

Differences in life-history and aging in two clonal groups of Daphnia cucullata Sars (Crustacea: Cladocera)

Life-history traits were compared in twenty-two clones of Daphnia cucullata from the pre-alpine lake Klostersee in southern Germany. Clones were kept under standardized conditions (18 °C; 16:8 L/D) both singly and with 25 females per glass. Clones belonged to two genotype groups distinguished by enzyme electrophoresis. 11 of the clones had the genotype FM at the phosphate glucose isomerase locus and 11 the genotype MM. Neonates and adults of the age class 10 to 21 days were larger in one group (FM) than in the other group (MM). The size specific growth rates within any age class depended on age and body size and differed between the genotype groups. The fecundity of young FM animals (< 16 days) was significantly higher than that of MM animals, but for older females fecundity did not differ significantly. Senescence started about 6 days earlier in FM than in MM. The data are consistent with results of a field study on Daphnia cucullata in the Klostersee (Jacobs, 1990).     

Zooplankton biomass rarely improves predictions of chlorophyll concentration in Canadian Shield lakes that vary in pH

To determine the frequency with which zooplankton influence chlorophyll a (Chla) levels, we explored annually-averaged data from oligotrophic and mesotrophic lakes that differed in morphometry, total phosphorus (TP) concentrations, and zooplankton community composition due to pH. The data were divided into two sets according to the type of filter used to collect chlorophyll. Residuals of the Chla: TP regressions were not related to lake morphometry, TN content, water clarity or pH. In the first data set there were no consistent relationships between residuals in Chla and twelve grazer biomass variables for 37 of the 38 lakes. The single exception had a very large population of Daphnia dubiaand low concentrations of Chla for its TP. In the second data set, 3 of 25 lakes had exceptionally low Chla concentrations for their TP. These lakes were acidic (pH < 6) and had very large biomasses of Holopedium gibberumcorrelated with negative Chla residuals, indicating significant grazing. At pH > 6, Daphnia spp. strongly influenced the significant correlations. We conclude that zooplankton contribute to the prediction of Chla beyond that possible by TP alone in acidic and non-acidic Canadian Shield Lakes, but evidence for strong suppression of chlorophyll by grazers was relatively rare (4 of 63 cases) on annual time steps.