Morphological Variation in Daphnia Cucullata Sars with Progressive Eutrophication of a Polymictic Lowland Reservoir

Morphological variation in Daphnia cucullata Sars was studied from 1994 to 2001 when fast increasing water trophy of the Siemianówka Reservoir was related to high TP and chlorophyll concentrations. During this period, D. cucullata biomass, as well as its contribution to the total cladocerans biomass clearly decreased. We found that among the six morphological parameters analyzed in D. cucullata, body length, height of the head and length of the tail spine had a tendency to decrease. The maximum values of these traits and carapace length also decreased. The morphology of D. cucullata appeared to be influenced by factors such as food availability, fish predation and an exclusive presence of D. cucullata among Daphniidae species. Increased water mixing in the reservoir may have also affected the changes in the head inclination of D. cucullata.     

Cyclomorphosis as a factor explaining success of a Daphnia hybrid in Tjeukemeer

Recent studies have shown that Daphnia hybrids are intermediate to their parental species for several life history traits, and that they combine advantageous traits of the parental species, thereby decreasing their risk of being preyed upon by planktivorous fish. In this study D. galeata, D. cucullata and their interspecific hybrid are compared with respect to cyclomorphosis and juvenile growth as a potential mechanism to avoid invertebrate predation. Helmets and tail spines are known to reduce predation risk, whereas fast juvenile growth reduces the period that Daphnia are vulnerable to predation. In laboratory cultures, I tested the hypothesis that, with regard to cyclomorphosis, juvenile hybrids are more comparable to the juveniles of D. cucullata, and with respect to growth rate, they are more comparable to D. galeata. The hybrid did not differ significantly from D. galeata in the time needed to reach 0.9 mm body length. In contrast, the relative spine and helmet length of juvenile hybrids were more comparable with those of D. cucullata, whereas in adult instars these characters were more similar to D. galeata. Thus during the first juvenile instars, the hybrids combine the relative long helmet and spine of D. cucullata, with the faster growth of D. galeata. This combination of characteristics might reduce hybrid vulnerability to invertebrate predation in field situations.     

Crowding‐induced changes in growth,reproduction and morphology of Daphnia

• 1 Daphnia may reach high population densities seasonally, or in patches, in lakes. To test the effects of chemicals released by high daphniid densities on their life‐history traits, nine species of Daphnia, D. magna, D. pulicaria, D. pulex, D. hyalina, D. galeata, D. laevis, D. lumholtzi, D. ambigua and D. cucullata, were grown in water from crowded Daphnia cultures in a flow‐through system in the presence of abundant food.
• 2 Water from Daphnia at 85 L^‐1 depressed growth rate, and lowered body size and clutch at first reproduction of six species of small‐bodied Daphnia (adult body length < 1.8 mm), but had no significant effects on larger species. Two clones of D. pulex differed in their growth rate in response to crowding, indicating that response patterns may vary within species.
• 3 Chemicals released by crowded D. magna reduced tail spine length in D. lumholtzi and D. cucullata by 37% and 11%, respectively, and induced changes in carapace morphology in D. lumholtzi and D. ambigua.
• 4 Chemicals released by crowded conspecifics may provide an additional, density‐dependent mechanism of population regulation; when large species of Daphnia coexist at a high population density with small species, these chemicals may reinforce the competitive advantage of large species.

     

Four different head shapes in Daphnia hyalina (Leydig) induced by the presence of larvae of Chaoborus flavicans (Meigen)

In many species of Daphnia spines, neck teeth, and enlarged or reshaped helmets are well-known as defences against invertebrate predators. Until now, Daphnia hyalina (Leydig, 1860), a common species in many European lakes, has appeared to be an exception to this rule.Here, we provide evidence that the larvae of Chaoborus flavicans (Meigen) also can induce morphological changes in D. hyalina. Specimens react morphologically to the presence of larvae in three ways: (a) by changes in the shape of head shield, (b) by the formation of spine(s) on the head, and (c) by the elongation of the tail spine. The frequency and intensity of these morphological changes are correlated positively with midge larvae densities. The most pronounced reactions occurred in young D. hyalina.The predator kairomone also induced changes in Daphnia body size.     

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.     

Competition in natural populations of Daphnia

I investigated the competitive relationships between two species of Daphnia, D. galeata and D. cucullata, and their interspecific hybrid. The term hemispecific competition was introduced to describe competition between parental species and hybrids. In eutrophic Tjeukemeer both parental species were found to compete with the hybrid, whereas competition between D. galeata and D. cucullata seemed limited. Although the effect of competition on life history traits of daphnids may be profound, the influence of the competitors on the seasonal dynamics of the Daphnia species seems limited.     

Extreme cyclomorphosis in Daphnia lumholtzi

1. Daphnia lumholtzi, not previously reported in North America, was found in a small reservoir in East Texas in January, 1991, This species possesses extremely long spines and large fornices; an allometric study was performed to detect any temporal differences in specific growth rates of the spines relative to the body. 2. In nature, mature females attained 1.8mm body length, excluding spines, but when the head and tail spines are included, the total length reached a maximum of 5.6mm. 3. Differences in the growth patterns of the head spine and the tail spine relative to the body existed for D. lumholtzi from January to March 1991. Both the head and the tail spines grew at a faster rate than the body during all 3 months although the rates varied between them. The results contradict the invertebrate predation hypothesis (Dodson, 1974) in that D. lumholtzi's head and tail spines continue to grow during adulthood instead of stopping after the juvenile instars. 4. The head spines grew at a constant allometric rate over time while the tail spine grew faster as the temperature increased. Both varied significantly in length over the 3 months, with animals having the shortest spines in February and the longest in March.     

Cyclomorphosis in Daphnia cucullata: morphometric and population genetic analyses

The cyclomorphic change of head and tailspine of Daphnia cucullatahas been analysed morphometrically and compared with temporalanalyses of enzyme variability. During June, round-headed animalsare rapidly replaced by specimens with high helmets; the rateof increase of the relative helmet length is more rapid thanthe rate of decrease later in the year. The relative sizes ofhelmet and tail spine are not always coupled. The tailspinelength remains approximately the same year round and grows withnegative allometry, so that small animals have the relativelylongest spines. In contrast, the helmet grows in a positivelyallometric fashion and exhibits high seasonal variation. Helmetgrowth is strongly correlated with water temperature but notwith food abundance. The morphometric results are consistentwith the hypothesis that helmet formation is a response to fishpredation while the tail spine provides protection against invertebratepredators. There is no indication of changes in the gene orgenotype frequencies at two loci during the period of rapidmorphological change. Thus cyclomorphosis apparently appearsin the absence of clonal replacement.     

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.     

Formation of morphological defences in response to YOY perch and invertebrate predation in two <Emphasis Type="Italic">Daphnia</Emphasis> species coexisting in a mesotrophic lake

This study examined the formation of morphological defences by two coexisting Daphnia species, the large-sized D. pulicaria (2 mm) and the small-sized D. mendotae (1.4 mm), in response to the presence of young-of-the-year (YOY) yellow perch (Perca flavescens) and invertebrate predators (Chaoborus, Leptodora) during summer in a mesotrophic lake. We hypothesized that due to differential size-selective predation risk by YOY fish and invertebrates, the large-sized and the small-sized Daphnia species would show different morphological responses to predation threats. We followed changes in two morphological traits (relative length of the tail spine in D. pulicaria and of the helmet in D. mendotae) among different periods during summer according to YOY fish and invertebrate predation. We defined four YOY fish predation periods based on the presence of YOY perch in the pelagic zone of the lake and the relative abundance of Daphnia preys in their gut contents, and two invertebrate predation periods based on exclusive or mutual occurrence of the invertebrate predators. The large-sized (D. pulicaria) and the small-sized (D. mendotae) species showed different morphological responses to YOY fish and invertebrate predators, respectively. The tail spine ratio of the juveniles and adults of D. pulicaria did not change in response to YOY fish predation or to invertebrate predation. A gradual increase in the helmet ratio was observed in the small-sized D. mendotae over the summer period. This change was related to the co-occurrence of the invertebrate predators (Chaoborus and Leptodora) and to YOY fish predation. The warmer temperature cannot be accounted for helmet elongation since it was constant across depths, and not related with the co-occurrence of D. mendotae and YOY perch. Guest editor: Piet Spaak Cladocera: Proceedings of the 7th International Symposium on Cladocera     

Breeding biology of Tenagomysis tasmaniae Fenton,Anisomysis mixta australis (Zimmer) and Paramesopodopsis rufa Fenton from south-eastern Tasmania (Crustacea: Mysidacea)

The number, size and developmental stage of young in the brood pouch of female Tenagomysis tasmaniae, Anisomysis mixta australis and Paramesopodopsis rufa was recorded throughout the year. Breeding was intensive from spring till the end of autumn for the three species. Calculation of the egg ratio for each species showed that their major reproductive peaks occurred during spring and summer. A winter depression in the breeding cycle was observed for T. tasmaniae and P. rufa, but A. mixta australis ceased breeding during winter. Seasonal variation in the length of gravid females and number of young carried was evident for these three species. Females were longer in spring and summer and carried more young than in autumn and winter. A linear relationship between female length and brood size was demonstrated for each species; annual and seasonal equations were calculated for females carrying each developmental stage. The seasonal equations showed that for a female of given length fecundity was greater during spring than any other season. Natality was estimated to be highest during late spring, summer and early autumn for the three species. No seasonal variation in the size of eggs was evident for the three species. The reproduction pattern of T. tasmaniae, A. mixta australis and P. rufa appears to be very similar to that reported for the majority of iteroparous coastal temperate mysids throughout the world.     

Zooplankton-induced unicell-colony transformation in Scenedesmus acutus and its effect on growth of herbivore Daphnia

The effects of colonial ecomorphs of the green alga Scenedesmus acutus on growth of Daphnia cucullata and D. pulex were examined. In ecologically relevant densities (up to 200 animals l^-1) the relatively small D. cucullata did not induce colony formation in Scenedesmus acutus, whereas the larger congener D. pulex significantly promoted colony formation. Both clearance rate and population growth rate (r) were significantly lower in D. cucullata when fed colonial Scenedesmus then when fed unicellular food. However, for D. pulex no effects of food type were observed. These results show that large Daphnia may influence its food in such a way that smaller congeners and competitors are negatively affected.     

Differences in genetic structure and ecological diversity between parental forms and hybrids in a Daphnia species complex

Hybridization is a common phenomenon in Daphnia species complexes. Hybrids often dominate in Daphnia populations; therefore it is worthwhile to look for principal differences between parental and hybrid populations with respect to their genetic structure and clonal differentiation. We studied natural populations of members of the Daphnia galeata/hyalina/cucullata complex in three lakes. In one of these lakes, one parental species (D. galeata) and one hybrid (D. galeata × cucullata) were investigated more intensively. The frequency of sexual reproduction was higher in parental populations, whereas clonal diversity was higher in hybrid populations. Ecological differentiation among clonal groups was more pronounced in the D. galeata × cucullata hybrid compared to D. galeata, whereas selection intensity was weaker. These results are discussed with respect to stability of clonal groups, multiple hybridizations and selective constraints.     

Anti-predator Defence in Three Daphnia Species

Over a two-year period, each of the three Daphnia species in two lakes with different pressure by fish and invertebrate predators exhibited different defensive reactions: predator avoidance in space (Daphnia hyalina), seasonal morphological changes (Daphnia cucullata), and a combination of both (Daphnia cristata). Body size and general susceptibility to predation, combined with predators' preferences were most likely responsible for the kind of defence in each of the three Daphnia.     

Impact of fish predation on coexisting Daphnia taxa: a partial test of the temporal hybrid superiority hypothesis

Fish predation was tested as a factor mediating the coexistence of Daphnia taxa in the shallow, hypertrophic Lake Blankaart. Naturally co-occurring populations of D. galeata and the hybrid D. galeata x cucullata were subjected to different levels of fish predation in in situ enclosures. In control enclosures without fish, the largest taxon D. galeata rapidly became dominant over the intermediate sized D. galeata x cucullata, mainly as a result of higher birth rates. In enclosures with fish, population densities of D. galeata dropped relative to D. galeata x cucullata, due to higher mortality rates. These results are in concordance with the `temporal hybrid superiority hypothesis', and can be explained by a higher vulnerability of the large and more conspicuous D. galeata to the size selective predation exerted by visually hunting planktivorous fishes. After approximately one month, however, population growth rates of D. galeata and D. galeata x cucullata in the enclosures with fish converged, due to a relative reduction in the mortality rate of D. galeata. This suggests that, in the presence of fish, D. galeata may co-exist with hybrids due to a decrease in its relative vulnerability to visual predation with time. Indeed, both D. galeata and the hybrid showed strong reductions in adult body size in the enclosures with fish, but this size reduction tended to be stronger in D. galeata than in D. galeata x cucullata. In addition, turbidity increased in the enclosures with fish and may additionally have reduced the relative advantage of D. galeata x cucullata with regard to mortality caused by visual predation.     

A new sparid, Acanthopagrus akazakii, from New Caledonia with notes on nominal species of Acanthopagrus

A new sparid species, Acanthopagrus akazakii, is described on the basis of 12 types collected from Noumèa, New Caledonia. Acanthopagrus akazakii is most similar to Acanthopagrus berda in overall appearance but differs from the latter in having 4¹/₂ scale rows between the 5th dorsal fin spine base and lateral line, anteriormost head scales broadly rounded without small scales anteriorly, upward- and downward-oriented portions present on anteriormost part of upper lip, a downward-oriented portion on anteriormost part of lower lip in specimens over ca. 160 mm in standard length, and upper head profile gently convex from snout tip to above eye throughout growth (vs. 3¹/₂ scale rows between 5th dorsal fin spine base and lateral line, anteriormost head scales rounded with small scales anteriorly, no upward and downward portions in anteriormost upper and lower lips, and upper head profile becoming concave from snout tip to above eye with growth). Furthermore, A. berda develops a strong concavity of the ventral edge of the first two infraorbitals above the posterior part of upper jaw with growth, whereas A. akazakii has a generally straight series throughout growth. Nominal species in A. berda are reviewed, with notes on nominal species of Acanthopagrus.     

Evolutionary stasis despite selection on a heritable trait in an invasive zooplankton

Invasive species are one of the greatest threats to ecosystems, and there is evidence that evolution plays an important role in the success or failure of invasions. Yet, few studies have measured natural selection and evolutionary responses to selection in invasive species, particularly invasive animals. We quantified the strength of natural selection on the defensive morphology (distal spine) of an invasive zooplankton, Bythotrephes longimanus, in Lake Michigan across multiple months during three growing seasons. We used multiple lines of evidence, including historic and contemporary wild‐captured individuals and palaeoecology of retrieved spines, to assess phenotypic change in distal spine length since invasion. We found evidence of temporally variable selection, with selection for decreased distal spine length early in the growing season and selection for increased distal spine length later in the season. This trend in natural selection is consistent with seasonal changes in the relative strength of non‐gape‐limited and gape‐limited fish predation. Yet, despite net selection for increased distal spine length and a known genetic basis for distal spine length, we observed little evidence of an evolutionary response to selection. Multiple factors likely limit an evolutionary response to selection, including genetic correlations, trade‐offs between components of fitness, and phenotypic plasticity.     

Coexistence in a sea urchin guild and its implications to coral reef diversity and degradation

Summary Coexistence between the coral reef inhabiting sea urchins Echinometra mathaei, Diadema savignyi and D. setosum was studied by comparing differences in body morphology, distribution, diet, susceptibility to predators, intra- and interspecific competition and settlement. The three species share similar diets and broad within-habitat distributions but differ in their microspatial preferences. E. mathaei is the smallest species, has the highest settlement rates and lives territorially within small burrows or crevices. D. savignyi is intermediate in size and lives frequently in intermediate size crevices or occassionally in social groups. D. setosum is the largest species and occassionally lives in large crevices or more frequently in social groups. Both Diadema have similarily low settlement rates. Competition experiments showed that E. mathaei was consistently the top competitor for crevice space. Diadema species shared larger crevices but competition occured within smaller crevices and was frequently won by the largest individual, regardless of species. D. savignyi may be the top competitor for crevice space between the Diadema species due to a reduced spine length/test size ratio which gives it a larger test for the same crevice size requirement. Predation rates were high for E. mathaei and low for both Diadema species. Coexistence is mediated by predation on the competitive-dominant while predation coupled with different body morphologies and behavior allows spatial resource partitioning of the reef's variable topography. Consequently, the three variables of predation, topographic complexity and differing body shapes create the observed species diversity. A reduction in predators due to stochastic fluctuations or from fishing pressure can lead to E. mathaei population increases and competitive exclusion of Diadema.     

Zooplankton dynamic of Lake Prespa (Macedonia)

The composition and temporal abundance patterns of zooplankton community in Lake Prespa pelagic zone were studied throughout an annual cycle (October 2008–September 2009). Eighteen species (10 Rotifera, 5 Cladocera, 2 Copepoda, 1 Mollusca) were evidenced. It includes Arctodiaptomus steindachneri, an endemic calanoid of the Western Balkans. The zooplankton density varied during the investigated period and certain seasonal successions in temporal transpositions of their maximal development were noticed. The total zooplankton density was much higher in spring and summer period. The representatives of subclass Copepoda were dominant during the whole year. The dominant species was A. steindachneri. At the second place (with exception of the winter period) were the Cladocera representatives. The dominant species was Daphnia cucullata. At the third and fourth place were rotifers and larval form of Dreissena polymorpha, respectively. The quantitative presence of the Copepoda representatives was significantly lower in terms of the previous investigations. The Cladocera quantity reached high values during this period, particularly in summer. Such altering in the quantitative ratio between Copepoda and Cladocera in favor of the Cladocera, especially high quantitative presence of D. cucullata, typical for eutrophic lakes, imply of significant changes in zooplankton community and changes of the water quality in Lake Prespa pelagial.     

Postsettlement growth of two estuarine crab species, Chasmagnathus granulata and Cyrtograpsus angulatus (Crustacea, Decapoda, Grapsidae): laboratory and field investigations

The estuarine grapsid crabs Chasmagnathus granulata and Cyrtograpsus angulatus belong to the most typical and dominant inhabitants of brackish coastal lagoons in southeastern South America. In a combined laboratory and field investigation of juvenile growth, we measured the increase in body size in these species under controlled conditions as well as in field experiments (in Mar Chiquita lagoon, Argentina), seasonal changes in size frequency distribution of a natural population, and growth related changes in selected morphometric traits of male and female juveniles (relations between carapace width, carapace length, propodus height and length of the cheliped, and pleon width). At 24°C, Cy. angulatus grew faster than Ch. granulata; it reached the crab-9 instar (C9; 13 mm carapace width) after 92 days, while Ch. granulata required 107 days to reach the C8 instar (7.4 mm). At 12°C, growth ceased in both species. The pleon begins to show sexual differences in the C5 (Cy. angulatus) and C8 instar (Ch. granulata), respectively, while the chelae differentiate earlier in Ch. granulata than in Cy. angulatus (in C4 vs C6). In the field, growth was maximal in summer, and was generally faster than in laboratory cultures. However, there is great individual variability in size (about 25% even in the first crab instar) and in size increments at ecdysis, increasing throughout juvenile growth. Our data indicate that, in the field, small-scale and short-term variations in feeding conditions, temperature, and salinity account for an extremely high degree of variability in the absolute and relative rates of growth as well as in the time to sexual differentiation. Received in revised form: 20 September 2001 Electronic Publication