Levels of polyandry in <Emphasis Type="Italic">Apis laboriosa</Emphasis> Smith from Nepal

Summary Actual and effective number of matings of Apis laboriosa Smith queens were estimated by analysing 40–192 workers or worker pupae of five colonies using microsatellites. Effective paternity frequencies varied from 13.63 to 31.57, with a mean of 20.19 (SD 6.07). Our results demonstrate that the Himalayan Giant honeybee A. laboriosa shows high levels of polyandry, typical of the genus.Received 30 January 2002; revised 24 October 2003; accepted 3 December 2003.     

Genetic variance of mating frequency in the honeybee (Apis mellifera L.)

Summary. The genetic variance of queen mating frequency was studied in honeybees (Apis mellifera carnica). Worker offspring (N = 966) of 28 naturally mated half sister-queens (r = 0.25) from seven unrelated breeding lines were genotyped at four DNA microsatellites. The mating frequencies of the queens were derived from the offspring genotypes. The number of observed matings per queen ranged from 10 to 28 with an average of 17.32 ± 1.10 (number of estimated matings: 24.94 ± 2.51; number of effective matings: 20.09 ± 1.73). Half-sib analyses of the breeding lines were used to estimate heritability. Heritability was h² = 0.449 ± 0.135 for the estimated number of matings and h² = 0.262 ± 0.103 for the number of effective males, which are both significantly different from zero. We conclude that a high genetic variance for polyandry in honeybees can be favored by balanced selection between individual queen and colony level.Received 16 October 2003; revised 4 May 2004; accepted 4 May 2004.     

Thermal investigations of a honey bee colony: thermoregulation of the hive during summer and winter and heat production of members of different bee castes

Summary The temperature at the centre, the periphery and the entrance of a honey bee colony (Apis mellifera carnica) was continuously determined during the summer season and the broodless time in winter. During the summer season the temperature in the brood nest averages 35.5°C with brief excursions up to 37.0°C and down to 33.8°C. Increasing environmental temperatures resulted in linear increases in the temperature of the hive entrance, its periphery and its centre. The temperature in the centre of an overwintering cluster is maintained at an average value of 21.3°C (min 12.0°C, max 33.5°C). With rising ambient temperatures the central temperature of a winter cluster drops whereas the peripheral temperature increases slightly. With decreasing external temperatures the peripheral temperature is lowered by a small amount while the cluster's centre temperature is raised. Linear relationships are observed between the central and the ambient temperature and between the central temperature and the temperature difference of the peripheral and the ambient temperatures. The slopes point to two minimum threshold values for the central (15°C) and the peripheral temperature (5°C) which should not be transgressed in an overwintering cluster. Microcalorimetric determinations of the heat production were performed on the three castes of the honey bee: workers, drones and queens of different ages. Among these groups single adult workers showed the highest heat production rates (209 mW·g^–1) with only neglectible fluctuations in the heat production rate. Juvenile workers exhibited a mean heat production rate of 142 mW·g^–1. The rate of heat production of adult workers is strongly dependent upon the number of bees together in a group. With more than 10 individuals weight-specific heat dissipation remains constant with increasing group sizes at a level approximately 1/17 that of an isolated bee. Differences are seen between the rates of virgin (117 mW·g^–1) and laying (102 mW·g^–1) queens. Laying queens showed less thermal fluctuations than virgin queens. High fluctuations in heat production rates are observed for drones. In both groups (fertile, juvenile) phases of high and extremely low activity succeed one another. The heat production of juvenile drones was 68 mW·g^–1, that of fertile drones 184 mW·g^–1 due to stronger locomotory activities.     

Patriline differences in emergency queen rearing in the honey bee, <Emphasis Type="Italic">Apis mellifera</Emphasis>

Summary In the polyandrous honey bee, Apis mellifera, workers can potentially increase their inclusive fitness by rearing full-sister queens. If the mother queen dies suddenly, workers feed a few larvae in worker cells with royal jelly and rear them into queens (emergency queen rearing). Using DNA microsatellite markers we determined the patriline of emergency queens reared in two colonies headed by naturally-mated queens before being made queenless. We found that some patrilines were reared more than others in one colony, but not in the other. These differences between colonies suggest that selective rearing is not always present and this might explain the mixed results of previous nepotism studies in the honey bee.Received 10 February 2003; revised 7 March 2003; accepted 17 March 2003.     

Experimental Conversion of Colony Social Organization in Fire Ants (<Emphasis Type="Italic">Solenopsis invicta</Emphasis>): Worker Genotype Manipulation in the Absence of Queen Effects

Colony social organization in the fire ant Solenopsis invicta appears to be under strong genetic control. In the invasive USA range, polygyny (multiple queens per colony) is marked by the presence of the Gp-9 ^b allele in most of a colony’s workers, whereas monogyny (single queen per colony) is associated with the exclusive occurrence of the Gp-9 ^B allele. Ross and Keller, Behav Ecol Sociobiol 51:287–295 (2002) experimentally manipulated social organization by cross-fostering queens into colonies of the alternate form, thereby changing adult worker Gp-9 genotype frequencies over time. Although these authors showed that social behavior switched predictably when the frequency of b-bearing adult workers crossed a threshold of 5–10%, the possibility that queen effects caused the conversions could not be excluded entirely. We addressed this problem by fostering polygyne brood into queenright monogyne colonies. All such treatment colonies switched social organization to become polygyne, coincident with their proportions of b-bearing workers exceeding 12%. Our results support the conclusion that polygyny in S. invicta is induced by a minimum frequency of colony workers carrying the b allele, and further confirm that its expression is independent of queen genotype or history, worker genotypes at genes not linked to Gp-9, and colony genetic diversity.     

Production,standing biomass and natural abundance of <Superscript>15</Superscript>N and <Superscript>13</Superscript>C in ectomycorrhizal mycelia collected at different soil depths in two forest types

Nutrient uptake by forest trees is dependent on ectomycorrhizal (EM) mycelia that grow out into the soil from the mycorrhizal root tips. We estimated the production of EM mycelia in root free samples of pure spruce and mixed spruce-oak stands in southern Sweden as mycelia grown into sand-filled mesh bags placed at three different soil depths (0–10, 10–20 and 20–30 cm). The mesh bags were collected after 12 months and we found that 590±70 kg ha^–1 year^–1 of pure mycelia was produced in spruce stands and 420±160 kg ha^–1 year^–1 in mixed stands. The production of EM mycelia in the mesh bags decreased with soil depth in both stand types but tended to be more concentrated in the top soil in the mixed stands compared to the spruce stands. The fungal biomass was also determined in soil samples taken from different depths by using phospholipid fatty acids as markers for fungal biomass. Subsamples were incubated at 20°C for 5 months and the amount of fungal biomass that degraded during the incubation period was used as an estimate of EM fungal biomass. The EM biomass in the soil profile decreased with soil depth and did not differ significantly between the two stand types. The total EM biomass in the pure spruce stands was estimated to be 4.8±0.9×10³ kg ha^–1 and in the mixed stands 5.8±1.1×10³ kg ha^–1 down to 70 cm depth. The biomass and production estimates of EM mycelia suggest a very long turnover time or that necromass has been included in the biomass estimates. The amount of N present in EM mycelia was estimated to be 121 kg N ha^–1 in spruce stands and 187 kg N ha^–1 in mixed stands. The ¹³C value for mycelia in mesh bags was not influenced by soil depth, indicating that the fungi obtained all their carbon from the tree roots. The ¹³C values in mycelia collected from mixed stands were intermediate to values from pure spruce and pure oak stands suggesting that the EM mycelia received carbon from both spruce and oak trees in the mixed stands. The ¹⁵N value for the EM mycelia and the surrounding soil increased with soil depth suggesting that they obtained their entire N from the surrounding soil.     

Virgin queens selectively destroy fully matured queen cells in the honeybee <Emphasis Type="Italic">Apis mellifera</Emphasis> L.

Summary Experiments were carried out to determine how newly emerged virgin queens destroy queen cells with broods which will be competitors for the succession of the colony. When queen cells with older, 13-day-old broods set to emerge within 1 day were presented together with those of younger broods to workers and newly emerged queens in the colony, we found that the older queen cells were preferentially destroyed. It was also shown that a virgin queen destroyed the queen cells with older brood (12–13 days old) first when they were presented together with cells with younger broods (9–10 days old) simultaneously in the experimental cage. However, no significant preference was detected in the destruction between queen cells with 10- and 7-day-old broods. We concluded that virgin queens selectively destroy the queen cells housing broods which will emerge shortly. The possibility that by the selective destruction of older queen cells, newly emerged queens can reduce their risks including death that might otherwise be caused by fights with competitors was discussed.Received 10 June 2003; revised 22 December 2003; accepted 15 January 2004.     

Kin structure and colony male reproduction in the hornet<Emphasis Type="Italic"> Vespa crabro</Emphasis> (Hymenoptera: Vespidae)

We estimated queen mating frequency, genetic relatedness among workers, and worker reproduction in Vespa crabro flavofasciata using microsatellite DNA markers. Of 20 colonies examined, 15 contained queens inseminated by a single male, 3 colonies contained queens inseminated by two males, and 2 colonies contained queens inseminated by three males. The genetic relatedness among workers was estimated to be 0.73±0.003 (mean±SE). For this high relatedness, kin selection theory predicts a potential conflict between queens and workers over male production. To verify whether males are derived from queens or workers, 260 males from 13 colonies were genotyped at four microsatellite loci. We found that all of the males were derived from the queens. This finding was further supported by the fact that only 33 of 2,990 workers dissected had developed ovaries. These workers belonged to 2 of the 20 colonies. There was no relationship between queen mating frequency and worker reproduction, and no workers produced male offspring in any of the colonies. These results suggest that male production dominated by queens in V. crabro flavofasciata is possibly due to worker policing.     

Nitrogen fixation by the woody legumeLeucaena leucocephala in Tanzania

Summary The nitrogen fixation rate in a 4-year-old stand of the woody legumeLeucaena leucocephala (Lam.) de Wit. was estimated in the field at a rather dry site in Tanzania by use of an acetylene reduction technique. The diurnal mean value during April–May was 35 nmol C₂H₄ mg^–1 (dry weight) nodules h^–1, with a variation between 22±8 and 48±12 nmol C₂H₄ mg^–1 (dry weight) nodules h^–1 in early morning and at midday, respectively. The nodule biomass was determined by auger sampling to be 51±16 kg (dry weight) ha^–1. Most of the nodules were found at the 10–30 cm soil depth level. A rough calculation of the amount of nitrogen fixed annually arrived at 110±30 kg ha^–1. The results give strong support for the use ofL. leucocephala for soil enrichment in less humid areas of tropical Africa.     

Colony dynamics in a primitively eusocial halictine beeLasioglossum (Dialictus) zephyrum (Hymenoptera: Halictidae)

Summary Proximate control of colony dynamics was studied in the primitively eusocial halictine beeLasioglossum (Dialictus) zephyrum using allozyme markers. The results indicate that workers produce on average 15% of the male brood (range=0–50%) in small laboratory colonies made up of unrelated, single-generation, uninseminated females. This proportion is not influenced by colony size, but is influenced by the relative size of the queen. Large queens are more successful in dominating their workers than are small queens, the queen being defined as the female that is the mother of most of the brood produced in the colony. Older and larger females tend to become queens. Thus, while small differences in age (up to 4 days) influence which female becomes a queen, her ability to control her workers is primarily influenced by her relative size. The proportion of reproduction that is co-opted by the queen is negatively correlated with colony reproductivity (the number of males/day/female). Colony reproductivity is also negatively correlated with the standard deviation in size among females.     

Bioenergetics and thermal physiology of American water shrews (<Emphasis Type="Italic">Sorex palustris</Emphasis>)

Rates of O₂ consumption and CO₂ production, telemetered body temperature (T_b) and activity level were recorded from adult and subadult water shrews (Sorex palustris) over an air temperature (T_a) range of 3–32°C. Digesta passage rate trials were conducted before metabolic testing to estimate the minimum fasting time required for water shrews to achieve a postabsorptive state. Of the 228 metabolic trials conducted on 15 water shrews, 146 (64%) were discarded because the criteria for inactivity were not met. Abdominal T_b of S. palustris was independent of T_a and averaged 38.64±0.07°C. The thermoneutral zone extended from 21.2°C to at least 32°C. Our estimate of the basal metabolic rate for resting, postabsorptive water shrews (96.88±2.93 J g^–1 h^–1 or 4.84±0.14 ml O₂ g^–1 h^–1) was three times the mass-predicted value, while their minimum thermal conductance in air (0.282±0.013 ml O₂ g^–1 h^–1) concurred with allometric predictions. The mean digesta throughput time of water shrews fed mealworms (Tenebrio molitor) or ground meat was 50–55 min. The digestibility coefficients for metabolizable energy (ME) of water shrews fed stickleback minnows (Culaea inconstans) and dragonfly nymphs (Anax spp. and Libellula spp.) were 85.4±1.3% and 82.8±1.1%, respectively. The average metabolic rate (AMR) calculated from the gas exchange of six water shrews at 19–22°C (208.0±17.0 J g^–1 h^–1) was nearly identical to the estimate of energy intake (202.9±12.9 J g^–1 h^–1) measured for these same animals during digestibility trials (20°C). Based on 24-h activity trials and our derived ME coefficients, the minimum daily energy requirement of an adult (14.4 g) water shrew at T_a = 20°C is 54.0 kJ, or the energetic equivalent of 14.7 stickleback minnows.     

Discovery of independent‐founding solitary queens in the yellow crazy ant Anoplolepis gracilipes in East Java,Indonesia (Hymenoptera: Formicidae)

We collected four solitary queens of the invasive ant Anoplolepis gracilipes under stones in East Java, Indonesia. They produced nanitic workers by claustral colony foundation. This is the first report of independent colony foundation by queens in this species. The discovery may give an important insight into discussion on the origin of this invasive ant.     

Queen power in relation to age and mating status in the Argentine antIridomyrmex humilis (Mayr)

Summary Queen power inIridomyrmex humilis during six arbitrarily chosen physiological stages of queens (virgin winged queens at the time of emergence, the same 4–5 days old, mated winged queens 5–6 days old, mated queens at the time of dealation, young egg-laying queens, and old egg-laying queens) was tested with regard to workers. Tested workers and queens were nestmates. The results were as follows. 1) Power of the old queens remained rather constant throughout the reproductive season. 2) Young queens were always less attractive than older ones. No changes were observed from the time of emergence to dealation. 3) At the time of egg-laying, these young queens became markedly attractive for workers but never as much as old egg-laying queens. Therefore, insemination and dealation do not trigger the increase in queen power. In contrast, oogenesis ending by egg-laying produces a significant increase in queen power.     

Axial rod growth and age estimation of the sea pen,Halipteris willemoesi Kölliker

Halipteris willemoesi is a large octocoral commonly found in the Bering Sea. It is a member of a ubiquitous group of benthic cnidarians called sea pens (Octocorallia: Pennatulacea). Sea pens have a skeletal structure, the axial rod, that in cross section exhibits growth rings. Pairs of adjacent rings, or ring couplets, were assumed to be annuli and were used to estimate the age and growth of H. willemoesi. Twelve axial rods, extracted from H. willemoesi collected in the Bering Sea, were selected to represent small (25–29 cm total length), medium (97–130 cm TL) and large (152–167 cm TL) colonies. Each rod resembled a tapered dowel; the thickest part (0.90–6.75 mm in diameter) was at about 5–10% of total length from the base tip, the distal part was more gradually tapered than was the base. The number of ring couplets increased with rod size indicating their utility in estimating age and growth. Estimated age among rods was based on couplet counts at the thickest part of each rod; the average estimated age (±SE) was 7.1 ±0.7, 19.3 ±0.5, and 44.3 ±2.0 yr for small, medium and large-size rods, respectively. Based on these estimated ages, average growth rate in total length was 3.9 ± 0.2, 6.1 ± 0.3, and 3.6 ± 0.1 cm yr^–1 for small, medium, and large-size colonies. The average annual increase in maximum rod diameter among all colonies was 0.145 ± 0.003 SE mm yr^–1; therefore, age prediction from maximum rod diameter was calculated (estimated age (yr) = 7.0 * (maximum rod diameter, mm) –0.2; R ² = 0.99). At maximum diameter, the average couplet width was relatively constant among the three colony sizes (0.072 ± 0.05 mm). X-ray diffraction and electron microprobe analyses revealed that the inorganic portion of the rod is composed of a high-magnesium calcite. Radiometric validation of these age and growth rate estimates was attempted, but high amounts of exogenous ²¹⁰Pb precluded using the disequilibria of ²¹⁰Pb:²²⁶Ra. Instead, ²¹⁰Pb activities were measured in a series of cores extracted along the axial rod. These activities ranged from 0.691 ± 0.036 (SE) to 2.76 ± 0.13 dpm g^–1, but there was no pattern of decay along the length of the rod; therefore, the growth rates and corresponding ages could not be validated. Based on estimated age from ring couplet counts, growth in total rod length is slow at first, fastest at medium size, and slows toward maximum size, with an estimated longevity approaching 50 yr.     

Assessing fine-root biomass and production in a Scots pine stand – comparison of soil core and root ingrowth core methods

Soil core and root ingrowth core methods for assessing fine-root (< 2 mm) biomass and production were compared in a 38-year-old Scots pine (Pinus sylvestris L) stand in eastern Finland. 140 soil cores and 114 ingrowth cores were taken from two mineral soil layers (0–10 cm and 10–30 cm) during 1985–1988. Seasonal changes in root biomass (including both Scots pine and understorey roots) and necromass were used for calculating fine-root production. The Scots pine fine-root biomass averaged annually 143 g/m² and 217 g/m² in the upper mineral soil layer, and 118 g/m² and 66 g/m² in the lower layer of soil cores and ingrowth cores, respectively. The fine-root necromass averaged annually 601 g/m² and 311 g/m² in the upper mineral soil layer, and 196 g/m² and 159 g/m² in the lower layer of soil cores and ingrowth cores, respectively. The annual fine-root production in a Scots pine stand in the 30 cm thick mineral soil layer, varied between 370–1630 g/m² in soil cores and between 210 – 490 g/m² in ingrowth cores during three years. The annual production calculated for Scots pine fine roots, varied between 330–950 g/m² in soil cores and between 110 – 610 g/m² in ingrowth cores. The horizontal and vertical variation in fine-root biomass was smaller in soil cores than in ingrowth cores. Roots in soil cores were in the natural dynamic state, while the roots in the ingrowth cores were still expanding both horizontally and vertically. The annual production of fine-root biomass in the Scots pine stand was less in root ingrowth cores than in soil cores. During the third year, the fine-root biomass production of Scots pine, when calculated by the ingrowth core method, was similar to that calculated by the soil core method. Both techniques have sources of error. In this research the sampling interval in the soil core method was 6–8 weeks, and thus root growth and death between sampling dates could not be accurately estimated. In the ingrowth core method, fine roots were still growing into the mesh bags. In Finnish conditions, after more than three growing seasons, roots in the ingrowth cores can be compared with those in the surrounding soil. The soil core method can be used for studying both the annual and seasonal biomass variations. For estimation of production, sampling should be done at short intervals. The ingrowth core method is more suitable for estimating the potential of annual fine-root production between different site types.     

Nest-founding behavior induced in the first descendants ofPolistes dominulus Christ (Hymenoptera: Vespidae) colonies

Summary First descendants' founding ability was tested inP. dominulus colonies by experimentally removing the original nests and queens 21–24 days after the first emergence. Foundation tests were carried out after three different periods had elapsed after the removal and the results were analyzed with regards to the effects of participation in social tasks, duration of stay within the colony and seasonal factors. 1.Immediately after the removal, the foundations that occurred were performed almost entirely by workers, regardless of the duration of their stay within the colony. The number of foundations depended on the season in which the females had emerged. 2.Two months and four months after the removal, foundations were made by only a small number of females (workers and non-workers). These females, which were therefore able to found a nest the year they were born, were mostly among the very first-born individuals in the colonies. 3.The following year after hibernation: females (workers and non-workers) which had stayed for only a short time within the colony (1–15 days) had the highest survival rates. The foundation rates among the surviving females depended only on seasonal factors, since the foundation rates were higher among the non-workers which had emerged later in the season. The results throw some light on the first descendants' nest-founding potential in colonies developing under natural conditions, where many workers in fact stay for only a short time at the nest.     

Comparisons of primary production and nutrients absorption by aMiscanthus sinensis community in different soils

Soil properties, primary production, nitrogen and phosphorus uptake in aMiscanthus sinensis community on serpentine gangue area were compared with that on nonserpentine area. Soil water content, soil pH and nitrogen content were quite different between the serpentine gangue area and nonserpentine area; but phosphorus content of the soil was similar between the two sites. The maximum above-ground net production in the serpentine gangue and nonserpentine areas was 4.5±0.2 kg m^–2 yr^–1 and 7.8±0.2 kg m^–2 yr^–1, respectively. The total maximum standing biomass in the serpentine gangue and nonserpentine areas was 8.5±0.8 kg m^–2 and 11.9±0.4 kg m^–2, respectively. Nitrogen uptake by plants in the nonserpentine area was 2.4 times greater than that in the serpentine gangue area. Phosphorus uptake by plants were similar for the two sites. The most probable reasons for the small biomass produced by theMiscanthus sinensis community in this serpentine gangue area are the low levels of nitrogen and water availability in the soil.     

Nest architecture of the ant <Emphasis Type="Italic">Formica pallidefulva</Emphasis>: structure,costs and rules of excavation

Summary The architecture of underground ant nests was studied in the ant Formica pallidefulva. These ants build shallow (30–45 cm deep) nests, which consist of more or less vertical shafts that bear chambers. Shafts are modular units of nest growth; nests are enlarged by adding more shafts or extending previously existing ones. The nests are top-heavy, their volume declining exponentially with depth. The total volume of the nest is strongly correlated with the number of worker occupying the nest (R² = 0.87). Some of the rules and templates used by workers for nest construction were determined: (a) chambers are formed in the direction of the tunnels leading up to them, (b) the amount of soil excavated per unit time increases with soil temperature and moisture content. The amount of time and energy required to construct a typical nest were approximated using digging ability parameters determined in the lab. We estimate that if a colony was to move twice a year, it would expend around 20% of its energy intake and at least 6% of its worker time on nest excavation.Received 16 December 2002; revised 10 July 2003; accepted 23 July 2003.     

Worker demography and behavior in a supercolonial ant colony: The case of the desert ant Cataglyphis niger

Insect societies headed by multiple queens (polygyny) raise evolutionary questions, such as how does genetic heterogeneity among colony members affect in-nest interactions; or, are all queens equally reproductive or equally treated by workers? Answering such questions requires intensive and continuous observations of in-nest behavior. Here, we addressed these questions in the polygyne supercolonial ant, Cataglyphis niger, using a barcoding system that enables the tracking of individual interactions, together with polymorphic DNA microsatellite markers that indicate the matriline and patriline of all individuals. Our findings that both queens and workers have low interrelatedness corroborate earlier reports regarding the supercolony structure of C. niger. Ovary inspection and worker genotyping revealed that all the queens contribute similarly to nest demography. Tracking positions of individual workers through time revealed that only a small proportion of them are constantly engaged in tending the queens and can be considered as retinue workers. However, genotyping these workers and the attended queens revealed no relationship to genetic relatedness, again typical of a true polygyne and supercolonial species. Unlike invasive supercolonial species, C. niger is native to Israel, enabling us to address questions regarding the driving forces, other than kin selection, that stabilize this society.     

The efficacy of Scenedesmus morphology as a defense mechanism against grazing by selected species of rotifers and cladocerans

Phytoplankton often develop various defense mechanisms in response to zooplankton grazing, such as spines and colonies. While it is now known that increased spine length and cells in a colony of members of the genus Scenedesmus, when zooplankton grazing is intense, helps in reducing zooplankton filtering rates, the effect of these defense mechanisms at the population level has been observed in few studies. Here we present data on the growth rates of four zooplankton species, Brachionus calyciflorus, B. patulus, Ceriodaphnia dubia and Daphnia pulex at two food levels using two species of colony-forming Scenedesmus spp.: S. acutus (cell length = 18.2 ± 0.4 µm; width = 4.2 ± 0.1 µm; average colony length = 90 µm; width: 21 µm) and S. quadricauda (cell length: 21 ± 0.5 width 7.5 ± 0.3 µm; average colony length: 84 µm; width: 30 µm). Whereas S. acutus had no spines, S. quadricauda had spines of 6–10 µm. Population growth experiments of the test rotifers and cladocerans were conducted in 100 ml containers with 50 ml of the medium with test algae. Algae concentrations used were: 13 and 52 mg dw l^–1 of each of the two algal species offered in colonial forms. We used an initial inoculation zooplankter density of 1 ind. ml^–1 for either of the rotifer species and 0.2 ind. ml^–1 for either of the cladoceran species. In all, we had 64 test containers (4 test species of zooplankton × 2 test species of algae × 2 algal densities × 4 replicates). We found a significant effect of algal size on the growth rates of all the four tested species of zooplankton. The population growth rates of zooplankton ranged from –0.58 to 0.66 and were significantly higher on diet of S. acutus than of S. quadricauda. Thus, our study confirms that the larger colony size and the formation of spines in S. quadricauda were effective defenses against grazing by both rotifers and smaller sized cladoceran Ceriodaphnia dubia but that larger-bodied Daphnia pulex could exploit both the algal populations equally.