Response of sea urchin pluteus larvae (Echinodermata: Echinoidea) to reduced seawater pH: a comparison among a tropical, temperate, and a polar species

Ocean acidification, as a result of increased atmospheric CO₂, is predicted to lower the pH of seawater to between pH 7.6 and 7.8 over the next 100 years. The greatest changes are expected in polar waters. Our research aimed to examine how echinoid larvae are affected by lower pH, and if effects are more pronounced in polar species. We examined the effects of lowered pH on larvae from tropical (Tripneustes gratilla), temperate (Pseudechinus huttoni, Evechinus chloroticus), and a polar species (Sterechinus neumayeri) in a series of laboratory experiments. Larvae were reared in a range of lower pH seawater (pH 6.0, 6.5, 7.0, 7.5, 7.7, 7.8 and ambient), adjusted by bubbling CO₂ gas. The effect of pH on somatic and skeletal growth, calcification index, development and survival were quantified, while SEM examination of the larval skeleton provided information on the effects of seawater pH on the fine-scale skeletal morphology. Lowering pH resulted in a decrease in survival in all species, but only below pH 7.0. The size of larvae were reduced at lowered pH, but the external morphology (shape) was unaffected. Calcification of the larval skeleton was significantly reduced (13.8–36.9% lower) under lowered pH, with the exception of the Antarctic species, which showed no significant difference. SEM examination revealed a degradation of the larval skeletons of Pseudechinus and Evechinus when grown in reduced pH. Sterechinus and Tripneustes showed no apparent difference in the skeletal fine structure under lowered pH. The study confirms the need to look beyond mortality as a single endpoint when considering the effects of ocean acidification that may occur through the 21st century, and instead, look for a suite of more subtle changes, which may indirectly affect the functioning of larval stages.     

pH effect on the susceptibility to parasitoid infection in the marine diatom Coscinodiscus spp. (Bacillariophyceae)

The pH on the frustule of individual cells of the marine centric diatoms Coscinodiscus granii and Coscinodiscus wailesii (Bacillariophyceae) was measured with pH microsensors in culture media with increasing pH values of 8.04, 8.14, and 8.22, respectively. In 85–96% of the C. granii cells the pH on the frustule was up to 0.4 units higher than that of the medium, reaching a maximum pH 8.95. Only in 2–3% the surface pH exceeded that of the medium by up to 0.7 pH units. These results strongly suggest that diatoms in batch cultures differ, at least temporarily, in their individual photosynthetic activities. Infection experiments with the parasitoid nanoflagellate Pirsonia diadema (Stramenopile) showed that flagellates failed to infect when the culture pH was 8.8 and above. pH measurements on freshly infected C. granii showed that the prevalence of infection was higher in tendency on diatoms with low surface pH. Application of these results to parasitoid-diatom interactions in natural waters suggests that within phytoplankton populations a strong photosynthetic activity might prevent diatom cells temporarily from infection by pH-sensitive parasitoids.     

Studies on the causes of mortality of the estuarine bivalve Macoma balthica under conditions of (near) anoxia

Survival of the bivalve Macoma balthica in (near) anoxic seawater was studied in a static system and a flow-through system and compared with emersed exposure to air and N₂. In the static system, a decrease in pH and exponential accumulation of sulphide in the incubation medium were observed, indicating excessive growth of (sulphate-reducing) bacteria. These changes in the chemical environment were prevented by the use of a flow-through system. However, this treatment hardly affected survival time. Median mortality times were 8.3 and 9.0 days for the static and flow-through incubation, respectively. Addition of the antibiotic chloramphenicol strongly increased survival time in both systems with corresponding values of 17.9 and 23.0 days. A similar value was obtained for survival in air (LT₅₀= 21.7 days). In a second experiment (1 year later), we obtained much lower values for anoxic survival in a static system, although laboratory conditions, season and temperature were similar. The pH values were adjusted to 6.5, 7.2 and 8.2 by buffering the media (25 mM Tris-HCl), and the corresponding LT₅₀ values were 5.5, 5.7 and 4.7 days, respectively. In the presence of chloramphenicol the values were 10.8, 10.9 and 9.5 days, respectively. These values show that a slightly acidic medium increased survival time. Exposure to an atmosphere of N₂ resulted in a survival time close to that in anoxic seawater without chloramphenicol (LT₅₀= 6.4 days). Overall the results indicate that proliferation of anaerobic bacteria associated with the bivalves was the main cause of death. Since chloramphenicol also displayed a strong positive effect in the flow-through system, which prevented the accumulation of released waste products and a decrease of pH, bacterial damage must have been by injury of the tissues of the clams and not by the release of noxious compounds to the medium. Bacterial outbreaks are a part of every anoxic event (eutrophication), and therefore, in their habitats, direct bacterial infection may also be the cause of clam mortality. It is concluded that laboratory studies on anoxic tolerance, or impact of sulphide, may produce artefacts when no precautions are taken to suppress bacterial proliferation. Received: 4 July 2000 / Accepted: 6 December 2000     

Effect of Water Variables On Lead Tolerance in Postlarvae of Penaeus Indicus (H. Milne Edwards)

Lead tolerance in Penaeus indicus post-larvae (PL) was studied in relation to the water variables: salinity, temperature and pH. the LC₅₀ for 96 hrs was 7.22 ppm at ambient conditions of salinity (20%_o), temperature (29°C) and pH (7.2). the post-larvae were found to be sensitive to salinity variations with a significant (P < 0.05) low LC₅₀ values at lower (2%_o) and higher (29%_o) ranges. Though a decrease in LC₅₀ value was observed both at higher (45°C) and lower (10°C) temperatures, it was significantly (P < 0.05) low only at higher temperature indicating more toxicity. the LC₅₀ values also showed a significant (P < 0.05) decrease in acidic (pH 2.8) and alkaline (pH 11.0) conditions. the data indicate that lead toxicity increases in the PL of P. indicus with variations in the water variables.     

The response of two ecologically important Antarctic invertebrates (Sterechinus neumayeri and Parborlasia corrugatus) to reduced seawater pH: effects on fertilisation and embryonic development

Ocean acidification, or the lowering of seawater pH, is caused by sequestration of atmospheric CO₂ into the oceans. This study investigated the effects of present-day pH 8.0, predicted ocean surface pH for the years 2100 and 2300 (pH 7.7 and pH 7.3, respectively) and an extreme pH (pH 7.0) on fertilisation and embryogenesis in the Antarctic nemertean worm Parborlasia corrugatus and sea urchin Sterechinus neumayeri. Fertilisation success was not affected by pH in P. corrugatus across a range of sperm concentrations. Fertilisation success in S. neumayeri declined significantly in pH 7.0 and 7.3 seawater, but only at a low sperm concentration. Seawater pH had no effect on the rate of egg cleavage in S. neumayeri, or the proportion of abnormal embryos 1-day post-fertilisation. P. corrugatus embryogenesis was also relatively robust to pH changes, with a significant effect detected only when the seawater pH was decreased to 7.0. While fertilisation and early cell division were relatively robust, later development through to the gastrula was sensitive to pH. In S. neumayeri, an effect of pH on development was evident by the gastrula stage, while there were significantly more abnormal P. corrugatus embryos in pH 7.0 up to the blastula stage, and in pH 7.0 and pH 7.3 at the coeloblastula stage. Our results are similar to the observations on other marine invertebrate species where fertilisation and early embryonic development are generally robust to lowered seawater pH, while the older coeloblastula and gastrula stages are more responsive. We also found no evidence to suggest that Antarctic species are more adversely affected by lower seawater pH compared with the findings for non-Antarctic counterparts. We conclude that in the two species we examined, near-future decreases in pH (decreases of ≈0.3–0.5 pH units) may not have a significant effect on fertilisation and early embryogenesis, while predicted longer term decreases (decreases of ≈0.7–0.77 pH units) could reduce fertilisation success in S. neumayeri if sperm concentrations are low and may increase abnormalities in P. corrugatus during later embryogenesis.     

Isolation and characterization of an Arthrobacter sp. strain HB-5 that transforms atrazine

A bacterial strain (HB-5) capable of utilizing atrazine as sole carbon and nitrogen source for growth was isolated from an industrial wastewater sample by enrichment culture. The isolate was identified as Arthrobacter sp. according to its phenotypic features, physiologic and biochemical characteristics, and phylogenetic analysis. The strain exhibited faster atrazine degradation rates in atrazine-containing mineral media than the well-characterized atrazine-degrading bacteria Pseudomonas sp. ADP. The broad optimum pH and temperature ranges observed for strain HB-5 indicate that it has potential for remediation of atrazine-contaminated sites. Strain HB-5 first metabolizes atrazine to yield hydroxyatrazine. Then, the bacterium metabolizes hydroxyatrazine to cyanuric acid, but could not mineralize atrazine.     

Ichthyotoxicity found in cultured media of Protogonyaulax spp.

Ichthyotoxicity was observed in the cultured media from Protogonyaulax catenella and P. tamarensis. No paralytic shellfish poison was detected in the cultured media. Test fish in the cultured media died showing signs of hypoxia. Histological observations of the gills of intoxicated fish indicated that the epithelial cells were swollen and exfoliated from the pillar cell. The cultured media had hemolytic activity against erythrocytes of various animals. The media also decreased the elasticity of eggs of rainbow trout Salmo gairdneri. From these results, it is concluded that the cultured media of both species of Protogonyaulax contain a cytotoxin which is lethal to fish.     

Lethal effects of Northwest Atlantic Ocean isolates of the dinoflagellate, Scrippsiella trochoidea, on Eastern oyster (Crassostrea virginica) and Northern quahog (Mercenaria mercenaria) larvae

The thecate dinoflagellate Scrippsiella trochoidea is a cosmopolitan, bloom-forming alga that has been generally considered non-toxic. Here, we report that environmentally relevant cell densities (10⁴ cells mL⁻¹) of Scrippsiella trochoidea strains isolated from the Northwest Atlantic Ocean caused 100% mortality in Eastern oyster (Crassostrea virginica) larvae during 3-day exposures while parallel control larvae exhibited 100% survival. S. trochoidea also exhibited lethal effects on Northern quahog (Mercenaria mercenaria) larvae (70% mortality during 3-day exposure) but were non-toxic to juvenile fish (Cyprinodon variegates). The cultures of S. trochoidea were more lethal to Northern quahog larvae than ten other species of harmful algae, including the highly toxic species Cochlodinium polykrikoides. Scrippsiella trochoidea cultures within later stages of growth were more toxic than exponential growth stages to bivalve larvae, and the toxicity was dose dependent. Furthermore, toxicity was maintained in the cultures that were sonicated, boiled, and frozen as well as in resuspended residues of the culture but was significantly lower in cell-free culture media. Collectively, these results suggest that S. trochoidea causes mortality in bivalve larvae through a physicochemical rather than strictly chemical mechanism, such as clogging of larval feeding apparatuses by materials produced by S. trochoidea (e.g., lipids, extracellular polysaccharides, and/or cell debris) which accumulate as cells in culture or blooms age. This is the first report of the lethal effects of Scrippsiella trochoidea on shellfish larvae.     

Photosynthetic utilization of CO2(aq) and HCO 3 - in Thalassia testudinum (Hydrocharitaceae)

The effects of total dissolved inorganic carbon (DIC), free carbon dioxide [CO_2(aq)], and bicarbonate (HCO ₃ ^- ) concentrations on net photosynthetic oxygen evolution of the marine angiosperm Thalassia testudinum Banks ex König collected from Biscayne Bay (1988) and from Tampa Bay (1990), Florida, USA, were examined. Rates of photosynthesis declined by 85% from pH 7.25 to 8.75 in buffered seawater media with constant DIC concentration (2.20 mM), suggesting a strong influence of CO_2(aq) concentration. A plateau in the pH-response curve between pH 7.75 and 8.50 indicated possible utilization of HCO ₃ ^- . Responses of photosynthesis measured in buffered seawater media of varying DIC concentrations (0.75 to 13.17 mM) and pH (7.8 to 8.61) demonstrated that photosynthesis is rate-limited at ambient DIC levels. Photosynthesis increased in media with increasing HCO ₃ ^- concentrations but near-constant CO_2(aq) levels, confirming HCO ₃ ^- assimilation. Calculated half-saturation constants (K _s )for CO_2(aq) and HCO ₃ ^- indicated a high affinity for the former [K _s (CO₂)=3 to 18 M] and a much lower affinity for the latter [K _s (HCO ₃ ^- )=1.22 to 8.88 mM]. Calculated V _max values for HCO ₃ ^- were generally higher than those for CO_2(aq), suggesting relatively efficient HCO ₃ ^- utilization, despite the apparent low affinity for this carbon form.     

Biodegradation kinetics of cymoxanil in aquatic system

A potential method to detoxify pesticides in aquatic system is using bioremediation. In this study, four microorganisms (Pseudomonas sp (EB11), Streptomyces sp. (EB12), Aspergillus niger (EB13) and Trichoderma viride (EB14) were isolated from cucumber leaves previously treated with cymoxanil using enrichment technique. These strains were evaluated for their potential to detoxify cymoxanil in aquatic system at the concentration level of 5×10^?4M. The effect of pH and temperature on the growth ability of the tested strains was also investigated by measuring the intracellular protein and mycelia dry weight for bacterial and fungal strains, respectively. Moreover, the remaining toxicity of cymoxanil after 28 days of incubation with tested strains was evaluated to confirm the complete removal of any toxic materials (cymoxanil and its metabolites). The results showed that the optimum pH for the growth of cymoxanil degrading strains (bacteria and fungi) was 7. A temperature of 30°C appears to be the optimum for the growth of either fungal or bacterial strains. Pseudomonas sp. (EB11) was the most effective strain in cymoxanil degradation followed Streptomyces sp (EB12), Trichoderma viride (EB14) and Aspergillus niger (EB13), with half-lives of 4.33, 9.5, 17.3 and 24.7 days, respectively. The degradation of cymoxanil by bacterial strains was much faster than fungal one. There is no remaining toxicity of cymoxanil detected in aqueous media previously treated with Pseudomonas sp. (EB11) for 28 days. The results suggest that bioremediation by Pseudomonas sp. (EB11) are promising for the detoxification of cymoxanil in aqueous media.     

Upper temperature tolerances of Tellina tenuis and T. fabula

The upper lethal tolerances of Tellina tenuis da Costa and T. fabula Gmelin were studied under a variety of temperature regimes in which both short-term and long-term mortality were noted; these included steady temperatures, acclimation at a steady temperature then exposure to a steady tempeature, and acclimation at gradually increasing temperatures. T. tenuis alone was also tested for variation in mortality rate according to the date of collection or to the size (i.e., age) of the mollusc. The results suggested that T. tenuis was consistently more tolerant than T. fabula, and that the difference in tolerance was around 5 C°. Both species readily acclimated to higher temperatures after steady temperature acclimation, but not under a regime of gradually increasing temperature. There was no significant, variation in the mortality rate of T. tenuis with season or age. It is suggested that, since the difference in temperature tolerance is not reflected in their respective geographical distributions, there may be some other stage in their life cycle, e.g. the larvae, which is equally susceptible to temperature extremes. The difference in tolerance is, however, reflected in their respective littoral and sublittoral distributions.     

Calcification in the articulated coralline alga Corallina pilulifera,with special reference to the effect of elevated CO2 concentration

Calcification in Corallina pilulifera Postels et Ruprecht displayed diurnal variations in aerated (350 ppm CO₂) culture media, with faster rates during the light than during the dark period. Addition of CO₂ (air+1250 ppm) inhibited calcification. This was attributable to the decreased pH resulting from CO₂ addition. Both photosynthesis and calcification were enhanced in seawater, with elevated dissolved inorganic carbon concentrations at a constant pH of 8.2.     

Interactive effects of pH,temperature and light during ammonia toxicity events in Elodea canadensis

Increased nutrient loading threatens many freshwater ecosystems. Elevated temperatures may increase the sensitivity to eutrophication in these ecosystems. Higher concentrations of possibly toxic reduced nitrogen (NH _x ) in the water layer may be expected as production and anaerobic breakdown rates will increase. Apart from temperature, NH _x and its effect on aquatic macrophytes will also depend on pH and light. We examined the interactive effects of NH _x , temperature, pH and light on Elodea canadensis in a full factorial laboratory experiment. Results demonstrate that high NH _x and high temperature together with low pH and low light causes the strongest toxic effects regarding relative growth rate and leaf tissue mortality. The adverse effects of high temperature and low light are most likely caused by increased metabolic activity and reduced photosynthesis, respectively. Severe toxicity at low pH compared to high pH can be ascribed to the ability of E. canadensis to induce a specialised bicarbonate-concentrating pathway at high pH, resulting in much higher carbon availability, needed for detoxification of NH _x . We conclude that NH _x toxicity will become more pronounced under higher temperatures, but that effects on aquatic macrophytes will strongly depend on pH of the water layer and specific metabolic adaptations of different species.     

Comparison of health patterns in two geochemically contrasted areas of northern Finland

In order to prepare the ground for a discussion of possible causal relationships between health patterns and the geochemistry of an area, health patterns in the form of age-adjusted cancer incidence values and rates of mortality due to (1) cancer of the stomach, colon or rectum, (2) lung cancer and (3) ischaemic heart diseases are compared between the populations of two geochemically contrasted communes of northern Finland (Kemijärvi and Kuusamo). Statistically significant differences (p < 0.001) are encountered in the mortality rates due to ischaemic heart diseases and marked differences in the incidence values of lung cancer, the former being higher in Kuusamo and the latter in Kemijärvi. The higher incidence of lung cancer observed in Kemijärvi is in accordance with previous results reported from the USA, where higher incidences have been observed in areas of granitic bedrock. Regarding the higher mortality rate due to ischaemic heart diseases In Kuusamo, which is geochemically characterised by higher potassium (p < 0.002) and calcium (p < 0.004) and lower Zn, V, Ti, and Fe (p > 0.002) in the soil (actually, in the fine till fraction), no easily demonstrable cause- and-effect relationship can be established.     

Mechanisms of habitat segregation between an invasive and an indigenous mussel: settlement,post-settlement mortality and recruitment

The mussel Mytilus galloprovincialis is highly invasive worldwide, but displays varying degrees of local and regional coexistence with indigenous mussels through spatial habitat segregation. We investigated the roles of settlement, post-settlement mortality, juvenile growth and recruitment in partial habitat segregation between the invasive M. galloprovincialis and the indigenous mussel Perna perna on the south coast of South Africa. We used two study locations, Plettenberg Bay and Tsitsikamma, 70 km apart, with two sites (separated by 300–400 m) per location, each divided into three vertical zones. There were no significant effects in Tsitsikamma, where daily settlement and monthly recruitment were significantly lower than in Plettenberg Bay. In Plettenberg Bay, settlement (primary and secondary) and recruitment of both species decreased upshore. Post-settlement mortality was measured over two consecutive 6-day periods during a spring tide and a neap tide. For both species mortality was low on the low-shore. High-shore mortality was consistently low for M. galloprovincialis, but increased dramatically for P. perna during spring tide. No data were obtained for growth of P. perna, but juvenile M. galloprovincialis grew more slowly farther upshore. P. perna recruited mainly in spring and summer, with a peak in summer far greater than for M. galloprovincialis. Recruitment of M. galloprovincialis was more protracted, continuing through autumn and winter. Thus local coexistence is due to a combination of pre- and post-recruitment factors differing in importance for each species. P. perna is excluded from the high-shore by recruitment failure (low settlement, high mortality). High survival and slow growth in juveniles may allow large densities of M. galloprovincialis to accumulate there, despite low settlement rates. With no differences between species in settlement or mortality on the low-shore, exclusion of M. galloprovincialis from that zone is likely to be by post-recruitment processes, possibly strengthened by periodic heavy recruitments of P. perna. At larger scales, larval retention and protracted recruitment contribute to the success of M. galloprovincialis at Plettenberg Bay, while recruitment limitation may explain why M. galloprovincialis is less successful at other sites.     

Differential sensitivity of marine infaunal amphipods to tributyltin

Three species of infaunal gammaridean amphipods, Rhepoxynius abronius (Phoxocephalidae), Eohaustorius washingtonianus, and E. estuarius (Haustoriidae) were tested in a water-only system to assess their sensitivity to tributyltin (TBT) without the influence of factors that could affect bioavailability. When mortality (LC₅₀) was the endpoint, the results indicated that R. abronius was 20 times more tolerant to tributyltin than either haustoriid species; however, when mortality plus reburial behavior (EC₅₀) was assessed, the difference was only about 10 times. The bioconcentration factor (BCF) was also consistently lower in R. abronius (11.1 to 16.5 times) than in the haustoriids; however, when the LD₅₀ was calculated, the concentration in the tissues associated with 50% mortality for each species was not significantly different. The large disparity in species' response is attributed to reduced uptake and a potentially greater ability to metabolize this compound by R. abronius. An analysis of TBT uptake confirmed that R. abronius was able to accumulate less TBT and hence maintain a low body burden for a given water concentration. The results of a separate uptake study were used to formulate a hypothesis for observed differences in reburial behavior. Because the rate of TBT uptake was lower in R. abronius, we propose that the slower rise in toxicant body burden allowed for a gradual response in this species which included a sublethal effect (non reburial), compared to a rapid rise in the body burden for E. estuarius which caused the response to quickly proceed from no effect to death.     

Ammonia excretion of Artemia sp. (crustacea: Branchiopoda) under axenic conditions)

Ammonia excretion of the Utah strain of Artemia sp. grown in axenic culture media was followed throughout the different stages of larval development. The ammonia accumulated in the culture medium does not affect ammonia excretion rates under our laboratory conditions. Growth rate, density and starvation all affect ammonia excretion. Ammonia excretion is directly related to the numerical growth index and inversely related to Artemia sp. density. Starved adult Artemia sp. excrete considerably lower amounts of ammonia than brine shrimp grown in nutritive medium. In view of the difficulties often encountered in estimating the excretion rates of small crustaceans, our culture conditions would appear advantageous for further studies on the purine metabolism of Artemia sp.     

Reproductive effort of colonial and solitary breeding tree sparrows Passer montanus L.

A total of 250 nestboxes were arranged in five plots in a suburban area of Budapest, Hungary (19°04E, 47°41N). In each plot, 25 were placed at 50 m intervals to simulate solitary breeding and 25 3–5 m apart to simulate colonial breeding. Length of nest building period, feeding frequency, nestling mortality, nestlings' diet, productivity and parental condition were compared for colonial and solitary breeding tree sparrows Passer montanus. Parents with long nest-building periods, including the majority of first-year females, produced fewer young than parents which built over short periods. Parents fed nestlings morefrequently and nestlings had lower mortality in second than first broods; whether or not a third brood was reared was determined by the costs invested in first and second broods. Females that laid a third clutch had reared fewer young in first and second broods and were heavier than females that reared many young in two broods. Colonial birds had higher feeding frequencies, more similar diets and suffered lower nestling mortality than solitary parents for first broods, but they fed less frequently, diets were less similar, and nestling mortality was higher in second and third broods. It is suggested that colonial breeders benefited from the social stimulation of simultaneous feeding in first broods, but the advantage of synchronicity in feeding declined in second broods and the sparser breeding spacing of solitary parents was more advantageous for feeding in second and third broods. Birds that changed nest spacing between broods had fed nestlings less frequently and had higher nestling mortality before changing than birds which retained their spacing. Parents which changed from colonies to solitary nests fed more frequently with lower nestling mortality in the next brood than parents which retained colonial nests for their second (and third) brood. Solitary parents did not show such a relationship. The rearing of three broods caused higher weight loss in colonial than solitary parents.Correspondence to: L. Sasvári     

Influence of culture media and environmental factors on mycelial growth and sporulation of Lasiodiplodia theobromae (Pat.) Griffon and Maubl

Lasiodiplodia theobromae, a common tea (Camellia sinensis) pathogen, usually does not sporulate or sporulates poorly in common media, which makes spore production difficult. In this study the effects of culture media, carbon source, nitrogen source, temperature, pH and light on mycelial growth and sporulation were evaluated. Among several carbon sources tested, glucose and sucrose were found superior for growth. Potassium nitrate supplemented media showed maximum growth amongst the tested inorganic nitrogen sources while peptone produced maximum growth among the tested organic nitrogen sources. Tea root extract supplemented potato dextrose agar medium was found to be the most suitable for mycelial growth and sporulation of L. theobromae. The fungus grow at temperatures ranging from 40 to 36 degrees C, with optimum growth at 28 degrees C and no growth was noted at 40 degrees C. There was no significant effect of different light period on growth of L. theobromae, but light enhanced sporulation. The fungus grow at pH 3.0-8.0 and optimum growth was observed at pH 6.0. Tea root extract supplemented potato dextrose agar medium with pH 6.0 was the most suitable for production of conidia of L. theobromae at 28 degrees C. Hence this media may be recommended for inoculum production for further studies.     

Variance in the male reproductive success of western gorillas: acquiring females is just the beginning

Variance in male reproductive success is expected to be high in sexually dimorphic mammals, even when it is modulated by the costs and benefits of group living. Here, we investigate the variance in reproductive success of male western gorillas (Gorilla gorilla), a highly dimorphic primate with long-term male–female associations, using 12.5 years of data collected at Mbeli Bai in northern Congo. Access to mates and offspring survival were both major sources of variance in male reproductive success. Males with larger harems had lower offspring mortality with no apparent reduction in female fertility or observed tenure length, so the size of harems did not seem to be limited by female feeding competition or by the risk of takeovers and infanticide by outsider males. The lower mortality in larger harems may reflect improved vigilance against predators, and females may cluster around males that enhance offspring survival. Thus, this study illustrates how a detailed analysis of the components of male reproductive success can shed light on the interrelated social and ecological aspects that affect it.