Competitive interactions between co-occurring invaders: identifying asymmetries between two invasive crayfish species

Ecosystems today increasingly suffer invasions by multiple invasive species. Complex interactions between invasive species can have different fitness implications for each invader, which can in turn determine the future progression of their invasions and result in differential impacts on native species and ecosystems. To this end, through pair-wise and group scale experiments, we examined possible interaction outcomes, competition effects and their potential fitness implications for two widespread invasive species of crayfish that increasingly co-occur in freshwater ecosystems of Europe (Pacifastacus leniusulus and Orconectes limosus). In all trials, P. leniusculus demonstrated the potential to outcompete O. limosus in both staged encounters and direct resource competition, being more likely to win heterospecific agonistic encounters and to acquire shelters at a higher rate. Observed dyadic dominance was translated to a broader social context of group-scale experiments, in which dominance of P. leniusculus was further strengthened by size differential between species. O. limosus was not able to compensate for competitive pressure by the dominant P. leniusculus and suffered wet weight loss and more frequent injuries in the presence of P. leniusculus. While both species are detrimental to native ecosystems, the ability of P. leniusculus to withstand competition pressure from another successful invasive species underscores its potential to establish dominant populations. Our results highlight the importance of understanding interspecies competition in prioritizing potential management activities or control efforts in contact zones.     

Relationships between fecal pellet production and feeding in the calanoid copepod Boeckella

The number, size (mm³ pellet ^–1) and total volume of fecalpellets produced (mm³ h^–1) by Boeckella titicacae increasedwith concentration of Chlorella and natural seston in threedifferent lakewaters from Bahia de Puno and Lago Grande of LakeTiticaca and nearby Laguna Arapa. The three lakewaters differedin the size and number of fecal pellets produced at seston concentrations>0.5 mg dry wt l^–1. However, the total volume of fecalpellets egested varied little. Large-sized food (>10 µg)resulted in larger pellets but smaller total fecal production.Food concentration had parallel effects on the production offeces and on feeding rates. Both fecal production and feedingincreased linearly with food concentration with saturation between5 and 7.5 x 10⁵ Chlorella ml^–1 (3.7–5.6 mg dry wtl^–1). Varied responses in fecal production when fed differentsize fractions of lake seston suggest Boeckella has a complexand flexible feeding behavior. Feeding preferences of B.titicacaeand B.occidentalis (Ivlev index) for two seston size fractionsare generally reflected by fecal production.     

Factors influencing the degree of sexual size dimorphism within and among calanoid copepod species

Populations of Diaptomus leptopus (Copepoda: Calanoida) and other calanoid copepods exhibit varying degrees of sexual size dimorphism. We examined whether intraspecific or interspecific variation in dimorphism could be explained by allometry, and we examined the relationship between adult size attained and development rate to determine any relationship between the two. We compared the degree of sexual size dimorphism in D. leptopus and in other calanoid copepods inhabiting temporary and permanent habitats. Allometry did not explain variation in sexual size dimorphism within or among populations or among species. Permanence of habitat affected the degree of dimorphism: dimorphism was greater within and among species inhabiting temporary environments. Non-significant differences in development rate were found among populations and significant differences were found between sexes of D. leptopus when reared under identical laboratory conditions: males developed more rapidly than females but there was no general relationship between development rate and adult size. Potential adaptive hypotheses to explain the differences between populations inhabiting temporary and permanent habitats are discussed.     

Diapause in the calanoid freshwater copepod Eudiaptomus graciloides

The seasonal appearance and the intensity of diapausing-eggproduction in Eudiaptomus graciloides in five lakes of differentsize and trophic level were studied. In all lakes, diapausingeggs were produced in autumn. In the large mesotrophic lakeSelenter See, the population’s shift to the productionof diapausing eggs was more complete than in other lakes. Weexamined day length, temperature and food as proximate factorsfor the production of diapausing eggs with laboratory experiments.Eudiaptomus graciloides produced diapausing eggs in all treatments,but a significantly higher percentage of diapausing eggs wasfound under short day conditions except when algal food wasabundant and temperature was high. To investigate the adaptivesignificance of diapause in E. graciloides, we compared thesurvival of adult and juvenile at different temperatures forE. graciloides with Eudiaptomus gracilis, a sympatric speciesthat does not exhibit diapause. At 8°C, adult E. graciloidessurvived better than adult E. gracilis and exhibited reducedgut contents and accumulation of storage lipids, traits characteristicof adult diapause. Eudiaptomus graciloides nauplii did not reachthe copepodid stage at 6°C, but E. gracilis nauplii exhibitedhigh mortality and developed very slowly. We hypothesize thatadult diapause and production of diapausing eggs facilitatethe survival of E. graciloides during cold periods and enhancecoexistence with its congener, E. gracilis, in temperate zones.     

The host—parasite relationship of two copepod species and two fish species

A total of 2494 Menidia beryllina and 717 M. peninsulae (Atherinidae) were examined from the Pensacola Bay area for parasitic copepods. M. peninsulae was infested with Bomolochus concinnus (Bomolochidae) and Ergasilus marticatus (Ergasilidae) and had incidences (and intensities) of 12.3% (1.6) and 4.2% (1.3), respectively. Only seven M. peninsulae were infested with both species of parasites. M. beryllina was infested only with E. manicatus and showed different incidences (and intensities) in two areas: Mulatto Bayou, 13.2% (1.9); Catfish Basin, 53.0% (2.3). Fishes with parasites were significantly longer than fishes without parasites and the case of increasing number of parasites with increasing fish length was reinforced. B. concinnus is a warm water parasite on M. peninsulae while E. manicatus acts as a cold water parasite on M. peninsulae and a warm water parasite on M. beryllina . The parasites tended to be overdispersed on their hosts but at the same time showed evidence of negative intraspecific associations within a host. These data suggest intraspecific avoidance by the parasite but active acquisition by the host.     

A calanoid copepod Gladioferens imparipes,holding to surfaces

Gladioferens imparipes, a calanoid copepod from estuaries in Western Australia, displays behaviour and associated morphology which is unusual among calanoids. Adult and copepodid stages make temporary attachment to underwater surfaces using fine hair sensillae on the surface of the prosome. A clear pattern can be seen in the number and arrangement of hair sensillae in early copepodid stages. Line drawings, photomicrographs and an S.E.M. are used to illustrate the structures. Laboratory studies with live copepods in a flow chamber show that adult animals may hold position against water currents. It is hypothesized that this behaviour influences the distribution pattern in an estuary where low velocity tidally induced water movement occurs, and may be of value in enabling the copepod to exist beyond the distribution of a major predator.     

Egg production rates of eight calanoid copepod species during summer 1997 off Newport, Oregon, USA

Measurements of hydrography, water transparency, chlorophyll (Chl) a and egg production rates (EPRs) by females of Calanus marshallae Frost, Calanus pacificus Brodsky, Eucalanus californicus Johnson, Epilabidocera longipedata Sato, Pseudocalanus mimus Frost, Centropages abdominalis Sato, Acartia longiremis Lilljerborg and Paracalanus parvus (Claus) were estimated at weekly intervals between 17 July and 2 September 1997. Production of eggs was determined in 24 h incubations to examine the effects of environmental variability on EPR, to detect the possibility of food limitation of EPR, and to evaluate the hypothesis that growth rates of females are size dependent. During the study, an anomalous downwelling event occurred, possibly in response to the 1997 El Niño, which allowed us to determine how El Niño events affect EPRs of coastal copepods. The larger copepods Calanus marshallae, Calanus pacificus and Centropages abdominalis showed the highest egg production and specific growth rates during the period of active upwelling (18 July-13 August, water temperatures 8-13C, Chl a concentration 4.7-16.2 l^-1 and water transparency 3-5 m). After 27 August, the 1997-98 El Niño arrived off Oregon, creating a downwelling situation. Upwelling winds ceased, the thermocline intensified, temperature and transparency increased (to >18°C and 16 m), and Chl a declined to <2 g l^-1. Densities of the common coastal species declined greatly as well. Paracalanus parvus became the dominant species, and Eucalanus californicus, Epilabidocera longipedata and Corycaeus anglicus became common in our samples. EPRs for the larger boreal copepods (Calanus and Centropages) declined greatly during El Niño;; the smaller copepods, Pseudocalanus mimus, A.longiremis and Paracalanus parvus, showed low but relatively constant egg production and specific growth rates during both upwelling and downwelling events. Over the entire study period, only three species produced eggs at its maximum rate (24-28 eggs female^-1 day^-1), Pseudocalanus mimus which averaged 4 eggs day^-1. All other species had EPRs that were two or five times below their maximum EPR. Thus, EPRs were not related to body size, contrary to our expectations. Hatching success was variable among species. Values as low as 20-40% were found for all species at least once during the study period, suggesting that occasionally a substantial portion of egg production may not be viable.      

Habitat and life history differences between two species of Gambusia

Life history strategies reflect trade-offs that tend to maximize fitness, such as investment in a few large or many small offspring. We compared life histories of two temperate livebearing fishes Gambusia affinis and G. nobilis, an endangered species which is virtually unstudied. The two species persist in environments that differ widely in abiotic and biotic factors in the same local area. Gambusia affinis were typically found in habitats with high productivity and wide fluctuations in temperature, salinity and dissolved oxygen, whereas G. nobilis occurred in more stable spring-fed habitats. We collected data on life-history traits: embryo mass, brood size (number of embryos), total maternal reproductive effort, population sex ratios, and size (mass and length) distributions of adults and juveniles. There was no difference between species in reproductive effort per brood, but they differed in investment strategy. Gambusia affinis females produced large broods with small embryos, whereas G. nobilis females produced broods of fewer, larger embryos. These differences in life history strategies reflect a tradeoff between individual productivity and differential mortality rates in different environments. At our field site G. affinis persists as an annual species with relatively high growth rates and corresponding reproductive patterns, whereas G. nobilis females have a slower reproductive tempo and may live multiple years.     

Reproductive variation and adult size in two co-occurring grasshopper species

ABSTRACT. 1. Egg sizes and clutch sizes of the grasshoppers Chorthippus brunneus (Thunb.) and Myrmeleotettix maculatus (Thunb.) were compared among three years and among three sites less than 1.3 km apart. Relationships between these reproductive traits and date of egg laying, body size and body condition were sought.
2. M.maculatus , the smaller species, laid fewer but larger eggs; and only the eggs of this species showed significant differences between sites and years.
3. A negative correlation between egg size and number per clutch was evident between species and years, but generally not among sites and among individuals of a population.
4. However, a hidden negative correlation between egg size and number was uncovered within populations when the relationship was examined for females of a given mature weight.
5. Variation in the number of eggs per clutch was explained statistically by a positive relationship between female body weight and egg number. Also, both interpopulation and intrapopulation comparisons revealed that for M.maculatus , but not for C.brunneus , females with long hind femurs laid large eggs.     

Shallow males, deep females: sex-biased differences in habitat distribution of the freshwater calanoid copepod Arctodiaptomus alpinus

In a small, circular high-altitude karst lake situated in the northeastern calcareous Alps of Austria the copepod Arctodiaptomus alpinus (Imhof) develops each spring from resting eggs after the lake basin fills in early spring. After mid-summer, late developmental stages and adults concentrate in a thin layer near the sediment throughout the lake basin. During daytime an average 86% and 71% of female and male copepods respectively resided near the bottom. Males exhibited stronger diel vertical migrations than females and part of the population concentrated in near-surface water at night, but the majority of copepods remained at the sediment during darkness. Macro-photographs and core samples revealed that the proportion of epibenthic male A. alpinus decreased constantly from >70% in 3 m depth to <20% below the 8-m isobath. The proportion of ovigerous females increased with depth, whereas non-ovigerous females were comparatively more abundant in shallow water. Concurrently the highest frequencies of copulating copepods were detected in areas of high male density. Above the 3-m isobath male A. alpinus formed dense swarms in scattered patches of the macrophyte Ranunculus eradicatus (>93% males; maximum abundance: 362 600 individuals m⁻²). Adults and larvae of the alpine newt Triturus alpestris are the top predators in this fishless alpine lake, and A. alpinus constitutes a major component of their diet. However, predator densities and different predation rates on male and female copepods were not sufficient to explain the observed horizontal distribution of A. alpinus . We argue that the optimization of successful sexual encounters in copepods is the ultimate driving force behind this segregation of both sexes of A. alpinus ('mate encounter hypothesis').     

Competition between the nymphs of two regionally co-occurring species of Notonecta (Hemiptera: Notonectidae)

1. Two species of freshwater invertebrate predator, Notonecta maculata and N. obliqua , showed a negative association in a series of small, man-made ponds in the Peak National Park, Derbyshire, U.K. The present study examines the potential role of interspecific interactions among nymphs on this regional distribution pattern.
2. The survival, development and feeding efficiency of nymphs were examined in laboratory and field mesocosm experiments with intra- and interspecific competition and contrasting environmental complexity.
3. Survival to adulthood and mean lifespan varied significantly in interspecific competition treatments in both laboratory and field experiments, with N . maculata showing higher survival in the simple environment and N. obliqua higher survival in the complex environment.
4. Variations in feeding efficiency were consistent with the survival trends: N. maculata had a higher efficiency in the simple environment, whereas N. obliqua had greater efficiency in the complex environment. There was evidence of a developmental response in feeding efficiency, with differences between species increasing with age.
5. These results suggest that the relative competitive abilities of the two species are affected by habitat complexity, and that competition between species may modify the species distribution where they co-occur.     

Reproductive biology and female frequencies of two co-occurring gynodioecious Thymus species

Plant Ecology - Gynodioecy, the co-occurrence of female and hermaphrodite plants in the same population is relatively common in the genus Thymus, where a high and variable female frequency has been...     

Divergent landscape effects on population connectivity in two co-occurring amphibian species

The physical and environmental attributes of landscapes often shape patterns of population connectivity by influencing dispersal and gene flow. Landscape effects on movement are typically evaluated for single species. However, inferences from multiple species are required for multi‐species management strategies increasingly being applied in conservation. In this study, I compared the spatial genetic patterns of two amphibian species across the northeastern United States and estimated the influence of specific landscape features on the observed genetic structure. The spotted salamander (Ambystoma maculatum) and wood frog (Rana sylvatica) share many ecological attributes related to habitat use, phenology and site fidelity. However, I hypothesized that important differences in their movement patterns and life history would create distinct genetic patterns for each species. Using 14 microsatellite loci, I tested for differences in the level of genetic differentiation between the two species across 22 breeding ponds. The effects of eight landscape features were also estimated by evaluating 32 landscape resistance models. Spotted salamanders exhibited significantly higher genetic differentiation than wood frogs. Different landscape features were also identified as potential drivers of the genetic patterns in each species, with little overlap in model support between species. Collectively, these results provide strong evidence that these two amphibian species interact with the landscape in measurably different ways. The distinct genetic patterns observed are consistent with key differences in movement ability and life history between A. maculatum and R. sylvatica. These results highlight the importance of considering more than one species when assessing the impacts of the landscape matrix on population connectivity, even for ecologically similar species within the same habitats.     

Naupliar growth versus egg production in the calanoid copepod Centropages typicus

Growth rates of the copepod Centropages typicus were assessedduring May 1998 in the Alboran Sea. The goal of the work wasto compare field growth rates of juvenile (artificial cohortmethod) and adult copepods (female egg production rates) underan extensive range of natural conditions. The results showedthat adult and juvenile growth rates were similar in some stations,while in others no such relationship was apparent. In contrastto the common belief that juvenile growth may be saturated undernatural conditions whereas adults may be food-limited, in mostof our study, adult growth rates were greater than juveniles.We discuss these results in the light of food availability atthe surveyed stations. In summary, the assumption that copepodgrowth rates estimated through egg production rates of adultsare equivalent to juvenile growth is not always valid.     

Plastic and constant developmental traits contribute to adaptive differences in co-occurring Polygonum species

Adaptive differences among species are often thought to result from developmentally constant trait differences that enhance fitness in alternative environments. Species differences in patterns of individual phenotypic plasticity can also have ecological consequences. Indeed, functionally related constant and plastic traits may interact to determine the phenotype's adaptive value in particular conditions. We compared juvenile shade avoidance traits (height and its components, internode length and node number) across two field density treatments in Polygonumpersicaria and P. hydropiper, annual plant species that co‐occur in pastures comprised of a mosaic of plant densities. We used selection analyses to test trait contributions to fitness in alternative density treatments. Seedlings of both species expressed plasticity for internode elongation in response to density; P. persicaria plants increased internode length and consequently height significantly more in high density than did those of P. hydropiper. As predicted by the shade avoidance hypothesis, increased height was adaptive for both species in high density stands, so P. persicaria plants had higher fitness in this environment. By contrast, node numbers were relatively constant across density treatments in both species: P. hydropiper seedlings consistently produced more nodes than did those of P. persicaria. This constant trait difference contributed to P. hydropiper's greater relative fitness at low density, where more nodes and hence leaves enable plants to better exploit available light. Differences between species in these juvenile shade‐avoidance traits did not result from the evolutionary constraints of lack of heritable variation or costs of plasticity. We discuss how these interspecific trait differences may have been generated by divergent selective histories resulting from differences in herbivore resistance. These results illustrate how adaptive differences in both plastic and constantly expressed traits may jointly contribute to ecological distribution, including coexistence in patchy habitats.     

Characteristics of digestive enzymes of calanoid copepod species from different latitudes in relation to temperature, pH and food

In calanoid copepods it is poorly understood how enzymatic activities and patterns are affected by abiotic and biotic factors. Such knowledge, however, is crucial to assess metabolic functioning and performance of organisms in different habitats. Therefore, our study focuses on digestive enzyme activities in relation to temperature, pH and food in the Arctic species Calanus glacialis and in Centropages hamatus and Temora longicornis from the North Sea. Enzyme activities were measured over a range from 0 to 70 °C (lipases/esterases, proteinases) and pH 5 to 9 (proteinases). In all species, relative proteinases activity peaked at 40/50 °C and pH 6; relative lipases/esterases activity peaked at 30 °C. Between 0 and 20 °C, lipase activity of C. glacialis was higher (40-70% of maximum) than that of the boreal copepods (25-64%), which suggests thermal adaptation of the lipid metabolism in the polar species. Incubating C. glacialis with the diatom Thalassiosira weissflogii showed (i) that enzyme activities increased especially in the alkaline range and (ii) that enzyme patterns, revealed by gel electrophoresis, differed from that of starving individuals, indicating that feeding induced enzyme expression. Such studies, linking abiotic and biotic conditions to enzyme functioning, can help elucidating the capacity of copepods to respond to environmental changes.     

Dissolved solids do not induce diapause in the calanoid copepod Aglaodiaptomus leptopus

The effect of different total dissolved solid levels (TDS) was investigated as a predictor of diapause induction in the calanoid copepod Aglaodiaptomus leptopus. We collected adult copepods in the field and monitored seasonal changes in diapause egg production. We determined total dissolved solid levels, conductivity and temperature values from five ponds. In experimental work, females bearing clutches, collected from the field, were monitored under laboratory conditions to determine the extent of diapause induced. Females without clutches were placed with males and then subjected to low, medium and high TDS levels. After six days, females bearing clutches were isolated to determine the proportion of clutches that were diapausing. The relationship between TDS and food levels was calculated using Williamson and Butlers food limitation index (f). We concluded that neither TDS nor temperature are predictors of the induction of diapause in this study. The obtained results suggest, however, that density possibly plays a role in predicting diapause induction.