Nitrogen, Phosphorus, and Potassium Nutrition of the Edaphic Ecotypes in Cynodon dactylon (L.) Pers.

A differential response to nitrogen, phosphorus, and potassiumwas observed for the three edaphic ecotypes in Cynodon dactylon(L.) Pers. naturally occurring at Malla, Shiwalik, and Chandigarh.The soils of these sites differ markedly with respect to availablenutrients. The better growth of the Malla population comparedto the Chandigarh population in low potassium and low nitrogenmedia could be related to the low availability of these nutrientsin the calcareous Malla soil. The Shiwalik population tendedto show an intermediate behaviour. Whilst the Malla populationwas susceptible to ‘phosphate toxicity’, the growthyield of the Shiwalik and the Chandigarh populations improvedwith increase in phosphorus in the medium. The three populationsalso differed from each other with respect to the uptake ofvarious nutrients. These results could be correlated with thenatural habitat conditions in which the three ecotypes occur.It was concluded that besides calcium which was shown to bepartly responsible for the restriction of the three ecotypesto their respective habitats, the availability of nitrogen,potassium, and also phosphorus may play an important role.     

A comparison of some populations of Eucalyptus viminalis Labill. growing on calcareous and acid soils in Victoria,Australia

A study of populations of Eucalyptus viminalis found on both acid and alkaline soils showed that seedlings differ in their tolerance of calcareous soils and their susceptibility to lime chlorosis. Seedlings from an open-forest population on calcareous dunes in the Otway Ranges, Victoria, averaged a significantly greater yield than seedlings of three other populations found on acid soils when all were grown on calcareous soil of pH 6.8. A tall ribbon gum form of E. viminalis, from Paradise gully in the Otways, was the least tolerant of alkaline conditions and showed severe signs of chlorosis and reddening of leaf margins. The latter appeared to be related to high uptake of phosphorus. This population showed rapid growth on a fertile acid loam. The data presented also emphasize the need to consider the variation in soils which can occur at any one site. Population differences were most marked when seedlings were grown on soils of pH near 7.0. Yield was reduced in the Otways calcareous population when seedlings were grown on the more leached soil from swales (pH 6.5) between dunes and on the highly calcareous soil from the dune crests (pH 7.8–8.0). The failure of any of the populations of E. viminalis studied to grow well on soils of high pH suggest that this species is not tolerant of highly calcareous soils. This may in part explain the absence of this species from the drier calcareous areas on the Mornington Peninsula and the Yanakie Isthmus, Victoria. Since seedlings had reduced root development on the soil collected from the Peninsula, E. viminalis may be restricted, in effect because of drought stress, to non-calcareous sites in the eastern part of Victoria.     

The effect of high pH and P on the development of lime-chlorosis in two seedling populations ofEucalyptus obliqua L'Hérit

Summary Glasshouse experiments have shown that the application of an acidulating agent to a calcareous soil can increase growth and alleviate severe chlorosis in an acidic population ofE. obliqua. In contrast, a calcareous population showed only a slight response to this treatment and maintained adequate growth and a low frequency of chlorosis on both control and treated calcareous soils. Foliar analyses of seedlings of the acidic population showed that alleviation of chlorosis was concomitant with a reduction in the levels of P, Ca and K, and an increase in uptake of Fe. However, the total Fe content of foliage was poorly correlated with the occurrence of severe chlorosis. Although this evidence suggested that the differential susceptibility ofE. obliqua to lime-chlorosis can be reduced by increasing the availability of Fe, the greater concentration of Fe in chlorotic seedlings indicated that lime-chlorosis may also be due to an inactivation of Fe within the plant (i.e. by P). This hypothesis was partly confirmed by a water culture experiment which showed that a combination of relatively high pH and high external levels of P could induce severe chlorosis in seedlings of the acidic population. In contrast, it appears that the calcareous population has a more efficient mechanism for absorbing Fe and holding it in an available form, even when external concentrations of P are high. It is suggested that plants which have an efficient mechanism for the uptake of Fe at relatively high pH and are less susceptible to the detrimental effects of P have been selected for on these alkaline calcareous soils.     

Influence of calcareous soil on Cry3Bb1 expression and efficacy in the field

Greater than expected injury by western corn rootworm (WCR) (Diabrotica virgifera virgifera LeConte) to Cry3Bb1 expressing maize hybrids (Zea mays L.) has been reported in southwestern Nebraska. Affected areas of some fields are often associated with high pH calcareous soils where maize growth is poor and iron chlorosis is common. As part of a comprehensive study to understand potential causes of unexpected injury, experiments were conducted during 2013 and 2014 to ascertain whether the calcareous soil conditions and associated poor maize growth negatively affect the expression of Cry3Bb1. Quantitative determination of Cry3Bb1 protein expression levels in root tissues was carried out on plants at V5–V6 growth stage using the enzyme-linked immunosorbent assay. Cry3Bb1 and non-Bt near isoline maize hybrids were artificially infested with Cry3Bb1-susceptible WCR eggs to measure survival and efficacy of Cry3Bb1 maize in calcareous and non-calcareous soils. Results showed that there was not a significant difference in expression of Cry3Bb1 protein between plants from calcareous and non-calcareous soils (18.9–21.2 µg/g fresh weight). Western corn rootworm survival was about sevenfold greater from the non-Bt isoline than Cry3Bb1 maize indicating that Cry3Bb1 performed as expected when infested with a Cry3Bb1 susceptible rootworm population. When survival from calcareous and non-calcareous soils was compared, no significant differences were observed in each soil. A significant positive correlation between soil pH and expression of Cry3Bb1 protein in roots was detected from samples collected in 2014 but not in 2013. No such correlation was found between soil pH and survival of WCR. Results suggest that Cry3Bb1 expression levels were sufficient to provide adequate root protection against WCR regardless of soil environment, indicating that lowered Cry3Bb1 expression is not a contributing factor to the greater than expected WCR injury observed in some southwestern Nebraska maize fields.     

Mobility, solubility and lability of fluid and granular forms of P fertiliser in calcareous and non-calcareous soils under laboratory conditions

Despite a long history of application of phosphorus fertilisers, P deficiency is still a major limitation to crop production on calcareous soils. Recent field research conducted in highly calcareous soils in southern Australia has demonstrated that both grain yield and P uptake of wheat (Triticum aestivum L.) is greater when fluid forms of P are used compared to granular forms. To improve our understanding of the mechanisms underlying this response to P in the field, we compared the lability, solubility and mobility of P applied as either a fluid (3 products) or granular (3 products) form to two calcareous and one alkaline non-calcareous soils in the laboratory. Over a five-week period, between 9.5 and 18 % of the P initially present in the fertiliser granules did not diffuse into the surrounding soil. The degree of granule dissolution was independent of the soil type. In contrast, P solubility, lability and diffusion were significantly greater when fluid products were applied to the calcareous soils, but not to the alkaline non-calcareous soil. These findings are discussed in relation to field trials results where fluid products outperformed granular fertilisers.     

Influence of poultry manure on phosphorus availability and the standard phosphate requirement of crop estimated from quantity-intensity relationships in different soils

Twelve samples comprising of non-calcareous, calcareous and acidic soils were used to study P availability parameters and standard phosphate requirement (SPR) in the presence and absence of poultry manure (PM). Adsorption of P and phosphate potential decreased while Olsen extractable P increased by PM addition, the extent varying with the type of soil. The calculated SPR to attain soil solution concentration of 0.2 microg P ml(-1) was invariably higher in fine textured soils. Acid and calcareous soils registered higher SPR compared to non-calcareous soils. PM addition lowered the SPR in all the soils. The observed P supplementation through PM ranged from 11.6% to 100% signifying no fertilizer P need in the latter. Calibration curves were drawn for calculating the SPR for a given soil solution P concentration without and with the addition of PM.     

Copper uptake and phytotoxicity as assessed in situ for durum wheat (Triticum turgidum durum L.) cultivated in Cu-contaminated, former vineyard soils

This work assessed in situ, copper (Cu) uptake and phytotoxicity for durum wheat (Triticum turgidum durum L.) cropped in a range of Cu-contaminated, former vineyard soils (pH 4.2–7.8 and total Cu concentration 32–1,030 mg Cu kg⁻¹) and identified the underlying soil chemical properties and related root-induced chemical changes in the rhizosphere. Copper concentrations in plants were significantly and positively correlated to soil Cu concentration (total and EDTA). In addition, Cu concentration in roots which was positively correlated to soil pH tended to be larger in calcareous soils than in non-calcareous soils. Symptoms of Cu phytotoxicity (interveinal chlorosis) were observed in some calcareous soils. Iron (Fe)–Cu antagonism was found in calcareous soils. Rhizosphere alkalisation in the most acidic soils was related to decreased CaCl₂-extractable Cu. Conversely, water-extractable Cu increased in the rhizosphere of both non-calcareous and calcareous soils. This work suggests that plant Cu uptake and risks of Cu phytotoxicity in situ might be greater in calcareous soils due to interaction with Fe nutrition. Larger water extractability of Cu in the rhizosphere might relate to greater Cu uptake in plants exhibiting Cu phytotoxic symptoms.     

Phylogeny of calcareous dinoflagellates as inferred from ITS and ribosomal sequence data

The phylogenetic relationships of calcareous dinoflagellates (i.e., Calciodinellaceae and Thoracosphaera) are investigated. Molecular data from the ribosomal 5.8S rRNA and highly conserved motifs of the ITS1 show Calciodinellaceae s.l. to be monophyletic when few non-calcareous taxa are included. They segregate into three monophyletic assemblages in a molecular analysis that considers the 5.8S rRNA and both the Internal Transcribed Spacer regions ITS1 and ITS2: a clade comprising species of Ensiculifera and Pentapharsodinium (E/P-clade), Scrippsiella s.l. (including fossil-based taxa such as Calciodinellum and Calcigonellum), and a heterogeneous group (T/P-clade) of calcareous (e.g., Thoracosphaera) and non-calcareous taxa (e.g., the highly toxic Pfiesteria). The potential to produce calcareous structures is considered as apomorphic within alveolates, and non-calcareous taxa nesting with calcareous dinoflagellates may have reduced calcification secondarily. Molecular results do not contradict general evolutionary scenarios provided by previous morphological (mainly paleontological) investigations.     

Mycorrhizal infection indicates the suitability of different management treatments for nature conservation in calcareous grassland

The vegetation composition of calcareous grasslands in Central Europe is mainly determined by the applied management treatments and the nutrient status of the ecosystems, but these factors cannot clearly be separated. Recently several authors reported a shift in growth-limiting plant nutrients from nitrogen to phosphate. Therefore, we tested the interaction between management and plant available phosphate in a long-term experiment in Southern Germany with different management treatments applied to calcareous grassland. We measured mycorrhization rates, which are assumed to be high at low phosphate availability, to explore the interaction between management, phosphate availability and vegetation composition. By correlating mycorrhization rates to vegetation and soil data we found that treatments involving nutrient removal show a greater degree of mycorrhization than treatments leading to nutrient enrichment. According to decreasing suitability for grassland conservation, the six different management treatments could be ranked as mowing, followed by mulching (twice a year, once a year and every second year), burning and undisturbed succession. It was shown, that mycorrhizal infection rates (1) can be used to evaluate different management treatments for their suitability to conserve the initial vegetation composition, and (2) may provide information about the processes of adaptation to the current management, namely to differences in plant available phosphate. However, as discussed for the undisturbed succession treatment, it might not be sufficient to look at the degree of mycorrhization only. These are related to phosphate availability, but to detect adaptation processes to management, the degree of mycorrhization should be analysed together with plant’s internal nutrient relocation patterns.     

Improving fertiliser efficiency on calcareous and alkaline soils with fluid sources of P,N and Zn

Alkaline calcareous or sodic soils represent an important proportion of the world's arable soils and are important for cereal production. For calcareous soils in general, despite high applications of P fertiliser for many years, P deficiency in cereals is common. Field experiments were conducted to test the relative ability of granular (e.g. DAP, MAP and TSP) and fluid fertilisers to supply P to wheat on grey calcareous and red brown calcareous sandy loam soils (Calcixerollic xerochrepts). A pot experiment was also conducted with these soils and with two non-calcareous alkaline soils to investigate the effects of placement on the efficiency of fertiliser performance. In 1998, fluid and granular sources of P, N and Zn were compared in the field by banding below the seed at sowing. In 1999, MAP applied as granular, and technical grade MAP applied as fluid, were compared as sources of P in rate response experiments. First year results showed that fluid sources of P, N and Zn produced significantly more grain than the granular product. In the following year, fluid fertilisers were found to produce significantly higher response curves for shoot dry weight, grain yield and P uptake in grain. At a commercial rate of 8 kg P ha^–1, fluid fertiliser produced between 22% and 27% more grain than the granular product. Soil moisture and fertiliser placement effects are implicated in the higher efficiency of fluid fertilisers.     

A comparative study of soil formation in primary stands of Scots pine (planted) and poplar (natural) on calcareous dune sands in the Netherlands

Initial soil formation under primary stands of Scots pine (planted) and European black poplar (natural) on calcareous dune sands was studied, paying particular attention to the humus forms and their spatial variability. The stands studied are both about 80 years old and are situated, at close distance, in the coastal dunes near Wassenaar (the Netherlands).Under Scots pine, soils with a mor-type humus form were observed, exhibiting slight podzolisation. Soil variability is rather slight and soil development is comparable to that under primary Scots pine stands on non-calcareous inland drift sands. Under poplar, mull-type humus forms occur which tend towards moder and exhibit a markedly stronger litter decomposition and bioturbation. In contrast to the soils under pine, soil variability is considerable. Results from chemical analyses of two representative soil profiles are in conformance with these trends.It is concluded that the observed trends in soil formation are in line with those described in the literature, and that a period of 80 years is sufficient for a strong vegetation related divergence in soil properties. Soil variability within the stands probably results from redistribution of litter by wind and/or gravity and will be rather site-dependent.     

The ‘masking effect’ of silviculture on substrate-induced plant diversity in oak-hornbeam forests from northern France

Traditional measures of plant diversity from phytosociological relevés are used to compare floristical richness and diversity between different oak-hornbeam forest stands of Thiérache (Northern France). Five sets of physiognomically similar forests are distinguished according to the geological substrate: schists and sandstones, loess on schists, loess on calcareous rocks, jurassic limestones, cretaceous marls and clays. Plant species diversity is investigated at two scale levels: a phytocoenotic level and a synusial one which is supposed to partly avoid a possible silviculture-related bias. Two pools of sets may be distinguished: a calcareous and a non-calcareous one. The calcareous shows the highest floristical diversity, and the non-calcareous one the highest phytosociological richness. For each inter-pool pair of sets, plant species diversity is significantly different at the phytocoenotic level as well as at the level of herbaceous or shrubby synusiae, but not at the arborescent one. Through the coppice-with-standards practice, silviculture was the main determining factor for arborescent synusiae while the herbaceous and shrubby synusiae, which support the major part of the diversity, remained strongly influenced by the substrate. It is suggested that silviculture has got a masking effect on plant species diversity within oak-hornbeam managed forests by making uniform tree layers that outshine the understory vegetation diversity.     

Density-dependent inhibition of expression of syncytiotrophoblastic markers by cultured human choriocarcinoma (BeWo) cells

In the presence of methotrexate, cultured human choriocarcinoma (BeWo) cells undergo a differentiative response that resembles normal trophoblastic development. In the current study, the effects of cell number and population density on drug-induced conversion of BeWo cells from the cytotrophoblastlike to the syncytiotrophoblastlike phenotype were investigated using as markers of differentiation formation of "giant" cells, a process shown to require exogenous purines, and expression of placental (heat-stable) alkaline phosphatase. Giant cell formation, assessed by determination of cell volumes, was reduced in crowded cultures, and addition of hypoxanthine to growth media partially restored methotrexate-induced cell enlargement. Cellular uptake of methotrexate, assessed by following the loss of methotrexate from cell culture fluids during drug exposures, was two-threefold greater in sparsely populated than in densely populated cultures. Although the concentration of methotrexate in culture fluids of crowded cultures declined during exposures of 48 hr, the amount of extracellular drug remaining at 48 hr was well above the threshold for induction of the differentiative response. When culture population was held constant and population density was manipulated by varying the substratum available to cells, methotrexate-induced cell enlargement was inversely related to population density. Expression of placental alkaline phosphatase, salvage of exogenous hypoxanthine, and synthesis of RNA were also reduced at high population densities. These results indicate that expression of markers of methotrexate-induced differentiation of BeWo cells was inhibited in a density-dependent manner that may have been related to reduced cellular uptake of the inducing agent and of exogenous nutrients (purines) from culture fluids.     

The effect of calcium on the germination,growth and mineral nutrition of acidic and calcareous populations ofEucalyptus obliqua L'Hérit

Summary A comparative study of two populations ofE. obliqua has shown that germination of an acidic population is severely reduced at high levels of Ca, whereas a calcareous population maintained high germinability over a wide range of Ca concentrations. In contrast, seedlings of both populations showed a similar overall response in yield to an increasing supply of external Ca. However, plants from the acidic population tended to show a lower survival rate and poorly growing plants accumulated higher levels of Ca and P at high external concentrations of Ca than the calcareous population. Plants showing severe chlorosis and symptoms of P and Ca toxicity also exhibited low K/Ca ratios.     

The role of calcium in buffering soils

Abstract. Calcium is the soluble cation that occurs in largest amount in most soils. It does not take part directly in the proton transfer reactions involved in pH-buffering, but it provides the cation charge balance for these reactions. It is also the complementary cation in formulations of chemical potential for many other ions in soils. The presence of free calcium carbonate in calcareous soils. The presence of free calcium carbonate in calcareous soils ensures a very high soil buffer capacity; d AB/ d pH ≃ 1000 Eq. m⁻³.
In acid mineral soils, dissolution and precipitation of aluminium ions contribute to the buffering processes, but most of the buffering in non-calcareous soils is caused by specific ion adsorption at variable-charge sites, in particular those associated with the dissociation of humus acids. Typical buffer capacity values of non-calcareous soils vary from 10 Eq. m⁻³ for sandy soils to 100 Eq. m⁻³ for peats. The pH changes associated with buffering are produced by leaching of calcium from soil, or by adding calcium to soil in liming materials.     

Why barren intervals? A taphonomic case study of the Scandinavian Alum Shale and its faunas

The Scandinavian Alum Shale Formation (Middle Cambrian to Lower Ordovician) accumulated under generally low oxygen concentrations. Syndepositional changes in the oxygen concentrations of the bottom water are reconstructed on the basis of the fossil fauna. Under relatively high oxygen concentrations, brachiopods and non-olenid polymerid trilobites inhabited the sea floor. Under lower oxygen concentrations the fauna was dominated by agnostids and at lowest oxygen levels by olenid trilobites. The enrichments of vanadium relative to nickel, as well as the enrichment of sulphur, match these faunal changes. A geochemical classification of the dysoxic environment is presented. The abundance of calcareous fossils decreases with increasing bottom water oxygen concentrations, indicating that the preservation of calcareous hard parts is most likely at the lowest oxygen concentrations. The poor preservation of calcareous fossils at relatively high oxygen concentrations is explained by the generation of corrosive pore waters during the reoxidation of sulphide compounds. Trilobitic and non-trilobitic intervals alternate up through the shale. Non-trilobitic intervals are either barren or contain non-calcareous fossils (phosphatic brachiopods, phosphatic 'ostracodes' and graptolites). The stratigraphical variation of trilobitic and non-trilobitic intervals is interpreted to reflect major changes in oxygen levels that might be linked to sea-level and climatic fluctuations.     

Regulation of gut function varies with life-history traits in chuckwallas (Sauromalus obesus: Iguanidae)

We examined the effects of hibernation and fasting on intestinal glucose and proline uptake rates of chuckwallas (Sauromalus obesus) and on the size of organs directly or indirectly related to digestion. These lizards show geographic variation in body size and growth rate that parallels an elevational gradient in our study area. At low elevation, food is available only for a short time during the spring; at high elevation, food may also be available during summer and autumn, depending on rainfall conditions in a given year. We hypothesized that low-elevation lizards with a short season of food availability would show more pronounced regulation of gut size and function than high-elevation lizards with prolonged or bimodal food availability. Hibernating lizards from both elevations had significantly lower uptake rates per milligram intestine for both nutrients, and lower small intestine mass, than active lizards. The combination of these two effects resulted in significantly lower total nutrient uptake in hibernating animals compared to active ones. The stomach, large intestine, and cecum showed lower masses in hibernators, but these results were not statistically significant. The heart, kidney, and liver showed no difference in mass between hibernating and nonhibernating animals. Lizards from low elevations with a short growing season also showed a greater increase in both uptake rates and small intestine mass from the hibernating to the active state, compared to those from high elevations with longer growing seasons. Thus, compared to those from long growing season areas, lizards from short growing season areas have equal uptake capacity during hibernation but much higher uptake capacity while active and feeding. This pattern of regulation of gut function may or may not be an adaptive response, but it is consistent with variation in life-history characteristics among populations. In areas with a short season, those lizards that can extract nutrients quickly and then reduce the gut will be favored; in areas where food may be available later in the year, those lizards that maintain an active gut would be favored. While other researchers have found much greater magnitudes of gut regulation when making comparisons among species, we find the different patterns of change in gut function between different populations of chuckwallas particularly intriguing because they occur within a single species.     

Identifying reference benthic diatom communities in an agricultural watershed (Guadiana River,SW Spain)

Determining the ecological status of river ecosystems should be based on reference (near-natural) conditions and on the use of their biological communities. Identifying reference conditions is particularly difficult in largely disturbed watersheds, such as those of agricultural landscapes. Ecotype reference sites and their characteristic diatom communities were identified in the Guadiana watershed. Three different ecotypes were defined: rivers with siliceous bedrocks, upper watershed calcareous rivers, and mainland watercourses. Thirteen key stressors divided in four categories (channel alteration, hydrological pressure, pollution pressure, and river habitat pressure) were used to select reference sites in each ecotype. Multivariate techniques were used to describe type-specific reference diatom community in each ecotype. Because of the difficulty to define reference sites in the mainland watercourses, some sites were selected under the criterion of minimum impacted sites. Diatoms inhabiting sites included species tolerant to high nutrient concentration and mineralized waters, like Nitzschia inconspicua, Gomphonema parvulum, and Navicula veneta. These taxa defined the best condition (near-to-reference) in an ecotype highly impacted by human activities, the best possible in the long-lasting impaired conditions in the large river watercourses in agricultural landscapes.     

Soil nutrient supply in natural and managed forests

The effect of a laboratory addition of 10, 100 and 500 mg Cd kg_{dry soil} ^-1 on ammonification and nitrification was studied using soil samples of two unpolluted grassland soils. Calcareous and non-calcareous soil were selected for this purpose. Various parameters of nitrifying activity were investigated simulataneously: activity during long-term laboratory incubations in the presence and absence of a substrate, mineralization potentials, and potential activity of both ammonium and nitrite oxidizers during short-term incubations in soil slurries. Cadmium was added as aqueous CdCl₂.Additions of both 100 and 500 mg Cd kg_{dry soil} ^-1 doses significantly lowered the ability of both soils to nitrify 100 g added NH₄ ⁺-N g_{dry soil} ^-1 as a substrate, which was reflected in a decreased rate of nitrate formation (maximum inhibition reached 60% in the calcareous soil and 45% in the non-calcareous soil). Furthermore, these two concentrations of Cd caused an abnormal accumulation of nitrite immediately after incorporation, particularly in the calcareous soil. The addition of 10 mg Cd kg_{dry soil} ^-1 intensified N-mineralization in both soils, probably as a consequence of a higher concentration of readily metabolized substrate originating from killed bacteria or fungi. An excess of nitrate was then formed as a final step. The harmful effect of cadmium was more pronounced in calcareous soil, probably due to the higher sensitivity of nitrite-oxidizers in these soil samples.     

RESOURCE INFLUENCED SEX ROLES OF ZAPROCHILINE TETTIGONIIDS (ORTHOPTERA: TETTIGONIIDAE)

Male tettigoniids donate nutrients to females at mating in the form of a spermatophylax. Male-donated nutrients function as paternal investment leading to a reversal in the sex roles of males and females. Reversal in the behavioral sex roles of a zaphrochiline tettigoniid was found to be directly related to the current availability of food resources in the environment. When resource availability was low, females were less fecund and males had lower and more variable accessory gland weights (the spermatophylax producing gland) than when resource availability was high. When resources were scarce, larger individuals had a reproductive advantage having more eggs or heavier accessory glands. All individuals were equally fecund or had equal accessory gland weights when resources were plentiful. During low resource availability males that had a spermatophylax to offer were choosy of their mates, and females were competitive. When resources were plentiful, males were less discriminative and females showed signs of discrimination. There was evidence that female sexual motivation decreased when environmental resources were plentiful (as indicated by mating frequency) and that only females of low nutritional status continued to mate. When resources were scarce, females achieved fecundities equivalent to those achieved during high resource availability through spermatophylax consumption. These data support the hypothesis that females continue to mate to compensate for low resource availability and that male parental investment may be important only in poor quality habitats. When resources are plentiful females do not need to remate. Thus resource availability may directly influence the number of sexually active males and females (operational sex ratio) and the form of sexual selection.