MHC, health, color, and reproductive success in sand lizards

Good genes are genetic elements that contribute to lifetime reproductive success, regardless of an individuals additional genotype. Their existence is debated, and most work has targeted their viability benefits to the offspring of choosy females. In the present study, we analyze a case of potential good genes effects in adult male sand lizards (Lacerta agilis). We show that males with a particular RFLP (Restriction Fragment Length Polymorphism) MHC genotype (O-males), as opposed to those that lack this genetic element (NO-males), have less ectoparasites under increasing physiological stress (indexed by baseline corticosterone level), and are not constrained by parasites at production of status coloration. Furthermore, O-males are more successful at mate acquisition and guard their partners longer. Ultimately, they have a higher genetic reproductive success as assigned by microsatellites.Communicated by W. Cooper     

Soil Lead at Elementary Public Schools: Comparison Between School Properties and Residential Neighbourhoods of New Orleans

Soil studies, conducted in Maryland, Minnesota and Louisiana, have described the urban pattern of lead contamination. They have shown that the highest amounts of lead cluster within the interior of the largest cities. The results of the New Orleans urban patterns of distribution of soil lead provided the basis for further study. The hypothesis was tested that elementary school properties have the same pattern of soil lead contamination as their neighbouring residential communities. Thirty New Orleans Public Elementary Schools were selected for this study. Surface samples (2.5cm or 1 inch depth) were collected from playgrounds and next to entrances of each school. Results showed that soil lead on school properties follows the same relative contamination patterns (pvalue10^–5) as soil lead on residential properties of neighbouring communities. Schools however, have significantly lower lead contamination than the neighbouring residential properties. Innercity school properties present a higher risk of soil lead exposure than mid and outercity schools. Soils next to innercity school entrances showed the highest lead, with 18.5% having concentrations over 400gg^–1. Systematic landscaping around the school entrances would significantly reduce the hazard from lead dust contaminated soils.     

Sorption of the natural endocrine disruptors,oestrone and 17 beta-oestradiol in the aquatic environment

Adsorption of hydrophobic contaminants at the particle/water interface is one of the key processes controlling their fate in the aquatic environment. The sorption of the natural female hormones oestrone and 17-oestradiol has been studied under simulated riverine conditions. Both the kinetics and the effects of varying fundamental environmental parameters (e.g. sediment properties) on the thermodynamic equilibrium partition coefficient (K _p) have been studied in continuous and batch sorption experiments, respectively. Results showed that the sorption of oestrone and 17-oestradiol by sediment was relatively slow, reaching equilibrium in 50 days. In addition, relatively small adsorption of both oestrone and 17-oestradiol onto the sediment was observed, with K _p values between 200 and 250 mL g^–1. The comparable K _p values of the two compounds reflect their structural similarity. It can be concluded that the two endocrine disruptors, oestrone and 17-oestradiol remain primarily in association with the aqueous phase.     

Variation of leaf gland volatile oil within a population of sweet gale (Myrica gale) (Myricaceae)

Summary The leaf gland volatile oils of ten sweet gale plants from a Scottish population were extracted in early summer. The results differed notably from reports of other populations in respect of the sesquiterpenes, -elemenone and germacrone, which were major components of the volatile oil. Three dihydrochalcones were also detected in the volatile oil. Variation within the population existed, particularly with respect to the relative importance of germacrone. Five plants were resampled in late summer and exhibited a marked reduction in -elemenone, a lesser reduction in germacrone and changes in the proportions of some monoterpenes.     

Female choice on male quantitative traits in lizards — why is it so rare?

Female choice on the basis of male traits has been described in an array of taxa but has rarely been demonstrated in reptiles. In the sand lizard (Lacerta agilis), and possibly in other non-territorial reptiles, a male's contribution to a female's fitness is restricted to his genes. In order to choose males of high genetic quality, females have to trade the fitness gain against the costs of active choice. In a Swedish population of sand lizards, long-lived males sired offspring with higher embryonic survival compared to offspring sired by short-lived males. In spite of this female sand lizards did not mate selectively with older and/or larger males. There appeared to be mo reliable cues to male longevity; age-specific male body size was highly variable. Furthermore, estimates of male nuptial coloration did not covary with ectoparasite load and, hence, females cannot use male coloration as a cue to heritable resistance to pathogenic parasite effects. When cues to male genetic quality are poor, or inaccurate, and males make no parental investment, we predict that female choice will be rare. Sand lizard females mating with many partners lay clutches with higher hatching success. Thus, females may obtain good genes for their young by multiple mating, thereby avoiding costs associated with mate choice.     

Reproductive effort in relation to maternal social rank in reindeer (<Emphasis Type="Italic">Rangifer tarandus</Emphasis>)

In polygynous mammals, high-quality females may increase their fitness by providing superior care to their offspring. Based on the agonistic interactions of female reindeer in an experimental herd during two consecutive years (1997 and 1998), we tested whether maternal social rank influenced: (1) winter body-mass change of females, (2) preparturition reproductive effort (measured as fecundity, the birth mass and the birth date of their calves), (3) preweaning maternal effort (measured as calves preweaning mortality, early preweaning and late preweaning growth rate and September body mass of calves), and (4) postweaning maternal effort (measured as calves body-mass change during their first winter). In the models, we included September females body mass as a covariate to separate the effects of maternal rank and body mass. We also tested whether the effect of social rank on maternal efforts was dependent on offspring sex. High-ranked females gained body mass whereas low-ranked females lost weight during the winter. Fecundity was higher and date of birth was earlier in high-ranked females than in subordinates, whereas no effect of females rank on birth mass of calves was found. Early preweaning growth rate and September body mass of calves increased with increasing females social rank, whereas late preweaning daily growth rate of calves was not influenced by females rank. Calves preweaning mortality was only influenced by year, which also explained most of the variance in the winter body-mass change of calves. The effects of females rank on the reproductive-efforts parameters studied were not specific to offspring sex. These findings suggest that females rank influences reproductive effort during the preparturition, as well as the preweaning, period, the effect being sex independent.Communicated by R. Gibson     

Nutritional phagostimulants function as male courtship pheromone in the German cockroach, <Emphasis Type="Italic">Blattella germanica</Emphasis>

Summary. Males of the German cockroach, Blattella germanica, secrete a pheromonal substance from the abdominal tergal glands, which elicits a feeding response in females during the sequential courtship behavior. The nuptial secretion consists predominantly of a synergistic mixture of sugars and phospholipids. Cholesterol and a series of amino acids, which are also components of the males glandular secretion, significantly enhanced the phagostimulant activity of the sugar components. The nuptial feeding behavior of the female cockroach is therefore elicited by a complex assortment of nutritive components in the male tergal secretion, including sugars, phospholipids, cholesterol, and amino acids. These results indicate that a mixture of primary metabolites, and not of specific secondary metabolites, serves as a pheromonal cue that appeals to the females gustatory sense and effectively brings her to the precopulatory position. Although the male secretion consists of nutrients, we suggest that these compounds probably do not represent a significant nutrient investment in females and their progeny but rather function as a signal in the mating sequence of B. germanica.     

Ground gleaning in horseshoe bats: comparative evidence from<Emphasis Type="Italic"> Rhinolophus blasii</Emphasis>,<Emphasis Type="Italic"> R. euryale</Emphasis> and<Emphasis Type="Italic"> R. mehelyi</Emphasis>

The 71 species of horseshoe bat (genus Rhinolophus) use echolocation calls with long constant-frequency (CF) components to detect and localize fluttering insects which they seize in aerial captures or glean from foliage. Here we describe ground-gleaning as an additional prey-capture strategy for horseshoe bats. This study presents the first record and experimental evidence for ground-gleaning in the little-studied Blasius horseshoe bat (Rhinolophus blasii). The gleaning bouts in a flight tent included landing, quadrupedal walking and take-off from the ground. The bats emitted echolocation calls continuously during all phases of prey capture. Both spontaneously and in a choice experiment, all six individuals attacked only fluttering insects and never motionless prey. These data suggest that R. blasii performs ground-gleaning largely by relying on the same prey-detection strategy and echolocation behaviour that it and other horseshoe bats use for aerial hawking.We also studied the Mediterranean horseshoe bat (R. euryale) in the flight tent. All four individuals never gleaned prey from the ground, though they appeared to be well able to detect fluttering moths on the ground. It is not known yet whether ground-gleaning plays a role in Mehelys horseshoe bat (R. mehelyi). In a performance test, we measured the ability of these three European species of middle-sized horseshoe bats (R. euryale, R. mehelyi and R. blasii) to take-off from the ground. All were able to take flight even in a confined space; i.e. the willingness to ground-glean in R. blasii is not related to a superior take-off performance. In contrast to ground-gleaning bats of other phylogenetic lineages, R. blasii appears not to be a specialist, but rather shows a remarkable behavioural flexibility in prey-capture strategies and abilities. We suggest that the key innovation of CF echolocation paired with behavioural flexibility in foraging strategies might explain the evolutionary success of Rhinolophus as the second largest genus of bat.Communicated by T. Czeschlik     

Plant volatiles affect oviposition by codling moths

Summary. Oviposition in wild codling moth females, collected as overwintering larvae from apple, pear and walnut, was stimulated by volatiles from fruit-bearing green branches of these respective hostplants. Analysis of headspace collections showed that eight compounds present in apple, pear and walnut elicited a reliable antennal response in codling moth females: (E)--ocimene, 4,8-dimethyl-1,(E)3,7-nonatriene, (Z)3-hexenyl acetate, nonanal, -caryophyllene, germacrene D, (E,E)--farnesene, and methyl salicylate. Any one of these compounds is found in many other non-host plants, and host recognition in codling moth is thus likely encoded by a blend of volatiles. A large variation in the blend proportion of these compounds released from apple, pear and walnut suggests a considerable plasticity in the female response to host plant odours. Wild females, collected as overwintering larvae in the field, laid significantly fewer eggs in the absence of host plant volatiles. The offspring of these females, however, reared on a semi-artificial diet in the laboratory, laid as many eggs with or without plant volatile stimulus. Tests with individual females showed that this rapid change in oviposition behaviour may be explained by selection for females which oviposit in the absence of odour stimuli, rather than by preimaginal conditioning of insects when rearing them on semi-artificial diet. Oviposition bioassays using laboratory-reared females are therefore not suitable to identify the volatile compounds which stimulate egglaying in wild females.     

Descriptions and durations of premolt setal stages in female Dungeness crabs,<Emphasis Type="Italic"> Cancer magister</Emphasis>

For crustaceans with a well-defined annual molting season, such as adult female Dungeness crabs (Cancer magister Dana), setal molt staging can, in principle, be used to predict molting destiny of individual crabs. Size-specific predictions of molting probability could, in turn, be useful for estimating mortalities due to molting. Female Dungeness crabs were collected January–March 1997 at depths of 10–30 m off the California coast, USA (41°N). Intermolt stage A₁–C₄ and premolt stages D₀, D₁, D₁, D₁ and D₂ were described based on features of the branchial epipod. Laboratory experiments suggested that stage D₁ was the earliest stage beyond which eventual molting was inevitable so that molting destiny could be determined. Estimated mean number of days from the beginning of stage D₁ to molting was 85. Estimated individual stage durations, based on laboratory experiments, were 24, 51, –4, 18, and 20 days, respectively, for stages D₀, D₁, D₁, D₁, and >D₂. The estimated –4 days for stage D₁ suggests that this stage must be very brief and raises a question of its utility in designation of molt stages. Molt staging of three field samples, 300–600 crabs in each, collected prior to the 1997 annual molting season, indicated an increase in the size and frequency of crabs staged D₁ or later as the time to molt approached, although crabs of 150 mm carapace width and larger showed few signs of molt preparation. Because the duration of the molting season (approximately 120 days) for adult female C. magister in northern California exceeds the estimated maximum duration of reliable prediction of molting destiny (85 days), it does not appear that molt staging can be used to predict molting destiny in this population. However, the procedures that we have used in this paper for application to female C. magister might be used with success for other crustaceans if the duration of stages D₁ to molting exceeds the duration of the molting season, and furthermore may be used for describing temporal molting trends.Communicated by J.P. Grassle, New Brunswick     

The oral cavity and bioluminescent organs of the cardinal fish species<Emphasis Type="Italic"> Siphamia permutata</Emphasis> and<Emphasis Type="Italic"> S. cephalotes</Emphasis> (Perciformes,Apogonidae)

The mechanism of bacterial-derived bioluminescence in the apogonid species Siphamia permutata and S. cephalotes presents some special structures, particularly the existence of two sites harboring bioluminescent bacteria, and not only one as previously described. One site, is the familiar disc-like, bacteria-harboring gland and paired ventral bioluminescent reflectors, typical for this genus, that merge and end at the level of the hypobranchial region. The second and more anteriorly situated site of luminescence features two sacs that originate in the gular region and protrude into the oral cavity, via the free space left by the laterally notched tongue. The apical parts of these luminous sacs harbor the bacteria whose light diffuses within the oral cavity. The tongue surface in the studied species is unique in character among the cardinal fishes, being criss-crossed anteriorly by protruding dermal ridges, rich in taste buds and mucus-producing cells. It would appear that at night when feeding, the luminous mouth cavity of these fish acts as a lure to attract the small prey, while the ridged structure of the tongues surface facilitates their collection and aggregation before they are swallowed.Communicated by O. Kinne, Oldendorf/Luhe     

Patterns of participation and free riding in territorial conflicts among ringtailed lemurs (<Emphasis Type="Italic">Lemur catta</Emphasis>)

Cooperation in animal social groups may be limited by the threat of free riding, the potential for individuals to reap the benefits of other individuals actions without paying their share of the costs. Here we investigate the factors that influence individual contributions to group-level benefits by studying individual participation in territorial defense among female ringtailed lemurs (Lemur catta). To control for potentially confounding factors, particularly group size, we studied two semi-free-ranging groups at the Duke University Primate Center. First, we used a combination of experimental and observational methods to investigate the costs and benefits of territorial defense for individual lemurs. We found three indications of costs: physical contact occurred during inter-group encounters, participation in territorial defense was negatively correlated with ambient temperature, and rates of self-directed behaviors increased during encounters. Benefits were more difficult to quantify, but observational and experimental tests suggested that individuals shared the gains of territorial defense by foraging in defended territories. Thus, during experiments in which one of the groups was prevented from defending its territory, the free-ranging group made more frequent incursions into the other groups territory. Second, we examined variation in participation in territorial defense. Individuals varied significantly in their rates of aggression and genital marking during inter-group encounters. The extensive variation documented among individuals was partially accounted for by dominance rank, kinship and patterns of parental care. However, we found no evidence to suggest that participation was enforced through punishment (policing) or exchange of benefits involving grooming. In conclusion, this study provides further insights into cooperative behavior in mammalian social groups by revealing how the costs and benefits of territoriality influence patterns of individual participation in the context of shared (collective) goods.Communicated by P. Kappeler     

Habitat and age of the giant squid (<Emphasis Type="Italic">Architeuthis sanctipauli</Emphasis>) inferred from isotopic analyses

The age and habitat of the giant squid, Architeuthis sanctipauli Velain, 1877, were determined based on isotopic analyses of the statoliths of three female specimens captured off Tasmania, Australia, between January and March 1996. Assuming that the aragonite of the statoliths formed in equilibrium with seawater, ¹⁸O analyses indicated that the squid lived at temperatures of 10.5–12.9°C, corresponding to average depths of 125–250 m and maximum depths of 500 m. The capture records indicated that these squid may have occasionally ranged still deeper, to as much as 1000 m. All the statoliths were labeled with bomb ¹⁴C (¹⁴C=+22.9 to +44.6), consistent with the depths inferred from ¹⁸O. A thin section through one of the statoliths revealed 351 growth increments grouped into check-ring structures every 10–16 increments. A model for statolith growth and the pattern of temporal change in ¹⁴C in the water column was used to estimate the ages of the three specimens. These estimates were very sensitive to the choice of depth range over which ¹⁴C values were integrated. Assuming that the capture depths represented the maximum habitat depths of these individuals, the calculations suggested an age of 14 years or less. More refined age estimates require a better understanding of the variation of ¹⁴C and temperature with depth in the areas in which the squids live.Communicated by J.P. Grassle, New Brunswick     

Dufour’s gland pheromone as a reliable fertility signal among honeybee (Apis mellifera) workers

When a colony becomes queenless and without the possibility of requeening, honeybee workers initiate reproduction and lay male eggs about a week later. Assays in which two bees were confined in a small arena revealed that they establish a reproductive dominance hierarchy, i.e., one worker demonstrates greater ovarian development than her paired bee. Reproductive dominance is independent of relatedness, and can be established between full sisters, cousins, or random nestmates. A social environment, however, is obligatory, as singly housed bees fail to develop ovaries on the same time scale. Allowing varying degrees of social interactions between the paired bees revealed that olfaction of volatile bee compounds, as well as tactile communication, seem to provide the necessary social environment. Ovarian development was accompanied by the production of queen-like Dufours gland secretion in these workers. Especially notable was the increase in the queen-like esters. This increase was tightly linked to ovarian development and not necessarily to the dominance status of the bees in the pair. Thus, the occurrence of queen-like esters can serve as a reliable fertility signal. Advertising ovarian status may recruit helper workers with less developed ovaries (and which are less likely to successfully reproduce before colony breakdown) to assist their nestmates and thereby gain inclusive fitness. Revealing the role of Dufours gland secretion as a fertility signal adds another dimension to our understanding of how queen pheromones operate. The mandibular-gland secretion is a good predictor of dominance hierarchy, being correlated with false-queen characteristics but not fertility, whereas Dufours gland secretion is a good predictor of fertility but not dominance hierarchy.Communicated by R.F.A. Moritz     

Precision and accuracy of stable isotope signals in otoliths of pen-reared cod (<Emphasis Type="Italic">Gadus morhua</Emphasis>) when sampled with a high-resolution micromill

A high-resolution sampling technique for stable-isotope measurements in otoliths is now available, but little is known about the precision and accuracy of the results when using this technique. Otolith samples of pen-reared cod (Gadus morhua L.) with well-defined temperature history were obtained using this high-resolution sampling technique. Samples from different-aged cod otoliths were collected in two yearly increments corresponding to the calendar years 1994 and 1995, and the stable-oxygen (¹⁸O) and -carbon (¹³C) isotope compositions of the samples were measured. Otolith ¹⁸O values had a clear seasonal pattern in response to the seasonal water temperature. Otolith material deposited in the same calendar years of different-aged fish showed similar ¹⁸O values. High precision of the temperature estimate using otolith ¹⁸O values was obtained when comparing results from different otoliths. The accuracy of the temperature estimates, relative to measured mean monthly water temperature, was also high, but reduced sampling resolution in the otoliths significantly reduced the accuracy due to attenuation of the ¹⁸O signals. We found that the otolith ¹³C values had a cyclic pattern, roughly in phase with the ¹⁸O values. This was most likely caused by temperature-increased metabolism. Indications of age-specific otolith ¹³C values are also presented.Communicated by M. Kühl, Helsingør     

Leaf volatiles and polyphenols in pear trees infested byPsylla pyricola. Evidence of simultaneously induced responses

Summary Feeding by the homopteranPsylla pyricola on leaves of pear trees induces the production of volatile compounds, such as (E,E)--farnesene and methyl-salicylate, as well as the production of polyphenols. The inference on induction is based on GC-MS and HPLC chromatograms from the same samples ofPsylla infested leaves, leaves from the same pear tree beforePsylla infestation and uninfested leaves from other pear trees.Psylla infestation greatly enhanced the production of volatiles ((E,E)--farnesene, methyl-salicylate and others) and triggered the production of new polyphenols, characterized by much longer retention times.However, the responses to infestation depend critically on leaf age (defined by leaf distance to apex). With respect to the leaf volatiles it appears that infested, old leaves produce fewer compounds and lower amounts of the volatiles than infested, young leaves. Moreover, there seem to be differences in pattern. Relative to (E,E)--farnesene, methyl-salicylate was found in much lower amounts in heavily infested, old leaves. With respect to polyphenols it was found that infested old leaves collected in August have polyphenols with the same retention times, but more or less equal amounts as uninfested young leaves collected in May. This shows thatPsylla infestation causes the induced response mostly in young leaves.The induced leaf volatiles may act as synomones to heteropteran bugs. As shown elsewhere,Anthocoris nemoralis responds significantly to (E,E)--farnesene and methyl-salicylate when offered in pure form against clean air in a Y-tube olfactometer. The effect of polyphenols on the performance ofP. pyricola is not yet known. Hence, a role in direct defence is still to be investigated.     

Diet and trophic position of Atlantic bluefin tuna (<Emphasis Type="Italic">Thunnus thynnus</Emphasis>) inferred from stable carbon and nitrogen isotope analysis

Stable ¹³C and ¹⁵N isotope analyses of scale, bone, and muscle tissues were used to investigate diet and trophic position of North Atlantic bluefin tuna (Thunnus thynnus Linnaeus) during residency in the northwest Atlantic Ocean off the northeast coast of the United States. Adult bluefin tuna scales collected from fish between June and October 2001 were significantly enriched in ¹³C compared to both muscle and bone across all months, while muscle was significantly enriched in ¹⁵N compared to either bone or scale throughout the same period. In muscle tissue, there was evidence of a shift over the summer from prey with ¹³C values (–17 to –18) that were characteristic of silver hake (Merluccius bilinearis) to species with ¹³C values of –20 to –21 that were similar to Atlantic herring (Clupea harengus) and sandlance (Ammodytes americanus). Depletion of ¹⁵N values in adult scales and bone compared to muscle tissue may be explained by bone and scale samples representing juvenile or life-long feeding habits, isotopic routing, or isotopic differences in amino acid composition of the three tissue types. Adult bluefin tuna were estimated to be feeding at a trophic position similar to pelagic sharks in the northwest Atlantic Ocean, while the trophic positions of yellowfin tuna (Thunnus albacares), albacore tuna (Thunnus alalunga), and juvenile bluefin tuna were indicative of a diet of up to a full trophic position below adult bluefin tuna. The close relationship between the juvenile bluefin ¹⁵N values and those of suspension feeders suggests that nektonic crustaceans or zooplankton may contribute significantly to the diet of bluefin tuna, a food source previously overlooked for this species in the northwest Atlantic Ocean.Communicated by J.P. Grassle, New Brunswick     

Rapid response to changing light environments of the calanoid copepod<Emphasis Type="Italic"> Calanus sinicus</Emphasis>

Calanus sinicus is a large calanoid copepod and a dominant species in the coastal waters of Japan. During a research cruise in Sagami Bay on 18 June 1996, we found C. sinicus performing an unusual diel vertical migration (DVM), a behavior that has not been reported in previous studies on this species. This study examined the DVM of C. sinicus under different light environments and revealed the copepods characteristic response to light. Field and laboratory results show that the DVM of C. sinicus is flexible and also confirmed its sensitivity and its rapid response to changing light environments. It is suggested that C. sinicus reacts to changes in absolute light intensity. This feature may be common in oceanic copepod species. The copepods quick reaction to light variation provides decreased predation risks and increased feeding opportunities, which make them a dominant survivor in coastal water habitats.Communicated by T. Ikeda, Hakodate     

The effect of diesel fuel on common vetch (Vicia sativa L.) plants

When petroleum hydrocarbons contaminate soil, the carbon:nitrogen (C:N) ratio of the soil is altered. The added carbon stimulates microbial numbers but causes an imbalance in the C:N ratio which may result in immobilization of soil nitrogen by the microbial biomass, leaving none available for plant growth. As members of Leguminosae fix atmospheric nitrogen to produce their own nitrogen for growth, they may prove more successful at growing on petroleum hydrocarbon contaminated sites. During a wider study on phytoremediation of diesel fuel contaminated soil, particular attention was given to the performance of legumes versus other plant species. During harvesting of pot experiments containing leguminous plants, a recurring difference in the number and formation of root nodules present on control and contaminated Common vetch (Vicia sativa L.) plants was observed. The total number of nodules per plant was significantly reduced in contaminated plants compared to control plants but nodules on contaminated plants were more developed than corresponding nodules on control plants. Plant performance of Common vetch and Westerwold's ryegrass (Lolium multiflorum L.) was compared to illustrate any difference between the ability of legumes and grasses to grow on diesel fuel contaminated soil. Common vetch was less affected by diesel fuel and performed better in low levels of diesel fuel contaminated soil than Westerwolds ryegrass. The total amount of diesel fuel remaining after 4 months in Common vetch planted soil was slightly less than in Westerwolds ryegrass planted soil.     

Are the Curative Properties of ‘Black Mud’ from the Dead Sea Due to the Presence of Bitumen (Asphalt) or Other Types of Organic Matter?

Deep black clayey sediments of the Dead Sea, previously covered with water at times of higher sea level, are being mined for therapeutic purposes and for the preparation of cosmetic products under the name Black Mud. It was claimed that the beneficial properties are due to the presence of bitumen (asphalt) as attested by the colour. Less commonly, the curative properties have been ascribed to humic acids or to organic matter derived from algae in the lake. Asphalt from the Dead Sea has been of major economic importance to the region and was used as a medication for 2000 years until the mid-20th century. Geochemical analysis of the Black Mud showed that it contains relatively little organic matter (ca. 0.6–0.8% organic carbon) and that the organic matter is mostly derived from the land masses surrounding the lake. Asphalt was totally absent and the concentration of humic acids very low. The black colour of the mud is rather due to the poorly crystallised iron sulphides. Based on this, the therapeutic properties of the mud are ascribed to its content of reduced sulphur species, its physical and chemical properties and possibly its brine content, but not to the presence of organic matter.