Large-Eddy Simulation of Coastal Upwelling Flow

Large-eddy simulations were carried out to study laboratory-scale realizations of coastal upwelling in an annular rotating tank with a sloping bottom. A two-layer stratified fluid was set into rigid body motion with the tank and then driven by the relative rotation of a solid top lid. The simulation code developed in this work was a three-dimensional incompressible Navier-Stokes solver using the message passing interface. The simulation runs were performed on a distributed memory massively parallel computer, namely, the IBM SP2. The simulation results were able to reveal the evolution of the complex upwelling structures in detail. The results were used to compare with and to complement two relevant series of coastal upwelling experiments. A Rayleigh-Taylor type of instability took place in the top inversion layer due to the unstable stratification after establishment of the upwelling front. The primary upwelling front was unstable to azimuthal perturbations and developed large amplitude baroclinic waves. The frontal wave structure consists of cyclone/anticyclone pairs. Whether cyclonic eddies containing the lower-layer fluid pinch off from the front depends on the _* value. The non-dimensional parameter _*=gh ₀/u _* f_s, which was first introduced by Narimousa and Maxworthy, combines the effects of stratification, rotation and surface stress and can be used to characterize the upwelling flow field. Our studies show that the frontal instabilities are much more intense and the upwelling front itself displays strong unsteadiness and cyclonic eddies containing the lower-layer fluid pinch off from the front when _* is significantly less than 5.8. For _*=5.8, the frontal instabilities are less intense and no pinched-off process is observed. To separate these regimes, a critical value of _* of about 5.4 is consistent with Narimousa and Maxworthy's results.     

Nuclear and mitochondrial DNA markers indicate unidirectional gene flow of Indo-Pacific to Atlantic bigeye tuna (<Emphasis Type="Italic">Thunnus obesus</Emphasis>) populations,and their admixture off southern Africa

A sharp genetic break separates Atlantic from Indo-Pacific bigeye tuna (Thunnus obesus) populations, as the frequencies of two major mitochondrial (mt) DNA types ( and ) found in this species are different across the tip of southern Africa. The level of nucleotide divergence between mtDNA types and is of the same order as that between reproductively isolated taxa. To further investigate the genetic structure of bigeye tuna over its distribution range and in the contact zone off southern Africa, bigeye tuna samples collected between 1992 and 2001 (including samples from a previous mtDNA survey) were characterized for four nuclear DNA loci and for mtDNA. Nuclear markers did not support the hypothesis that and mitochondria characterize sibling species. Significant allele-frequency differences at one intronic locus (GH2) and one microsatellite locus (µ208) were found between Atlantic and Indo-Pacific samples, although the level of nuclear genetic differentiation (Weir and Cockerhams =0.025 to 0.042) was much lower than in mtDNA ( =0.664 to 0.807). Probabilistic Bayesian assignment of individuals to a population confirmed that southern African bigeye tuna samples represent a simple mixture of individuals from Atlantic and Indian stocks that do not interbreed, with a higher contribution from Indian Ocean individuals (about 2/3 vs. 1/3).Communicated by M.S. Johnson, Crawley     

Residues of Organochlorinated Pesticides in Eggs of Water Birds from Tai Lake in China

The levels of organochlorine compounds in eggs of water birds from the colony on Tai Lake in China were studied. The eggs were collected in 2000 and belonged to the following species: 65 samples of black-crowned night heron (Nycticorax nycticorax), 36 samples of little egret (Egretta garzetta), 26 samples of cattle egret (Bubulcus ibis) from 13 clutches and 43 samples of Chinese pond heron (Ardeola bacchus) from 17 clutches. Dichlorodiphenyltrichloroethane (DDT) and its derivates (DDE and DDD), hexachlorocyclohexane (HCH) and its isomers (-HCH, -HCH, -HCH, -HCH), heptachlor, heptachlor epoxide, aldrin, dieldrin, endrin, endrin aldehyde, -endosulfan, -endosulfan, and endosulfan sulfate were determined in the laboratory by gas chromatography. The data showed that DDE had the highest levels in all the samples, followed by -HCH. The mean levels of DDE among the water bird species were in the order as follows: black-crowned night heron (5464.26ng/g, dry weight) > Chinese pond heron (2791.12ng/g, dry weight) > little egret (1979.97ng/g, dry weight) > cattle egret (660.11ng/g, dry weight). DDT and its metabolites accounted for 90% of the total organochlorines, except that it was only 73% for cattle egret. The differences of the residue among the bird species were statistically significant and could be attributed to their variations in prey and habitat. Although the DDE burdens in Tai Lake were much lower than 8 g/g (wet weight) which are thought to have significant adverse effects on black-crowned night herons, they would be expected to increase the risk of adverse effects on survival of chicks of herons and egrets, particularly black-crowned night heron, based on the critical value of 1 g/g (wet weight) DDE. The burdens of HCHs in this study were higher and the cyclodienes were lower than those found elsewhere.     

Circadian rhythmicity of locomotor behavior and oviposition in female Teleogryllus commodus

Summary Locomotor activity of the Teleogryllus commodus is under circadian control, with _LL=25.3 h and _DD=23.4 h. In LD 12:12 h, running occurs either exclusively during darkness (57%), mainly in the dark phase (35%), or with substantial activity peaks in both phases (8%). For oviposition, _LL>24 h and _DD<24 h; in LD 12:12 h, 80% of the studied deposited their eggs mainly during the light phase and 20% primarily during darkness. In the discussion, a temporal correlation between stridulation, spermatophore formation, and locomotion, and oviposition is established, which serves to make an encounter and mating between the sexes as likely as possible.Dedicated to Prof. Dr. M. Gersch on the occasion of his 70th birthday     

Ancient divergences and recent connections in two tropical Atlantic reef fishes<Emphasis Type="Italic"> Epinephelus adscensionis</Emphasis> and<Emphasis Type="Italic"> Rypticus saponaceous</Emphasis> (Percoidei: Serranidae)

Reef habitats of the tropical Atlantic are separated by river outflows and oceanic expanses that may preclude larval dispersal or other population connections in shorefishes. To examine the impact of these habitat discontinuities on the intraspecific phylogeography of reef-associated species we conducted range-wide surveys of two amphi-Atlantic reef fishes that have dispersive pelagic larval stages. Based on 593 bp of mtDNA cytochrome b from the rock hind Epinephelus adscensionis and 682 bp from the greater soapfish Rypticus saponaceous (n=109 and 86, respectively), we found evidence of relatively ancient separations as well as recent surmounting of biogeographic barriers by dispersal or colonization. Rock hind showed slight but significant population genetic differentiation across much of the tropical Atlantic Ocean (_ST=0.056), but deep divergence between the southeastern United States and seven other localities from the Bahamas to the south, central and east Atlantic (mean pairwise d=0.040, overall _ST=0.867). The geographic distribution of the two rock hind lineages is highly unusual in genetic studies of Caribbean Sea reef fishes, because those lineages are separated by less than 250 km of open water within a major biogeographic region. In contrast, highly significant population genetic structure was observed among greater soapfish from the SW Caribbean, Brazil, and mid-Atlantic ridge (_ST=0.372), with a deep evolutionary separation distinguishing putative R. saponaceous from West Africa (mean pairwise d=0.044, overall _ST=0.929). Both species show evidence for a potential connection between the Caribbean and Brazilian provinces. While widespread haplotype sharing in rock hind indicates that larvae of this species cross oceanic expanses of as much as 2000 km, such a situation is difficult to reconcile with the isolation of populations in Florida and the Bahamas separated by only 250 km. These findings indicate that populations of some species in disjunct biogeographic zones may be isolated for long periods, perhaps sufficient for allopatric speciation, but rare gene flow between zones may preclude such evolutionary divergence in other species.Communicated by P.W. Sammarco, Chauvin     

Allozyme and mitochondrial DNA variation in orange roughy,Hoplostethus atlanticus (Teleostei: Trachichthyidae): little differentiation between Australian and North Atlantic populations

Allozyme and mitochondrial DNA (mtDNA) genetic variation was compared in orange roughy (Hoplostethus atlanticus Collett) collected from waters off southern Australia and from waters about 22 000 km away in the North Atlantic west of Scotland. Samples were screened for 11 polymorphic allozyme loci and with 9 restriction enzymes. Significant heterogeneity between the two areas was detected for three allozyme loci (ADA ^*, CK ^* and GPI-1 ^*), and the overall G _ST (gene-diversity statistic) value of 1% was small but significant. Significant mtDNA haplotype heterogeneity was observed after ²- of haplotype frequencies but not after a G _ST analysis. Nucleotide sequence-diversity analysis showed very low net divergence (0.0023%) between the two samples. The Australian orange roughy had a lower allozyme heterozygosity and a lower mitochondrial DNA nucleon diversity than the North Atlantic sample. The very limited, although significant, allozyme and mitochondrial DNA heterogeneity between these areas suggests that there is some gene flow between these two populations. The species appears to be widespread, with its presence reported from the southern Pacific, southern Indian, and northern and southern Atlantic Oceans, and it is likely that gene flow between the antipodes is mediated by stepping-stone exchange between adjacent populations rather than by direct migration.     

Lead-based paint in dwellings: The potential for contamination of the home environment during renovation

The quantity and particle size characteristics of lead in dust released during three different paint removal techniques was determined under controlled conditions and in situ in a dwelling. Air-lead and deposited dust-lead levels were highest after sanding but burning-off and hot-air removal methods also produced significant contamination. The importance of dust particle-size and lead is discussed in relation to the potential hazard to home renovators via inhalation and to children via the hand-to-mouth route.This paper is also being published in Proceedings of the Conference on Lead in the Home Environment and is one of several selected from SEGH sponsored conferences for simultaneous publication in this journal.     

Genetic heterogeneity among blacktip shark,<Emphasis Type="Italic"> Carcharhinus limbatus</Emphasis>, continental nurseries along the U.S. Atlantic and Gulf of Mexico

Genetic population structure of the blacktip shark, Carcharhinus limbatus, a commercially and recreationally important species in the southeast U.S. shark fishery, was investigated using mitochondrial DNA control region sequences. Neonate blacktip sharks were sampled from three nurseries, Pine Island Sound, Terra Ceia Bay, and Yankeetown, along the Gulf of Mexico coast of Florida (Gulf) and one nursery, Bulls Bay, on the Atlantic Ocean coast of South Carolina (Atlantic). Sequencing of the complete mitochondrial control region of 169 neonates revealed 10 polymorphic sites and 13 haplotypes. Overall haplotype diversity and percent nucleotide diversity were 0.710 and 0.106%, respectively. Haplotype frequencies were compared among nurseries to determine if the high mobility and seasonal migrations of adult blacktip sharks have maintained genetic homogeneity among nurseries in the Atlantic and Gulf. Chi-square analysis and AMOVA did not detect significant structuring of haplotypes among the three Gulf nurseries, P(²)=0.294, _ST=–0.005 to –0.002. All pairwise AMOVA between Gulf nurseries and the Atlantic nursery detected significant partitioning of haplotypes between the Gulf and Atlantic (_ST=0.087–0.129, P<0.008), as did comparison between grouped Florida Gulf nurseries and the Atlantic, _CT=0.090, P<0.001. Based upon the dispersal abilities and seasonal migrations of blacktip sharks, these results support the presence of philopatry for nursery areas among female blacktip sharks. Our data also support the treatment of Atlantic and Gulf blacktip shark nursery areas as separate management units.Communicated by P.W.Sammarco, Chauvin     

Relationship of daily and circatidal activity rhythms of the fiddler crab,<Emphasis Type="Italic"> Uca princeps</Emphasis>, to the harmonic structure of semidiurnal and mixed tides

Intertidal organisms may employ circatidal rhythms to track the tidal cycle, but tidal patterns may vary within a species range and necessitate adaptation to the local tides. Circatidal rhythms were examined in populations of the eastern Pacific fiddler crab Uca princeps (Smith) from four sites with differing tidal characteristics, La Paz (24°10N; 110°21W), San Blas (21°33N; 105°18W) and Manzanillo (19°6N; 104°24W), Mexico (lower amplitude, mixed semidiurnal tides) and Mata de Limon, Costa Rica (9°55N; 84°43W) (high-amplitude, semidiurnal tides). Local tides were characterized by harmonic constants of M₂, S₂, K₁, and O₁, partial tides that largely determine their semidiurnal and diurnal features. Rhythmic structure in continuously recorded locomotor activity of individual crabs held under laboratory conditions was described by cosinor and periodogram methods of time-series analysis. Both daily and circatidal rhythms were found in crabs studied in light–dark cycles set to local conditions at the time of collection. Crabs at all four sites shared a tendency toward bimodality, with a mid-morning activity peak and varying degrees of nocturnal activity. Circatidal rhythms closely matching the period of the 12.42-h M₂ partial tide were consistently present at all sites except Manzanillo. At Mata de Limon, the circatidal rhythm clearly dominated locomotor activity, but was strongly modulated by a daily rhythm in a repeating pattern at a semilunar interval. In contrast, the amplitude of the daily rhythm was higher than that of the circatidal rhythm in crabs from the three mixed tide sites on the Mexican coast, where the tidal pattern is dominated by a diurnal inequality arising from the diurnal K₁ and O₁ partial tides. These results suggest that populations of U. princeps use both daily and circatidal timing systems to track local forms of the tide generated by their M₂, S₂, K₁, and O₁ geophysical counterparts.Communicated by J.P. Grassle, New Brunswick     

Genetic population structure in two tropical sponge-dwelling shrimps that differ in dispersal potential

The spatial context in which host races of parasitic animals originate is a central issue in the controversial theory of sympatric speciation. Sponge-dwelling shrimps in the genus Synalpheus provide a good system for evaluating the possibility of resource-associated divergence in sympatry. I used allozyme electrophoresis to assess the genetic population structure of two Caribbean Synalpheus species sampled in 1988 to 1990 at a hierarchy of spatial scales. S. brooksi Coutière is a host-generalist, using several sponge species in an area, and develops directly, with no planktonic larval stage. G-tests and estimates of F _ST revealed highly structured populations in this species, with significant differentiation among samples from individual reefs within a region, and strong divergence among regions (Panama, Belize, Florida). Moreover, samples of S. brooksi taken from the two sponges Spheciospongia vesparium (Lamarck) and Agelas clathrodes (Schmidt) in Panama, and separated by 3 km, showed significant differentiation at both of the loci that were polymorphic in these populations. Genetic distances between these host-associated populations averaged >60% greater than distances between samples from the same host species and were comparable to, or greater tha, those for some inter-regional comparisons. These genetic data corroborate a previous finding of demographic differences between the same populations. The second species, S. pectiniger Coutière, occurs only in Spheciospongia vesparium, and produces swimming larvae. Although allele frequencies in this species differed significantly among the three regions, S. pectiniger showed no differentiation within regions, and significantly lower differentiation (F _ST) among regions than its direct-developing congener. These data suggest that genetic population structure in these two commensal crustaceans is related to dispersal potential, and that restricted dispersal may allow the divergence of host-associated populations on a local scale.     

Preliminary report on the digestive enzymes present in the digestive gland ofPerna viridis

Quantitative data on digestive enzymes of bivalves are very limited and so this study was performed to obtain such information, and to find out whether some of these enzymes are well adapted to conditions in the digestive gland. The green musselPerna viridis was obtained from Clementi West Market in 1985. The results show that various carbohydrases are present in the digestive gland. The activities of-amylase, cellulase and-glucosidase are higher than that of other carbohydrases, whilst the activities of both lipase and protease, especially the former, are low. The pH optima of the following enzymes are also given:-amylase = 5.8; laminarinase = 5.5;-glucosidase = 5.8; trehalase = 4.8;-glucosidase = 5.5. The only enzyme which was not well adapted to conditions in the digestive gland (pH 6.2) was trehalase. The wide spectrum of enzymes present in this bivalve indicates that it has the ability to utilize a wide range of nutrients efficiently.     

Chitinolytic enzymes in the integument and midgut-gland of the shrimpPalaemon serratus during the moulting cycle

Chitinase andN-acetyl--D-glucosaminidase activity were quantified inPalaemon serratus (Pennant) integument and midgut gland during the moulting cycle. Studies were performed on specimens collected near Concarneau, France, in July 1989. The changes in specific activity are different in the two organs: in the midgut gland enzymatic activity is high throughout the whole moulting cycle with a weak peak at the early premoult Stage D1, whereas in the integument the activity of both enzymes is very low throughout post- and intermoult stages and rises only after premoult Stage D1. The highest specific activity is reached in D1 for chitinase and somewhat earlier (D1) forN-acetyl--D-glucosaminidase. The increase in specific chitinolytic activity coincides with an increase in ecdysteroids.     

Effects of light intensity on nitrate and nitrite uptake and excretion by Chaetoceros curvisetus

Michaelis-Menten uptake kinetics were observed at all light intensities. With constant illumination, the V_max and K₁ in nitrate uptake over the natural light intensity range of 0 to 2000 E were 0.343 g-at NO₃–N(g)^-1 at protein-N h^-1 and 26 E, respectively. Nitrate uptake was inhibited at higher light intensities. The K_s for nitrate uptake did not vary as a function of light intensity remaining relatively constant at 0.62 g-at NO₃–N 1^-1. With intermittent illumination, the V_mzx for light intensity in nitrate uptake over a light intensity range of 0 to 5000 E was 0.341 g-at NO₃–N(g)^-1-at protein-N h^-1. No inhibition of nitrate uptake was observed at higher than natural light intensities. Chaetoceros curvisetus will probably never experience light inhibition of nitrate uptake under natural conditions.     

Growth of the Atlantic surfclam,Spisula solidissima,from Georges Bank to the Delmarva Peninsula,USA

Age/shell length data for offshore surfclam, Spisula solidissima (Dillwyn, 1817), populations were used to estimate the parameters of the von Bertalanffy growth model by time period and region, from Georges Bank in the north to the Delmarva Peninsula in the south. Randomization tests were used to compare curves. We tested the a priori hypothesis that growth curves would change over time in the south (i.e., New Jersey and Delmarva) but remain constant in the north (i.e., Long Island and South New England). This hypothesis was proposed because surfclam population structure in the south had been altered by the hypoxic event of 1976, and possibly by intense, long-term commercial harvesting. Northern regions, unaffected by these factors, served as natural controls. Based on a comparison of data collected in 1980 with pooled data from 1989 and 1992, the hypothesis was supported. Both the growth coefficient (k) and maximum shell length (L ) declined between two time periods in the two southern regions, while during the same time interval, no change occurred in the two northern regions. Differences in growth between regions were often statistically significant. For example, compared with the southern regions, the growth coefficient on Georges Bank was larger, and those clams attained a smaller maximum length. In a comparison of adjacent regions from Delmarva to S. New England, k increased from south to north. This could imply faster growth in cooler water, as well as no relationship between growth and primary productivity. Alternatively, size-selective mortality, imposed by the commercial fishery, was discussed as a mechanism that might account for this unexpected pattern.     

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 structure of plankton communities of the Indian Ocean

Species diversity and trophic structure of plankton communities have been considered as most important biological features. The Fisher () and Simpson () indices and the information index H=- pi log₂ pi (Margalef) were used for evaluation of species diversity. The indices have been calculated within the group of filter feeders by numbers ( and H) and biomass ( and H) of the species; the best results were obtained in the latter case. The index H=- pi log₂ pi is proposed for the quantitative evaluation of trophic structure, where a portion of an individual trophic group is taken as pi. The plankton material was collected by the R.V. Vityaz in the northern Indian Ocean in January through February, 1960. Analysis of characteristics of zooplankton distribution, species and trophic diversity in connection with characteristics of the water regime resulted in the definition of 3 types of the plankton communities: (a) those in intensive divergence zones; (b) those in poor divergence zones; (c) those in regions with stable water stratification or with feebly marked convergence, distiguished by different values of biomass, species diversity, and trophic group ratios.     

Translocation of fixed carbon from symbiotic bacteria to host tissues in the gutless bivalve Solemya reidi

Naturally-occurring lipophilic inducers of larval settlement and metamorphosis wer isolated and identified for Phragmatopoma californica, a gregarious tube worm from southern California. Organic solvent extraction of the sand/organic cement matrix of tubes diminished the inducing capacity of the tube matrix. The inducing capacity was restricted to a single, highly active, HPLC-purified fraction of the organic solvent extract. Chemical analysis of this fraction revealed a mixture of free fatty acids (FFAs), dominated by eicosapentaenoic acid (20:5, 20%), palmitic acid (16:0, 14%) and palmitoleic acid (16:1, 12%). In assays of the nine FFAs that each contributed 3% or more to the active fraction, only 16:1, 18:2, 20:4 and 20:5 induced larval settlement and metamorphosis, while the others were ineffective. The larval response was contact-dependent, highly specific, and concentration-dependent, with a significant response to 16:1 and 20:4 at as low as 10 g FFA spread onto 1 g of sand (surface area 36 cm²). Active FFAs were extracted at approximately 14 g g^-1 sand from the tube matrix, although the levels encountered by larvae in nature are believed to be higher.     

The evolution of cheating and selfish behavior

Summary We examine the necessary conditions for the spread of genes that determine selfish and cheating behaviors and the rate of spread of these genes through structured populations, in order to address the question of the invadability of altruistic systems by anti-social mutations. We find that, although cheaters always have a higher relative fitness than altruists within groups, population structures which permit the evolution of altruism also preclude invasion by anti-social mutations. These results are related to a discussion by Hamilton (1971) concerning the limits to the evolution of altruistic and selfish behaviors.     

High turnover of inorganic carbon in kelp habitats as a cause of δ13C variability in marine food webs

In a study to assess qualitatively the importance of organic matter derived from kelp production in the Aleutian Islands of subarctic Alaka, replicated samples of autotrophic sources and primary and secondary consumer organisms were sampled for ¹³C among sources, sites, (treatment) islands, and years. Unanticipated variation in the ¹³C of kelps occurred among overtly similar sites at different islands. Variation in the ¹³C of the surface canopy-forming kelp Alaria fistulosa was particularly extreme, ranging from-15.5 to-28.0 compared to the understory kelps, Laminaria spp. A. fistulosa ¹³C varied by as much as 6 to 7 among similar sites at a given island within years, and by as much as 3 to 4 between years at the same sampling site. In serveral cases, ¹³C variation was weakly tracked by some consumer organisms, suggesting that even detritus pathways through the food web can be localized and tightly coupled. Dynamic cycles in the concentration and ¹³C of dissolved inorganic carbon (DIC) and aqueous CO₂ concentration ([CO₂]_aq) were measured at three sites on one island. The ¹³C or organic carbon fixed by A. fistulosa, calculated from diurnal DIC concentration and ¹³C measurements, varied by 15 and varied inversely with [CO₂]_aq concentrations. Local DIC variability, probably resulting from high productivity and decreased turbulence in dense kelp habitats, provides a possible mechanism of variation in kelp ¹³C. The short-term variability in the ¹³C of organic carbon fixed by kelps indicates that sampling methodology and design must assess this potential variation in marine macrophyte ¹³C before making assumptions about the transfer of ¹³C-invariate organic matter to higher trophic levels. On the positive side, a predictable relationship between [CO₂]_aq concentration and kelp ¹³C offers a potentially robust means to assess productivity effects on CO₂ limination in kelps and other complex aquatic macrophyte habitats.     

The erect ‘penis’ is a flag of submission in a female-dominated society: greetings in Serengeti spotted hyenas

In East Africa, spotted hyenas live in large clans in a highly structured society dominated by females. A clan is a fission-fusion society where members are often solitary or in small groups. Spotted hyenas have a ritualized greeting during which two individuals stand parallel and face in opposite directions. Both individuals usually lift their hind leg and sniff or lick the anogenital region of the other. The unique aspect of greetings between individuals is the prominent role of the erect penis in animals of both sex. Female spotted hyenas have fused outer labiae and a pseudo-penis formed by the clitoris which closely resembles the male penis and can be erected. During greetings subordinates signalled submission with gestures which were not necessarily reciprocated by the dominant participant. Asymmetries were most pronounced in greetings between adult females where the probability of asymmetries increased with the divergence in rank between partners. Greetings between adult females and males were uncommon and restricted to males above median rank, principally the alpha male. Models of primate affiliative behavior assume that benefits derived from social relationships with different individuals are not equal and that individuals are selected to maximize the benefits they receive from social relationships with others. The observed distribution of greetings between partners of different rank matched the predictions of these models. An examination of non-adaptive hypotheses on the evolution of the pseudo-penis demonstrated that the conventional scenario linking (initial) virilization of female genitalia with selection for female dominance does not explain either the initial virilization, nor the evolution of the pseudo-penis to its current form and use. We sketch a new scenario that links (1) initial virilization to the occurrence of neonatal siblicide amongst members of a twin litter, and (2) costs of maintenance, pseudo-penile control over copulation and male submission. Our analysis confirms previous adaptive hypotheses on the function of the pseudo-penis in greetings and suggests new hypotheses to account for hitherto unexplained features. Correspondence to: M.L. East