Metabolic rates of juvenile scalloped hammerhead sharks (Sphyrna lewini)

Oxygen consumption of juvenile scalloped hammerhead sharks, Sphyrna lewini, was measured in a Brett-type flume (volume=635 l) to quantify metabolic rates over a range of aerobic swimming speeds and water temperatures. Oxygen consumption (log transformed) increased at a linear rate with increases in tailbeat frequency and swimming speed. Estimates of standard metabolic rate ranged between 161 mg O₂ kg^-1 h^-1 at 21°C and 203 mg O₂ kg^-1 h^-1 at 29°C (mean-SD: 189ᆣ mg O₂ kg^-1 h^-1 at 26°C). Total metabolic rates ranged from 275 mg O₂ kg^-1 h^-1 at swimming speeds of 0.5 body lengths per second (L s^-1) to a maximum aerobic metabolic rate of 501 mg O₂ kg^-1 h^-1 at 1.4 L s^-1. Net cost of transport was highest at slower swimming speeds (0.5-0.6 L s^-1) and was lowest between 0.75 and 0.9 L s^-1. Therefore, these sharks are most energy efficient at swimming speeds between 0.75 and 0.9 L s^-1. These data indicate that tailbeat frequency and swimming speed can be used as predictors of metabolic rate of free-swimming juvenile hammerhead sharks.     

Genetic evidence for cryptic speciation in the marine bryozoan Alcyonidium gelatinosum

The common subtidal marine bryozoan Alcyonidium gelatinosum (L.) is one of the most confusing species of a taxonomically very difficult genus. It shows massive variation for almost all morphological characters. A number of sympatric colonies collected off the coast of Lundy (Bristol Channel) could be clearly divided on morphological grounds into two quite distinct types. Using starch gel electrophoresis the products of a range of genetic enzyme loci were compared between the two morphotypes and also between these and other Bristol Channel populations of A. gelatinosum. Minimal differences were observed between one Lundy morphotype and the other allopatric populations, which it was consequently concluded were conspecific. The second morphotype showed considerable genetic differentiation [genetic identity (Nei, 1972) =0.417] from the other Lundy population, with no common alleles at some loci. The probability of obtaining the observed results by chance in a single population is significantly low (P<10^–14). It is therefore concluded that the two morphotypes of A. gelatinosum from Lundy are not conspecific.     

Inter-island movements of scalloped hammerhead sharks (Sphyrna lewini) and seasonal connectivity in a marine protected area of the eastern tropical Pacific

Marine top predators are common at offshore bathymetric features such as islands, atolls, and seamounts, where most pelagic reef fish reside, while certain sharks perform inter-island movements between these formations. Scalloped hammerhead sharks are known to school in great numbers at small islands and seamounts in the eastern tropical Pacific (ETP) and are very susceptible to fisheries while moving into the open sea. It is, therefore, essential to understand hammerhead inter-island movements and environmental effects to provide baseline information for their conservation and management within and beyond an insular marine protected area. Movements of scalloped hammerheads were analyzed in the Galapagos Marine Reserve (GMR) and ETP, and environmental factors were linked to their movements. Hammerheads were tagged (N = 134) with V16 coded pingers (July 2006 to July 2010) in the northern Galapagos and detected at listening stations around four islands in the GMR and two isolated islands in the ETP, 700 and 1,200 km away. Hammerheads formed daytime schools at specific locations, but dispersed at night. Overall, more daytime than nighttime detections were recorded at all receivers in the northern Galapagos Islands, and more detections in the up-current sides of these islands. Hammerheads remained more days at the northern islands during part of the warm season (December–February) compared to the cool; however, fewer individuals were present in March–June. Movement modes were diel island excursions (24-h cycles) in the northern Galapagos and inter-island in the GMR and ETP at different scales: (1) short back-and-forth (<50 km, SBF), <5 days cycles, (2) medium distance (50–300 km, MDT), 5–20 days, and (3) long distance (>300 km, LDT), 15–52 days. The high degree of inter-island connectivity of hammerheads within the northern GMR is striking compared to the almost nil movement to the central GMR. A seasonal migratory pattern to locations offshore is indicated by (1) fewer hammerheads observed in the northern GMR during part of the warm season (March–June) and (2) evidence of LDT movements from the northern GMR to other islands in the ETP. LDT movements of mature female hammerheads are possibly associated with pupping areas. Our results indicate that currents, season, and individual behavior mainly drive inter-island movements of hammerheads at small (SBF) and medium (MDT) scales. These findings have important implications for the management of a highly mobile and endangered top predator within a marine protected area and beyond.     

Genetic evidence for cryptic speciation in allopatric populations of two cosmopolitan species of the calcareous sponge genusClathrina

Many sponge species are considered to be cosmopolitan. However, the systematics of marine sponges are very difficult because of the paucity of taxonomically useful characters, and hence the apparently cosmopolitan nature of many species may be simply a consequence of this. In this paper, geographically distant populations of two pairs of cosmopolitan calcareous sponges of the genusClathrina were compared genetically.C. clathrus andC. cerebrum were collected by SCUBA diving between January and March 1989 from two localities: the Mediterranean Sea at La Vesse, near Marseille, Frances, at 9 to 12 m depth, and from the South West Atlantic at Arraial do Cabo, about 200 km east of Rio de Janeiro, at 2 to 10 m depth. Very high levels of gene divergence were found between the allegedly conspecific populations. The levels of genetic identity,I, observed are so low (I=0.128 and 0.287) that the populations clearly cannot be considered conspecific. New species names ofC. aurea sp. nov. andC. brasiliensis sp. nov. are therefore assigned to the southwest Atlantic counterparts ofC. clathrus andC. cerebrum, respectively. It is concluded that, at least for the species studied, and probably for many other species in taxonomically difficult groups, the actual distributions of single species may be far more geographically restricted than is generally assumed.     

Highly directional swimming by scalloped hammerhead sharks,Sphyrna lewini,and subsurface irradiance,temperature, bathymetry,and geomagnetic field

The homing behavior of scalloped hammerhead sharks (Sphyrna lewini) to and fro between Espiritu Santo Seamount and Las Animas Island and the surrounding pelagic environment was studied to reveal their mechanisms of navigation in the oceanic environment. Four sharks were tagged with ultrasonic transmitters and tracked at the former location and one shark at the latter site during July, August, or September between 1981 and 1988. Hammerhead swimming movements were highly oriented: the mean coefficient of concentration (r) for sets of ten consecutive swimming directions recorded during eight homing movements by three hammerhead sharks ranged from 0.885 to 0.996. Drift within a current could not explain this directionality, since highly variable directions were recorded from a transmitter floating at the sea surface after becoming detached from a shark. Forward swimming momentum was an unlikely explanation, since highly directional swimming was maintained for a period of 32 min with only a gradual change in course. To maintain directionality over this period, an environmental property should be necessary for guidance. The hammerheads swam at night, with repeated vertical excursions ranging from 100 to 450 m deep, out of view of either the sea surface or the sea floor. The sharks' vertical diving movements were compared to distributions of spectral irradiance (relative to elasmobranch scotopic and photopic visual sensitivities), temperature, and current-flow directions in the water column. No relationships were evident between these properties and the sharks' oriented swimming movements. Movements of scalloped hammerhead sharks to and from a seamount were compared to topographic features in bathymetry and geomagnetic field leading away from the seamount. Sharks swam repeatedly over fixed geographic paths, and these paths occurred less often along submarine ridges and valleys than maxima and minima in the geomagnetic field. No significant difference existed between the degree of association of points from the sharks' tracks and points from track simulations and 20° changes in the slope of the depth record. On the other hand, significantly more points from the sharks' tracks were associated with slope changes in the magnetic intensity record than points from track simulations. A magnetic intensity gradient of 0.037 nanoteslas/m (nT/m) existed at 175 m depth, where a shark swam directionally, and this gradient was three times steeper than that measured at the sea surface and exceeded that recorded at a depth of 200 m. The hammerheads are hypothesized to find the seamount using geomagnetic topotaxis. The shark could be attracted to and move back and forth along ridges and valleys, features in the relief of magnetic field intensities occurring over a geographical area.     

Genetic adaptation to local habitats and speciation processes within the genus Gammarus (Amphipoda: Crustacea)

A recent event of allopatric speciation (a time span of about 4 000 years) is reported from a comparison between two guilds of amphipod gammarids from the central Baltic Sea and from the Limfjord (inner Danish waters), respectively. For at least two of the five species populations involved, a genetically fixed habitat specialization has led to the establishment of reproductive barriers. It has, however, not been possible to correlate this speciation event with electrophoretically determined isozyme changes. It is argued that the lack of correlation is due to the presence of an initial time lag before a speciation event can be detected as isozyme differences between newly formed species populations as predicted by the neutrality hypothesis for evolution at the molecular level.     

Genetic evidence of population heterogeneity and cryptic speciation in the ommastrephid squid Martialia hyadesi from the Patagonian Shelf and Antarctic Polar Frontal Zone

Horizontal starch gel electrophoresis was used to investigate levels of genetic differentiation between four samples of the nominate squid species Martialia hyadesi Rochbrune and Mabille, 1889, obtained from regions of the Patagonian Shelf and Antarctic Polar Fron-tal Zone over 1000 km apart. M. hyadesi is an ecologically important South Atlantic ommastrephid squid and it is probable that, in the future, fishing effort will be increasingly directed towards this species. Details regarding the population structure of the species are therefore required. In comparison with the other three samples of M. hyadesi, one of the samples from the Patagonian Shelf (PAT 89II) exhibited fixed allelic differences at 16 of the 39 enzyme loci which were resolved (genetic identity, I=0.51). This high level of genetic differentiation contradicts the apparent morphological similarity between samples, indicating the presence of a cryptic or sibling congeneric species. Deviations from Hardy-Weinberg equilibrium and significant differences in allele distribution were also detected within and between the other three putative M. hyadesi samples, suggesting that the species fails to maintain effective panmixia across its geographical range. The occurrence of both temporal (1986 cf. 1989) and geographic structuring within the species complex is consequently indicated, caused possibly by an overlap of reproductively isolated stocks (stock mixing) outside their respective breeding areas. Low levels of genetic variability were detected throughout the samples examined, estimates of average heterozygosity per locus within the two species detected being in the order of 0.01 and 0.002. These values are discussed in relation to levels of genetic variability reported for other squid species, and in comparison with values typically expected for marine invertebrates.     

Seasonal changes in movements and habitat preferences of the scalloped hammerhead shark (Sphyrna lewini) while refuging near an oceanic island

Movements and habitat preferences of sharks relative to a central location are widely documented for many species; however, the reasons for such behaviors are currently unknown. Do movements vary spatially or temporally or between individuals? Do sharks have seasonal habitat and environmental preferences or simply perform movements at random at any time of the year? To help understand requirements for the designation of critical habitats for an endangered top predator and to develop zoning and management plans for key habitats, we examined vertical and horizontal movements, and determined habitat and environmental preferences of scalloped hammerhead sharks (Sphyrna lewini). We tracked seven hammerheads for 19–96 h at Wolf Island (1.38ºN, 91.82ºW) between 2007 and 2009 using ultrasonic transmitters with depth and temperature sensors, and we profiled temperature through the water column. Movements of individual hammerheads fell in two classes: constrained (remaining near the island) and dispersive (moving offshore to pelagic environments). The central activity space or kernel off the southeast side of Wolf Island was small and common to most, but the area varied among individuals (mean ± SE 0.25 ± 0.2 km²), not exceeding 0.6 km² for any of the sharks, and not changing significantly between seasons. In general, hammerheads showed preference for the up-current habitat on the eastern side of Wolf Island in both the warm and cold seasons. However, the depth of sharks varied with season, apparently in response to seasonal changes in the vertical structure of temperature. Hammerheads performed frequent vertical excursions above the thermocline during offshore movements and, in general, were observed to prefer temperatures of 23–26 °C found above the thermocline. At times, though individuals moved into the thermocline and made brief dives below it. Our results provided evidence that hammerheads (1) are highly selective of location (i.e., habitat on up-current side of island) and depth (i.e., top of the thermocline) while refuging, where they may carry out essential activities such as cleaning and thermoregulation, and (2) perform exploratory vertical movements by diving the width of the mixed layer and occasionally diving below the thermocline while moving offshore, most likely for foraging.     

Phylogeographic patterns of the mysid Mesopodopsis slabberi (Crustacea, Mysida) in Western Europe: evidence for high molecular diversity and cryptic speciation

The phylogeographic patterns among populations of Mesopodopsis slabberi (Crustacea, Mysida), an ecological important mysid species of marine and estuarine habitats, were analysed by means of DNA sequencing of a fragment of the mitochondrial cytochrome c oxidase subunit I (COI) and the 16S ribosomal RNA genes. Samples of M. slabberi collected from five Atlantic and two Western Mediterranean populations were investigated. Very high levels of within-population molecular diversity were observed in all samples (mean h=0.807 and π=0.0083), with exception of the Mediterranean Ebro population which contained only one haplotype. Differentiation among populations was high, and a clear phylogeographic break was observed between the Atlantic and Mediterranean populations. Moreover, a strong differentiation was detected between both populations in the Western Mediterranean Sea (Alicante and Ebro delta), while two divergent lineages occurred in sympatry within the Atlantic Mondego estuary. The high congruence between both the COI and 16S rRNA sequence data, the reciprocal monophyly of the different mitochondrial clades and the levels of nucleotide divergence between them suggest the presence of a complex of cryptic species within M. slabberi. Estimations of divergence time between the different mitochondrial lineages indicate that a split occurred during the late Miocene/early Pliocene. Such a divergence could be concordant with vicariant events during sea-level drops within the Mediterranean region at that time. However, within the Mediterranean Sea, the potential of divergence through ecological diversification cannot be ruled out.     

The utility of the 16S gene in investigating cryptic speciation within the Brachionus plicatilis species complex

Recent reports indicate an extensive amount of molecular evolution separating cryptic taxa as well as significant population structure at a microgeographical scale. Appropriate molecular markers are particularly suitable for distinguishing cryptic biological species. In this study, we examine the phylogenetic utility of 16S rRNA in elucidating the evolutionary relationships within the recently described euryhaline Brachionus plicatilis species complex. In addition, we assess the applicability of this marker in the genetic identification and monitoring of rotifer populations. We have sequenced a 378-bp fragment of the mitochondrial 16S rRNA gene in laboratory reference strains, hatchery clones as well as collections from a wild population of the subsaline Lake Koroneia (Northern Greece). Also, restriction fragment length polymorphism (RFLP) analysis was performed with eight restriction endonucleases. Rotifer samples are distinguished into six genetically divergent lineages. Average sequence divergence between lineages is 0.1038. The evolutionary relationships and divergence time-scales revealed with the 16S sequence data are in agreement with previous analyses using different mitochondrial and nuclear markers. The 16S region appears to have several advantages over other regions of the genome regarding use of species-specific primers, ease of amplification from single specimens and undiluted informational content over both recent and more ancient separations. It has also exhibited maximum discriminatory power (100% success) between lineages during RFLP analysis. The 16S assayed region has proven especially informative and consistent in detecting, supporting and establishing the lineage status within the B. plicatilis species complex both from a phylogenetic perspective and as an identification tool.     

Diel movement patterns of the scalloped hammerhead shark (Sphyrna lewini) in relation to El Bajo Espiritu Santo: a refuging central-position social system

Summary Movement patterns of scalloped hammerhead sharks in the vicinity of El Bajo Espiritu Santo, a seamount in the Gulf of California, were determined by tracking by ultrasonic telemetry 13 sharks and marking 100 sharks. The 13 tracked sharks swam back and forth along the seamount ridge throughout the day. They did not swim in different directions to reduce swimming effort when currents changed from a parallel to a perpendicular orientation to the ridge. Sharks tracked up to 8 km away into the pelagic environment soon returned to the seamount. From such trackings and repeated observations of marked sharks over periods of several weeks, it is believed that most sharks disperse and return to the seamount in a rhythmical fashion. The separate departures of individual hammerheads in five paired trackings indicated that the sharks left the seamount either in small groups or singly. For these reasons, we argue that the social system of the scalloped hammerhead shark can be described as a refuging system.     

Genetic evidence for separation of two sharks,Carcharhinus limbatus andC. tilstoni,from Northern Australia

Allozyme electrophoresis was used to show that the two morphologically different groups ofCarcharhinus limbatus co-existing in waters to the north of Australia were distinct species. Forty-eight loci were examined in 967 specimens of the normal form and 20 specimens of the dark pelvic form collected between 1982 and 1985. Two loci exhibited nearly fixed allelic differences between the two forms, indicating that they must be considered as two distinct species. Further genetic analysis ofC. limbatus specimens from the West Indies (the typelocality) and South Africa showed that the rare dark pelvic form is the trueC. limbatus, while the normal form in Australian waters isC. tilstoni.     

Genetic evidence for the occurrence of a cryptic species with the littoral nemerteans Lineus ruber and L. viridis (Nemertea: Anopla)

Lineus ruber and L. viridis are common intertidal heteronemerteans found on the coasts of northern Europe and the east coast of North America. The two species resemble each other morphologically and were synonymised for many years prior to being separated on the basis of larval development. In this study, specimens of L. ruber and L. viridis were collected from sites along the west and southwest coast of Britain, northern France and the east coast of North America. The external morphology and allele frequencies of isozymes of the two species were compared from all sites. The external morphology of L. ruber and L. viridis was similar but they could generally be separated by colour. Allele frequencies for up to 13 enzyme loci between sympatric populations of L. ruber and L. viridis indicated that these two species are genetically very different (Nei's genetic identity=0.090 to 0.083). Allele-frequency data also indicated the presence of a third genetic type occurring sympatrically with populations of both L. ruber and L. viridis on the coasts of Britain and France but not on the coast of North America. Fixed differences in allele frequencies between populations of the third genetic type and sympatric populations of L. ruber and L. viridis were observed across multiple loci. Genetic identity between the third genetic type and sympatric populations of L. ruber and L. viridis were extremely low (Nei's genetic identity =<0.078). Such large genetic differences between populations indicate a barrier to gene flow and reproductive isolation. The aberrant type, which exhibits sufficient morphological variation to prevent individuals being distinguished from those of both L. ruber and L. viridis, therefore represents a separate species. The low genetic identities found in intrageneric comparisons of species found in this study have been found in other studies on nemerteans. They may indicate systematic problems within these groups or other phenomena such as morphological stasis.     

Incipient speciation within a subgenus of rockfish (<Emphasis Type=Italic>Sebastosomus</Emphasis>) provides evidence of recent radiations within an ancient species flock

The high frequency of speciation events associated with species flocks (i.e., radiations of closely related species) provides invaluable insight into the speciation process. Investigations of the speciation process in the marine environment are rare, and therefore, the genetic analysis of the rockfish genus Sebastes, considered an ancient marine species flock, provides an opportunity to investigate this process in the sea. Using both mitochondrial and nuclear markers, we analyzed five closely related species within the rockfish subgenus Sebastosomus. Our goal was to understand the evolutionary history and genetic relationships among species within this group and to provide evidence of recent speciation events within the subgenus. In the genetic analysis of the subgenus, we found different stages of the speciation process, with greater genetic divergences among three of the five species, evidence of recent divergence between two of the five species, Sebastes entomelas and S. mystinus, and significant genetic divergence between two lineages within S. mystinus revealing a signature of incipient speciation. We also found frequency differences of the two S. mystinus lineages among sample locations and found no evidence of introgression between the lineages at the location where both coexist. Although Sebastes is an example of an ancient species flock, this study provides evidence of ongoing speciation within the genus and reveals stages of this process from incipient to distinct species.     

Genetic evidence for two sibling species within Polydora cf. ciliata (Polychaeta: Spionidae) from the Sea of Japan

Two morphologically indistinguishable but genetically very distinct polychaete species of the family Spionidae are found in the Peter the Great Bay of the Sea of Japan. Both species bore into molluscan shells and occur sympatrically. They are close in morphology to Polydora ciliata (Johnston, 1838) and some other members of the redefined P. ciliata species complex. The systematics of the two species is still under consideration by one of us (V.I.R.). A total of nine enzyme systems comprising 12 isozyme loci were examined in both the species by means of starch gel electrophoresis. Nei's genetic distance between them proved surprisingly high (D= 1.942) being the highest among estimates so far obtained for pairs of polydorid species. The levels of intraspecific genetic variation in both species (Polydora sp. 1, H _e= 0.287 ± 0.064; Polydora sp. 2, H _e= 0.111 ± 0.045) are comparable to those of other electrophoretically studied polydorids. The problem of taxonomic integrity of cosmopolitan marine invertebrate species is briefly discussed as well as the usefulness of isozymes as species-specific characters suitable for unequivocal discrimination between sibling polydorid species. A list of enzymes characterized by interspecifically variable numbers of expressed isozymes within polydorids is given and recommended for such a discrimination. Received: 22 July 1997 / Accepted: 19 February 1998     

Genetic evidence for multiple parentage in eastern kingbirds (Tyrannus tyrannus)

Summary Electrophoretic analysis was performed on 28 families of eastern kingbirds (Tyrannus tyrannus) from northern Michigan to estimate the occurrence of multiple parentage. Out of 19 families used in the final analysis, at least one putative parent was excluded in 9 families, or 18 out of 60 offspring (30% of offspring). Distribution of exclusion types conforms most closely to a model of quasi-parasitism, rather than extra-pair fertilizations, with secondary females laying their eggs in the primary female's nest, but random brood parasitism cannot be ruled out as an additional or alternative source of stray genes. Based on the model of random parasitism, an estimated 39% of all offspring in this population may be unrelated to one or both of the putative parents, or 53% based on a model of quasi-parasitism. Heretofore, eastern kingbirds have been considered to be exclusively monogamous; no behavioral evidence for alternative reproductive strategies has ever been reported for this species.     

Genetic evidence for inter-oceanic subdivision of bigeye tuna (Thunnus obesus) populations

Bigeye tuna (Thunnus obesus Lowe, 1839) are a commercially important species of tuna found in the Atlantic, Indian, and Pacific oceans. To initiate an analysis of bigeye tuna population-structure, three PCR–RFLP assays were developed based on the published mtDNA control-region sequences of four bigeye tuna. Population analyses using these three restriction assays on a total of 248 individuals resulted in an array of 13 composite haplotypes. A total of 347 nucleotides of mtDNA control-region sequence was characterized for 11 of the 13 composite haplotypes. Phylogenetic analyses demonstrated that the DNA sequences belong to two monophyletic clades. However, only one of the three restriction assays was able to discriminate between the two clades. The other two assays were confounded by excessive homoplasy. Both parallel (independent occurrences of the same nucleotide change) and convergent (different nucleotide changes within the same restriction site) changes of restriction sites were observed. These results emphasize the importance of DNA sequence-analysis for the interpretation of restriction-site polymorphism data. Analyses of the frequency distribution indicated that samples of bigeye tuna from the Atlantic Ocean were genetically distinct from those found in the Indian and Pacific oceans. Thus, these results reject the null hypothesis of a single global population of bigeye tuna. Received: 16 June 1997 / Accepted: 25 July 1998     

Movements of tiger sharks (Galeocerdo cuvier) in coastal Hawaiian waters

The giant clam Tridacna crocea harbors in the mantle tissue symbiotic microalgae commonly called zooxanthellae. Isolated zooxanthellae release glycerol into the medium in the presence of mantle tissue homogenate (MH), but it is not clear whether the cells do so in situ. In order to determine the photosynthetic products released by zooxanthellae in the mantle of the giant clam we traced photosynthetic fixation products from ¹³C- and ¹⁴C-bicarbonate both in the clam and in isolated zooxanthellae (IZ) in the presence or absence of MH. After 15 min incubation in the absence of MH the IZ released less than 0.6% of the fixed labeled carbon, mainly as glucose. The major intracellular photosynthates were neutral lipids, which constituted 20 to 40% of the total extractable ¹⁴C. In the presence of MH, the IZ released up to 5.6% of the total fixed ¹⁴C, mostly as glycerol, and the major intracellular photosynthate was glucose. In an intact clam incubated in sea water containing ¹⁴C-bicarbonate, 46 to 80% of the fixed ¹⁴C was translocated from the zooxanthellae to the host tissues. Most of the ¹⁴C in the hemolymph, in the isolated zooxanthellae and in intact mantle tissue (containing zooxanthellae) was recovered as glucose. No ¹⁴C-glycerol was detected in the mantle after 1 to 30 min incubation, and, even after 60 min, far less ¹⁴C-glycerol was synthesized than by IZ in the presence of MH. The possibility that in clam tissue glycerol is converted to glucose was examined by tracing the labeled carbon from ¹⁴C-glycerol injected into the adductor muscle. After 5 min incubation, no labeled glucose was found in the hemolymph, but after 60 min, some 20% was found as glucose. Thin slices containing zooxanthellae, cut from the surface of the mantle, fixed inorganic carbon supplied as NaH¹⁴CO₃ in the medium and mainly released ¹⁴C-glucose. The addition of MH to the surrounding medium did not affect the release rate or form of release product. When the slices were cut into smaller pieces, however, the ratio of glycerol to glucose in the release product increased. These results indicate that in the presence of MH the metabolism of isolated zooxan- thellae was different from that of zooxanthellae in the mantle. In the presence of MH, isolated zooxanthellae release mostly glycerol, whereas in the mantle they release glucose. Received: 18 February 1998 / Accepted: 4 December 1998     

Movements of blue sharks (Prionace glauca) in depth and course

Acoustic telemetry was used to follow 22 blue sharks,Prionace glauca (Linnaeus), over the continental shelf and slope in the region between George's Bank and Cape Hatteras between 1979 and 1986. The sharks frequently made vertical excursions between the surface and depths of several hundred meters. The oscillations, which were repeated every few hours, were largest in the daytime and were smaller in amplitude and confined to depths near the thermocline at night. This behavior was prominent in trials from August through March, but was not seen from June through July. Diving is discussed in terms of a hunting tactic and behavioral thermoregulation. Most of the sharks moved in a southeasterly direction from the release point and many maintained a constant course day and night for several days. The sharks may orient to the earth's magnetic field, or to the ocean's electric fields, allowing them to swim on a constant heading in the absence of celestial cues. These possibilities are discussed in the appendix.     

Genetic evidence for mixed parentage in nests of the emu (Dromaius novaehollandiae)

Parentage in emus (Dromaius novaehollandiae) was examined by microsatellite analysis using four independent loci. Of 106 chicks sampled in one breeding season from 18 nests, 54 (51%) were not fathered by the nesting male, 12 (11%) were not from the observed mate of the sitting male, and 9 (8%) represented intra-specific brood parasitism, having no alleles in common with either nest parent. Some males (11%) fathered all chicks in their nests, but the majority showed high levels of cuckoldry. Those males commencing incubation earliest in the season tended to have the highest levels of paternity in their own nests. These results reveal a high frequency of extra-pair fertilisations and resultant cuckoldry in a predominantly socially monogamous bird and support recent reports which have described the emu mating system as a complexity of polyandrous, promiscuous and monogamous behaviour. Parentage assignment of chicks resulting from extra-pair fertilisations revealed an evenly scattered pattern of paternity that did not show any particular male dominance in reproductive success. These results lead to a reassessment of behavioural observations of emus, the consequences of parentage distribution, and theories about mating systems and sexual selection. The frequency of extra-pair copulations and intra-specific brood parasitism suggests patterns of descent that differ greatly from those implied by social monogamy. Received: 27 November 1999 / Received in revised form: 8 January 2000 / Accepted: 8 February 2000