Effects of dissolved ammonium addition and host feeding with Artemia salina on photoacclimation of the hermatypic coral Stylophora pistillata

 Effects of nutrient treatments on photoacclimation of the hermatypic coral Stylophora pistillata (Esper) were studied. Studies on photoacclimation of colonies from different light regimes in the field were evaluated and used to design laboratory experiments. Coral colonies were collected in the Gulf of Eilat (Israel) from January to March 1993. Exterior branches of colonies from different depths (1 to 40 m) displayed different trends in production characteristics at reduced and very low levels of illumination. From 24 ± 3% to 12 ± 2% of incident surface photosynthetic active radiation (PAR_o), zooxanthella population density and chlorophyll a+c per 10⁶ zooxanthellae increased, a trend seen in the range of light levels optimal for coral growth (90 to 30% PAR_o). The P _max of CO₂ per 10⁶ zooxanthellae decreased, while P _max of CO₂ per 10³ polyps increased, indicating an increase in zooxanthella population density at low light levels. Proliferous zooxanthella frequency (PZF, a measure of zooxanthella division) declined significantly at light levels <18 ± 3% PAR_o. At the lowest levels of illumination (<5% PAR_o), zooxanthella population density decreased, as did the PZF; chl a+c per 10⁶ zooxanthellae was unchanged. In 28-d experiments, exterior coral branches from the upper surfaces of colonies from 3 m depth (65 ± 4% PAR_o) were incubated in aquaria under bright (80 to 90% PAR_o), reduced (20 to 30% PAR_o), and extremely low (2 to 4% PAR_o) light intensities. At each light intensity, the corals were maintained in three feeding treatments: sea water (SW); ammonium enriched SW (SW + N); SW with Artemia salina nauplii (SW + A). An increase in P _max of CO₂ per 10³ polyps was found in corals acclimated to reduced light (20 to 30% PAR_o) in nutrient-enriched SW, while in SW, where the increase in zooxanthella population density was smaller, it did not occur. Nutrient enrichments (SW + N at 2 to 4% PAR_o and SW + A at 20 to 30% PAR_o) increased zooxanthella population density, but had no effect on chl a+c per 10⁶ zooxanthellae. Acclimation for 14 d to reduced (10 to 20% PAR_o) and extremely low (1 to 3% PAR_o) light intensities shifted ¹⁴C photoassimilation into glycerol and other compounds (probably glycerides), rather than sugars. Both ammonium addition and feeding with Artemia salina nauplii resulted in an increase in photosynthetic assimilation of ¹⁴C into amino acids. We conclude that acclimation to reduced light consists of two processes: an increase in photosynthetic pigments and in zooxanthella population density. Both processes require nitrogen, the increase in zooxanthella population density needing more; this adaptation is therefore limited in nitrogen-poor sea water. Received: 19 June 1998 / Accepted: 13 June 2000     

Mating behaviour of Macrophthalmus hirtipes (Brachyura: Ocypodidae)

 We examined the mating behaviour of the New Zealand ocypodid crab Macrophthalmus hirtipes in the laboratory between February and June 1998. This species has a discrete breeding season. Mating and moulting were not linked and only intermoult females with mobile gonopore opercula were attractive to males. Allometry and compatibility of gonopods and gonopores of different-sized crabs was investigated. Under laboratory conditions, the opercula of intermoult females remained mobile on average for 11.4 d, but the duration of receptivity did not appear to be under female control. The operational sex ratio in the laboratory fluctuated greatly, but was always male-dominated. During the period of opercular mobility, females mated many times with several different males. Matings in the absence of burrows were relatively short (mean duration = 23 min, max. = 122 min) and the mating behaviour of M. hirtipes lacked courtship and mate-guarding. Males used a search-intercept method to acquire mates, with very low levels of intrasexual competition. There was no evidence of mate preference in M. hirtipes, and males spent just as long mating with ovigerous females as with non-ovigerous ones. Although M. hirtipes has ventral-type spermathecae, as do several other ocypodid crabs, it is unclear whether this promotes last-male sperm precedence. The role of burrows in modifying the mating behaviour of M. hirtipes in the field remains to be established. Received: 7 January 2000 / Accepted: 5 June 2000     

Mitochondrial DNA sequence variation and phylogeography of oceanic insects (Hemiptera: Gerridae: Halobates spp.)

Relatively few insects have invaded the marine environment, and only five species of sea skaters, Halobates Eschscholtz (Hemiptera: Gerridae), have successfully colonized the surface of the open ocean. All five species occur in the Pacific Ocean, H. germanus White also occurs in the Indian Ocean, whereas H. micans Esch- scholtz is the only species found in the Atlantic Ocean. We sequenced a 780 bp long region of the mitochondrial cytochrome oxidase subunit I gene (COI) for a total of 66 specimens of the five oceanic Halobates species. Our purpose was to investigate the genetic variation within species and estimate the amount of gene flow between populations. We defined 27 haplotypes for H. micans and found that haplotype lineages from each of the major oceans occupied by this species are significantly different, having sequences containing five to seven unique base substitutions. We conclude that gene flow between populations of H. micans inhabiting the Atlantic, Pacific, and Indian Ocean is limited and hypothesize that these populations have been separated for 1 to 3 million years. Similarly, there may be limited gene flow between H. germanus populations found in the Pacific and Indian Ocean and between H. sericeus populations inhabiting the northern and southern parts of the Pacific Ocean. Finally, we discuss our findings in relation to recent hypotheses about the influence of oceanic diffusion on the distribution and population structure of oceanic Halobates spp. Received: 29 July 1999 / Accepted: 23 November 1999     

Reproductive variability over a four-year period in the sea urchin Evechinus chloroticus (Echinoidea: Echinodermata) from differing habitats in New Zealand

 The endemic New Zealand echinoid, Evechinus chloroticus (Valenciennes), was sampled approximately monthly from September 1990 to October 1994 at three sites in Tory Channel, Marlborough Sounds, New Zealand. These channel sites (outer, mid and inner) were up to 20 km from the open ocean and differed in their shore type, exposure to wave action and macrophyte abundance. E. chloroticus showed an annual reproductive cycle at each site, with gametogenesis commencing in the late austral winter and spawning in summer. Maximum gonad indices (reproductive potential) varied spatially, with the outer site generally having higher maximum gonad indices than the inner site, and temporally, with maximum gonad indices occurring earlier in 3 of the 4 seasons at the outer site than the inner site. The mid and inner sites showed much greater variation in maximum gonad indices (range 15.83 to 26.99% and 11.87 to 20.90%, respectively) than the outer site (range 19.31 to 22.95%). Reproductive output (weight of gametes released per gram echinoid) also varied, with the different sites showing significantly different outputs in the different years. A regression of maximum reproductive potential against reproductive output was significant ( p < 0.001), and had a positive slope with an r ² of 0.79. While, the initiation of gametogenesis was relatively synchronous between sites and years, and is possibly cued by increasing daylength, it progressed at different rates among populations. Spawning did show spatial and temporal variability, occurring near the time of highest sea-surface temperatures (∼15 °C). The observed variations in reproductive cycle may be related to small-scale variability in diet and environmental conditions. Furthermore, asynchronous spawning, variable spawning duration, and variable reproductive output are likely to strongly influence annual recruitment variability in E. chloroticus, with different larval subpopulations contributing unequally in different years. The ecological consequences of this, both for the ability of E. chloroticus to propagate itself in space and time and for the management of the developing E. chloroticus fishery in New Zealand, are discussed. Received: 9 December 1998 / Accepted: 13 June 2000     

Phylogeny and biogeographic history of hake (genus Merluccius), inferred from mitochondrial DNA control-region sequences

 Phylogenetic analyses of the left domain of the mitochondrial DNA control-region sequence have been used to examine the relationships among species of the genus Merluccius (Rafinesque, 1810), and to compare these with hypotheses based on morphological, meristic and allozyme characters. Analysis of aligned sequences revealed that transition bias was much lower than in mammalian mtDNA, and that nucleotide composition of control-region sequences was biased toward A and T. We have roughly calibrated a molecular clock for the genus, based on the rise of the Isthmus of Panamá, which is believed to have created a barrier to dispersal between marine species of the Atlantic and Pacific Oceans. Our mtDNA-based phylogeny was highly congruent with allozyme-based phylogenies, but poorly so with a previously described phylogeny based on morphology. Specifically, our phylogeny resolved two well-supported principal clades, one of American (west Atlantic and east Pacific) species and the other of Euro–African (east Atlantic) species. This suggests an evolutionary history during which the ancestral lineage of Merluccius was divided between two geographic regions, with subsequent dispersal and vicariant events resulting in the evolution and distribution of extant taxa. However, the relationships between some taxa within the American clade could not be resolved. We suggest that this is consistent with an hypothesis of a rapid origin and radiation of these taxa. Received: 12 December 1998 / Accepted: 15 October 1999     

Meiofauna response to a dynamic river plume front

 The benthic response to a plume front was studied in two areas of the northern Adriatic (Mediterranean Sea) differently influenced by the Po River freshwater input. Sediment samples were collected in June 1996 and February 1997 from 12 stations. The adopted sampling strategy was able to identify the front line in real time by satellite images and to locate sampling stations along an inner–outer plume gradient in order to cover the benthic area beneath the river plume, where enhanced biological production was expected, and open-sea sediments not directly influenced by freshwater inputs. Meiofaunal parameters were compared to the physical conditions and to phytodetritus inputs, organic matter accumulation and bacterial secondary production. The sediments of the Adriatic Sea were characterised by high concentrations of phytopigments (0.6 to 13.9 μg g⁻¹ for chlorophyll a and 1.2 to 17.7 μg g⁻¹ for phaeopigments) and biopolymeric organic carbon (0.15 to 3.02 mg g⁻¹). The plume system extended for a large sector of the northern Adriatic. In the northern area, a large and highly dynamic plume area was coupled with a sediment organic matter concentration significantly higher than in open-sea sediments. In the southern sector, where the plume area and the front line did not change markedly during the year, plume–benthic coupling was evident only in the sediments beneath the front, and corresponded to phaeopigment accumulation. Bacterial parameters and secondary production were high and significantly higher in the frontal area than at open-sea stations. Meiofauna density (1342 to 8541 ind. 10 cm⁻²) did not change either by season or between areas and was significantly correlated with phaeopigments and bacterial secondary production. Meiofauna displayed different responses to plume inputs in the two sampling areas. In the northern sector, meiofauna density was coupled with organic matter distribution and displayed highest values beneath the plume. In the southern sector, the densities of copepods, turbellarians and kinorhynchs displayed highest values under the front in summer, and the same applied to total meiofauna density in winter. Juvenile decapods and copepod nauplii significantly increased their densities in sediments beneath the front. Data presented in the present study suggest that plume inputs and frontal systems, enhancing phytodetritus accumulation and benthic bacterial response, might influence density, composition and distribution of meiofaunal assemblages. As river plumes are highly variable systems affecting the trophic characteristics of the sediments underneath, their dynamics should be considered when analysing mesoscale spatial changes of meiofaunal assemblages. Received: 30 November 1999 / Accepted: 24 May 2000     

Grazing rates of organic matter and living fungal biomass of decaying Spartina alterniflora by three species of salt-marsh invertebrates

 The pathway for the flow of salt-marsh grass production into marsh food-webs is still not well defined. We compared the abilities of three marsh macroinvertebrates [salt marsh periwinkles, Littoraria irrorata (Say) (=Littorina irrorata), salt-marsh coffee-bean snails, Melampus bidentatus (Say); and a talitrid amphipod, Uhlorchestia spartinophila Bounsfield and Heard] to access standing-dead leaves of smooth cordgrass (Spartina alterniflora Loisel). The invertebrates were incubated with naturally-decaying leaves, and the rates of removal of organic matter and living fungal biomass (ergosterol) were measured. The impact of invertebrate activity upon fungal growth rates was measured as rates of fungal-membrane synthesis (incorporation of radioacetate into ergosterol). The removal rates of organic leaf biomass per mg individual biomass were highest for amphipods (700 μg mg⁻¹ d⁻¹) and lowest for periwinkles (90 μg mg⁻¹ d⁻¹), but the relatively large biomass of the snails made their removal rates per individual greater than those of amphipods. Net removal of ergosterol by all three invertebrates was >50% for yellow-brown (early-decay) leaf blades. For fully-brown (advanced-decay) blades, >50% removal of ergosterol was found only for periwinkles; exposure to coffee-bean snails and amphipods resulted in a net ergosterol reduction of ≤20%. The lower net reduction of living fungal biomass by coffee-bean snails and amphipods may have been due to fungal-growth stimulation (2.3-fold stimulation in coffee-bean snails and 1.5-fold stimulation in amphipods). Grazing by periwinkles did not stimulate fungal growth, possibly because of its high intensity. Grazing by these three salt-marsh shredders may affect marsh-grass shoot-decay in different ways. Periwinkles may abbreviate the period of fungal production, and incorporate the decaying material relatively quickly into snail biomass and fecal-pellet rain to the sediments. Coffee-bean snails and amphipods may enhance and prolong fungal production, along with the formation of fecal-pellet rain. All three invertebrates fed preferentially on leaf blades rather than leaf sheaths, and feeding rates of gastropods were higher during the night than during the day. Received: 25 November 1998 / Accepted: 4 November 1999     

Cadmium accumulation in the female shore crab Carcinus maenas during the moult cycle and ovarian maturation

 The influence of moulting and ovarian maturation on cadmium accumulation in the tissues of female shore crabs Carcinus maenas exposed to 1 mg Cd l⁻¹ in the water was investigated. Cadmium accumulation in all tissues examined was markedly increased in crabs in the postmoult stages (A and B) compared to crabs in all other moult stages. During the moult cycle, average cadmium accumulation in the midgut gland ranged from 29 μg Cd g⁻¹ dw at premoult stage (D₂) to 589 μg Cd g⁻¹ dw at postmoult stage (A). Average cadmium concentrations in the haemolymph ranged from 0.56 μg Cd ml⁻¹ at intermoult stage (C₄) to 4.6 μg Cd ml⁻¹ at postmoult stage (A), while the gills accumulated from 103 μg Cd g⁻¹ dw in intermoult stage (C₃) to 352 μg Cd g⁻¹ dw in postmoult stage (A). Cadmium concentration in gills and haemolymph was also significantly higher in crabs in late premoult stage (D₃) compared to C₄-crabs, while midgut gland cadmium concentration remained elevated in C₁- and C₃- intermoult stages relative to C₄. During ovarian maturation the cadmium accumulation in midgut gland, gills, ovaries and haemolymph decreased. Average cadmium concentration in the midgut gland decreased from 63 μg g⁻¹ dw in ovarian Stage I to 19 μg g⁻¹ dw in ovarian Stage VI. The same pattern was observed for gills, haemolymph and ovaries. The present study demonstrates that cadmium accumulation in the female shore crab strongly depends on the physiological status of the animal. A possible association between physiological calcium requirements and cadmium accumulation during moulting is discussed. Received: 20 January 2000 / Accepted: 20 July 2000     

A comparative study of feeding and growth in two coexisting species of carnivorous gastropods

The effects of variable food supply on growth rate and feeding behaviour in two coexisting intertidal gastropods were examined through field and laboratory experiments involving mark and recapture procedures, growth experiments under controlled food supply and video observations of feeding. All cohorts of both Japeuthria species grew most in summer, with the highest growth rate being achieved by the youngest cohort of both sexes. In contrast, almost all cohorts (except the youngest one) showed zero to negative growth in winter. The growth rates of Japeuthria species clearly varied depending on feeding frequencies and the initial size of individuals. Females of both species had a significantly higher growth rate than males, though the difference became less marked with decreasing food supply. The total time spent on feeding and the length of the first feeding bout generally increased as the duration of the interval between meals was increased across treatments. From both the field growth experiment and the observations of feeding behaviour it is clear that J. ferrea had a higher frequency of feeding than J. cingulata in the field; 1- to 3-d intervals of feeding frequency in the former and 3- to 6-d intervals in the latter. Most notably, the two species showed a reversal in growth rates with changing conditions of food supply. When food was relatively abundant J. ferrea achieved better growth than J. cingulata, but J. cingulata grew better than J. ferrea under low food supply. In view of the fact that J. ferrea seems to be competitively superior to J. cingulata under normal levels, the present observation that J. cingulata can perform better than J. ferrea when food availability is low gives an important advantage to the latter in terms of its coexistence with the former. Received: 9 March 1999 / Accepted: 8 October 1999     

Functional mouthpart morphology of the squat lobster Munida sarsi, with comparison to other anomurans

 The mouthparts of Munida sarsi Huus, 1935, including the labrum and paragnaths, are described using LM and SEM, with special attention to the setal composition and its function. We also used high-resolution macrovideo equipment to record the feeding processes and monitor setae in action. This combination of morphological and preliminary behavioral data enabled us to arrive at a more comprehensive understanding of the functional morphology. M. sarsi was found to be an opportunist and capable of handling and eating most kinds of animal tissue. The species can also be a selective deposit feeder, sorting the sediment by quality and not by size. This sorting is performed by the setae on the medial rim of the bases of maxilla 1, maxilla 2 and maxilliped 1. Many of the other mouthpart functions ranging from food detection to ingestion are mapped with comparison to what has been found for other anomurans. Grooming was shown to be a significant function of many mouthparts, especially the endopod of maxilliped 3. In general the mouthparts closest to the mouth have the least complex movements and functional scheme, with the mandibles being able to perform only one movement, i.e. cutting or crunching food objects. Some functions are attributed to the paragnath, the endopod of maxilla 2 and the epipod of maxilliped 1. Received: 3 October 1999 / Accepted: 22 March 2000