Reproduction, larval development and functional relationships of the burrowing, spionid polychaete Dipolydora armata with the calcareous hydrozoan Millepora complanata

Dipolydora armata (Langerhans, 1880) is a small (4 to 5 mm) spionid polychaete found burrowing in the calcareous hydrozoan Millepora complanata Lamarck, 1816, on coral reefs at Barbados, West Indies. It excavates complex networks of interconnecting burrows and forms aggregations of worms in cavities within branches of the coral. Adult worms have a mixed feeding mode (suspension feeding and deposit feeding). Size–frequency distributions of worms in branch samples suggest that they mature in a single year and that reproduction occurs throughout the year. Burrow openings on the surface of the coral develop distinctive, erect spines caused by combined growth of worm tubes and host tissue. Millepore zooids were absent in the vicinity of tube openings and on spines, and thus the potential feeding surface of the coral will be reduced in heavily colonized branches. Burrows and openings were densest at the bases of millepore branches where weakening of the skeleton would be expected to occur. The absence of openings near the branch tips suggests difficulty in larval settlement there, amongst stinging zooids. Reproduction␣and larval development of the worms were examined, and a sequence of larval stages from one to 20 segments and a juvenile stage of 22 segments are described. Eggs are deposited in brood sacs attached to the burrow wall, and the larvae feed upon nurse eggs (adelphophagy). The presence of larvae and juveniles occurring free in the burrows suggests that larval development may be completed within the host coral as an alternative or in addition to a planktonic larval phase. Lack of provisional larval setae, early development of adult capillary setae, production of special spermatophores and a protracted breeding cycle in D. armata are all traits which would favour complete development within the host skeleton. Received: 6 March 1997 / Accepted: 25 October 1997     

A correction to: biological and oceanographic insights from larval labrid (Pisces: Labridae) identification using mtDNA sequences

Previously published mtDNA cytochrome b sequences for Xyrichtys novacula (Linnaeus) and two morphologically distinct types of Xyrichtys larvae (Pisces: Labridae) included human cytochrome b sequence, presumably due to sample contamination and/or poor preservation of sample DNA. Those sequences had been used to identify the two types of Xyrichtys larvae as X. novacula, but owing to the contamination, the identifications are invalid. Fresh specimens were collected: X.␣martinicensis (Cuvier and Valenciennes), X. novacula, X. splendens (Castelnau), and Xyrichtys sp. larvae of each of the two morphotypes (dorsal-forward eye and ventral-forward eye). The cytochrome b fragment was amplified from each specimen using the polymerase chain reaction. Comparison of the sequences with human cytochrome b sequence confirmed that the new sequences were not contaminated. The interspecific differences in Xyrichtys sequences were less than previously reported, but still greater than observed in many fish genera. Distance and parsimony analyses indicated that X. novacula and X. martinicensis were more closely related to each other than to X. splendens. This conclusion differs from our previous conclusion that X. martinicensis and X. splendens were more closely related to each other than to X. novacula. Distance and parsimony analyses also demonstrated that both larval morphotypes were X. novacula and, thus, our previous conclusions regarding larval ecology and oceanographic transport remain unchanged. Received: 12 March 1997 / Accepted: 24 October 1997     

Population biology of Dyopedos monacanthus (Crustacea: Amphipoda) on estuarine soft-bottoms: importance of extended parental care and pelagic movements

The suspension-feeding amphipod Dyopedos monacanthus (Metzger, 1875) is a common epibenthic amphipod that lives on self-constructed “mud whips” (built from filamentous algae, detritus and sediment particles) in estuaries of the northern North Atlantic Ocean. The population biology of D. monacanthus at a shallow subtidal site in the Damariscotta River Estuary (Maine, USA) was examined between July 1995 and July 1997. The resident population at the study site was dominated by adult females during most months of the year. High percentages of subadults were found in late summer/early fall. Often, between 10 and 20% of the adult females were paired with males, and the percentage of ovigerous females varied between 40 and 100%, indicating continuous reproduction. The percentage of parental females varied between 40 and 80% during most months, but dropped to levels below 20% during summer/early fall. The average size of amphipods on their own mud whips was ∼4 mm during the summer/early fall, after which it increased continuously to >7.0 mm in March or April, and then dropped again. In March and April, the average number of eggs and juveniles female⁻¹ was ∼100 eggs and 55 juveniles, while during the summer/early fall the average number of eggs female⁻¹ was <20 and that of juveniles female⁻¹ was <10. Many juveniles grew to large sizes (>1.4 mm) on their mothers' whips in winter/early spring but not in the summer/fall. The average number of amphipods at the study site was low in late summer/early fall (<50 individuals m⁻²), increased steadily during the winter, and reached peak densities of >3000 individuals m⁻² in April 1996 (>1600 individuals m⁻² in May 1997), after which densities decreased again. The decrease of the D.␣monacanthus population at the study site coincided with a strong increase of amphipods found pelagic in the water column. This behavioural shift occurred when temperatures increased and benthic predators became more abundant and active on shallow soft-bottoms, suggesting that D. monacanthus at the study site is strongly affected by predation. The effects are direct (by predation on amphipods) and indirect (by reducing duration of extended parental care and enhancing pelagic movements). Both extended parental care and pelagic movements are important behavioural traits of D.␣monacanthus (and other marine amphipods), and significantly affect its population dynamics. Received: 18 January 1998 / Accepted: 27 May 1998     

Factors influencing food choice by the seaweed-eating marine snail Norrisianorrisi (Trochidae)

Through two-choice gustatory experiments, a␣preference hierarchy was established␣for␣the␣herbivorous gastropod Norrisianorrisi Sowerby, with laminarialean kelps preferred over all other seaweeds. Among the kelps, laminae of Macrocystispyrifera were slightly preferred over Egregiamenziesii, and both were strongly preferred over sporophylls of Eiseniaarborea. E.arborea, the least preferred kelp, was consistently chosen over other algae common in the snail's habitat (Halidrysdioica, Dictyotaflabellata, and Pterocladiacapillacea) and over seaweeds believed to be edible and palatable based on their morphology, structure, and secondary chemistry (Endarachnebinghamiae, Mazzaellaflaccida, and Ulvalobata). The morphologies and structural toughness of tested seaweeds varied significantly as did their nutritional (% carbon, % nitrogen, C:N ratio, and % ash) contents and phlorotannin concentrations; however, snails preferred to feed on kelps regardless of nutritional content, toughness or phlorotannin concentration; and among kelps preferred to feed on the least tough species (based on penetrometer measurements), which also were those containing the lowest phlorotannin concentrations. Preference for kelp was not upheld in experiments using agarose thalli to which freeze-dried powder, of either the kelp Eiseniaarborea or non-kelp Endarachnebinghamiae was added, suggesting the destruction of attractant chemicals during the making of the artificial foods. Our data suggest that the preference of N. norrisi for kelps over other potentially edible and palatable seaweeds may not be related to nutritional content, but instead may have evolved in response to factors such as availability, habitat provision, or refuge from predation. Received: 27 September 1996 / Accepted: 7 October 1997     

Elemental (C, N and P) analysis of metamorphosing bonefish (Albula sp.) leptocephali: relationship to catabolism of endogenous organic compounds, tissue remodeling, and feeding ecology

Whole-body carbon (C), nitrogen (N) and phosphorus (P) content, and stable-isotope composition (¹³C:¹²C and ¹⁵N:¹⁴N), were followed during metamorphosis of bonefish (Albula sp.) larvae (leptocephali). Metamorphosing larvae depend entirely on endogenous carbon compounds (some of which contain N and P) as an energy source. Two fundamental questions are (1) Do the demands of extensive tissue remodeling during metamorphosis require the efficient retention of N and P during the catabolism of carbon compounds? (2) What effect does the lack of feeding have on stable-isotope composition? Our results showed that both C and N decreased by ～35 to 40%, reflecting the utilization of neutral lipid (triacylglycerols) and N-containing compounds (phosphatidylethanolamine and keratan sulfate glycosaminoglycan) as energy sources, and indicating that larvae have little or no capacity to retain N. Evidence suggested that collagen breakdown, measured as a loss of hydroxyproline content, also contributed to N loss. Stable-isotope ratios, expressed as ¹³C and ¹⁵N, showed no statistically significant differences in early and advanced metamorphosing larvae. In contrast to C and N, phosphorus was conserved during metamorphosis and most probably is utilized in the increased bone mineralization occurring in advanced larvae. We show, however, that advanced larvae are P-limited and that normal ossification is dependent upon a supply of exogenous P obtained after the resumption of feeding. The N:P ratio of 12.3 in early larvae decreased to 8.1 in advanced larvae owing to the conservation of P as N was lost. The mean ¹⁵N value in early metamorphic larvae (11.6‰) is consistent with results from other studies, and provides further support for the view that premetamorphic leptocephali feed at a very low trophic level.     

Settlement-competency periods of larvae of three species of scleractinian corals

The pattern of settlement over time of three broadcast spawning coral species (Cyphastrea serailia, Acanthastrea lordhowensis, and Goniastrea australensis) from the Solitary Islands (30°00′S; 153°20′E) was studied in 1995 and 1996 in order to determine the maximum length of time these larvae could remain in the water column and still retain the ability to settle and metamorphose. Larvae were maintained in aquaria and the number which had settled on biologically-conditioned tile pairs was monitored every 5 to 10 d. While the majority of larvae settled quickly after becoming competent, some larvae survived and settled for extended periods after spawning. Competency periods ranged from 26 d for C. serailia to 56 d for G. australensis and 78 d for A. lordhowensis. These data greatly extend the known competency periods for larvae of broadcast-spawning corals and indicate the potential for transport of broadcast-spawned coral larvae over large distances. Medium to long-distance larval dispersal of the species studied provides a mechanism for their widespread distribution in subtropical regions, on reefs which are often widely spaced and relatively isolated. Received: 27 May 1997 / Accepted: 27 November 1997     

Karyotypic characterization of the great sturgeon, Huso huso, by multiple staining techniques and fluorescent in situ hybridization

A karyotype analysis by several staining techniques was carried out on the great sturgeon, Huso huso (Linnaeus, 1758). The karyotype (2n = 118 ± 2) was composed of 42 pairs of meta-/submetacentric chromosomes and 17 pairs of acrocentrics/microchromosomes. Constitutive heterochromatin was mainly located at the centromeric regions of the acrocentric chromosomes. The biarmed chromosomes showed weak C-bands. Fluorescent staining with GC-specific chromomycin A₃ showed clearly recognizable fluorescent regions, whereas a more uniform base composition was revealed by the AT-specific 4,6-diamidino-2 phenylindole. After Ag-staining, nucleolar organizer regions could be observed on the short arms of two medium-sized submetacentrics and on two acrocentrics. Digoxigenated 28S and 5S rDNA probes, prepared from Acipenser naccarii DNA and hybridized to metaphase chromosomes, showed signals on six and two chromosomes, respectively. The telomeric sequence (TTAGGG) _n detected by FISH was located at both ends of each chromosome. Results are discussed in relation to karyotype organization and evolution in sturgeons. Received: 1 April 1998 / Accepted: 21 July 1998     

Effects of UV radiation of different wavebands on pigmentation, 15N-ammonium uptake, amino acid pools and adenylate contents of marine diatoms

Three marine diatoms Lauderia annulata Cleve, Odontella sinensis (Greville) Grunow and Thalassiosira rotula Meunier were exposed to ultraviolet (UV) radiation of different wavebands under controlled laboratory conditions (0.035 vol% CO₂, 18 °C). Several changes in the patterns of pigments in these organisms were seen depending on the waveband of UV radiation and species examined. UV-B and UV-B plus UV-A radiation led to a reduction in the overall pigment content of all three diatoms. The uptake of ¹⁵N-ammonium was less affected by 5-h UV-A (WG 320) but significantly reduced after UV-B and UV-B plus UV-A exposure. The pattern of free amino acid pools varied depending on the applied UV wavebands and the tested diatom. The main protein-bound amino acids of T. rotula decreased after 5-h UV irradiance except leucine. Contents of adenosine 5′-mono-, di-, and triphosphate (AMP, ADP and ATP) were affected differently by UV radiation; ATP values increased at the end of UV-B and UV-B plus UV-A exposure. These results have been discussed with reference to the impact of the different UV sources and the influence on the nitrogen metabolism in connection to pigments and supply with energy. Received: 13 May 1997 / Accepted: 11 October 1997     

Genetic protein variation in red mullet (Mullus barbatus) and striped red mullet (M. surmuletus) populations from the Mediterranean Sea

Starch-gel electrophoresis of allozymes was used to differentiate the two red mullet species (Mullus barbatus L. and M. surmuletus L.) in the Mediterranean Sea and, further, to investigate the genetic stock structure of M. barbatus in the eastern Mediterranean area. Twenty putative enzyme-coding loci were examined in eight M. barbatus samples caught in the Aegean and Ionian Seas (Greece) and in the Gulf of Lion (France), and two M. surmuletus samples caught in the Aegean and Gulf of Lion. A high degree of genetic polymorphism was found in both species. Species-specific electrophoretic patterns were found in PGI* and PGM*. Estimates of variance of allele frequencies among samples (F _ST) and ² analyses both revealed significant differences (P < 0.05) among the M. barbatus samples. Most of the genetic variation was among samples regardless of region. The mean value of Nei's genetic distance between the two species was 0.329. Genetic distance among M. barbatus samples was low (maximum Nei's D = 0.012), with the sample from Platania differing most from other M. barbatus samples. This is probably be due to founder effects existing at this area. These results suggest that allozyme analysis may provide important information on the genetic structure of the red mullet to ensure sustainable management of this species. Received: 7 May 1997 / Accepted: 13 October 1997     

Instrument Choice When Regulators and Firms Bargain

We compare outcomes with an emissions tax and an emissions standard when a firm and regulator engage in cooperative bargaining over the stringency of the regulation. Bargaining is motivated by giving the firm a choice of abatement technologies. If the firm's preferred technology differs from the regulator's, the first-best outcome is not an equilibrium of the traditional noncooperative game in which the regulator is a Stackelberg leader. The regulator may therefore choose to offer the firm a more lenient regulation if it agrees to switch technologies. We find that the resulting bargaining outcomes differ for a tax and a standard even though information is symmetric, and we identify conditions under which each instrument yields lower social costs.