Reef coral reproduction in the eastern Pacific: Costa Rica, Panamá and Galápagos Islands (Ecuador). IV. Agariciidae, recruitment and recovery of Pavona varians and Pavona sp.a

The reproductive ecology of two eastern Pacific zooxanthellate coral species was examined as part of a continuing series of studies relating bleaching/mortality events caused by the El Niño–Southern Oscillation disturbance, and is described for study sites in Costa Rica, Panamá, and the Galápagos Islands (Ecuador). This study deals with the sibling agariciid species Pavona varians and Pavona sp.a over a 13?yr period (1985 to 1997). Both Pavona species are broadcast-spawners with some gonochoric, but mostly sequential hermaphroditic colonies. Minimum colony sizes (and ages) at first reproduction were 5?cm (5?yr) and 3?cm (2 to 3?yr), respectively, in P. varians and Pavona sp.a. In the Panamá and Galápagos populations, gonochoric colonies spawn eggs or sperm at least monthly. Six fecundity attributes were not significantly different in the two species, but the eggs of P. varians are white to beige and positively buoyant, and those of Pavona sp.a are dark green and neutrally to negatively buoyant. Eggs of both species lack zooxanthellae. Both species are reproductively active year-round, with maximum activity in the dry season in the nonupwelling Gulf of Chiriquí, and in the wet season in the upwelling Gulf of Panamá. Spawning is predominantly during full moon, and possibly also at new moon at most study sites. Spawning in P. varians and Pavona sp.a is 12?h out of phase, with the former species spawning ～1?h before sunrise and the latter about 1?h after sunset. The fecundity of Pavona spp. at Caño and the Galápagos Islands was much greater (19?900 to 27?900 eggs cm^?2?yr^?1) than at all Panamá sites (14?800 to 19?800 eggs cm^?2?yr^?1). Intraspecific crosses in both species resulted in swimming planula larvae after 25 to 36?h. Recruitment of P. varians was highest in Panamá, moderate in Costa Rica, and nil in the Galápagos Islands, matching, respectively, the contributions of P. varians to the pre-1982/1983 El Niño coral-population abundances in these areas. Recruitment success of P. varians at Uva Island was significantly related to maximum monthly positive sea surface-temperature (SST) anomalies that occurred in the year preceding recruitment over the period 1982 to 1996; recruitment failed when SST anomalies exceeded 1.6 to 1.9?C° during the severe ENSO events of 1982/1983 and 1997/1998.     

Metabolic differences between “demersal” and “pelagic” development of the Antarctic sea urchin Sterechinus neumayeri

 Early development of the Antarctic sea urchin Sterechinus neumayeri was examined under two differ-ent culture regimes: one to simulate development near-bottom (“demersal development”) and the other to simulate the development of embryos in the water column (“pelagic development”). When embryos of both treatments reached the hatching blastula stage at 5 d post-fertilization (−1.5 °C), the blastulae that had undergone demersal development evidenced significant differences (by ANOVA or suitable non-parametric comparison) in the following: a thicker blastoderm layer (12%, P < 0.001), higher ash-free dry weights (19%, P < 0.01), lower mass-specific respiration rates (50%, P < 0.001), higher incorporation rates of ³⁵S-methionine into protein (23%, P < 0.003), and a differential pattern of protein synthesis. When embryos developed demersally, they remained in the jelly-coat material released with the eggs at spawning. Quantitative isolation of this jelly-coat material in S. neumayeri demonstrated that it contained a significant amount of organic matter, 115 ng ash-free dry mass per egg, equivalent to 17% of the egg's initial organic mass. Uptake of external nutrients during embryogenesis may be a significant component of the physiological energetics of this polar invertebrate by allowing the utilization of jelly-coat material released by a female during spawning. Received: 21 April 1999 / Accepted: 5 June 2000     

Comparative morphology and cytology of siphons and siphonal sensory organs in selected bivalve molluscs

 The form, anatomy and cytology of the water-passages in siphon-possessing and siphon-lacking species of a selected group of bivalve molluscs from the Red Sea (Callista florida, Circe currogata, C. crocea, Lucinia dentifera, Dosinia histrio, Pitar hebraea, Tridacna maxima, Pteria aegyptica), and from the Mediterranean Sea (Spondylus spinosus, Pinctada radiata, Pinna nobilis, Donax trunculus, D. semistriatus, Mactra stultorum, Tapes decussatus, Petricola lithophaga, Brachidontes pharaonis) were studied by means of light and electron microscopy. In the mytiloids, ostreoids and ptrioids studied, the water-passages are covered by a ciliated epithelium and the few tentacles around their openings are identical to those found on the mantle edges. Contrary to this, in the veneroids studied, the siphons are covered by a microvillar epithelium and their tentacles, especially of the inhalant siphon, reveal characteristic branching. The so-called Type I, Type II and Type III ciliated sensory organs on and within the siphons are similar to those observed in other bivalves, differing from each other in the dimension, number and form of cilia. In the studied mytiloid species only Type I and Type II organs were found. The form and structure of the siphons and the distribution of sensors on them can be used as markers for ecological as well as phylogenetic considerations. Received: 28 January 2000 / Accepted: 28 May 2000     

Parthenogenetic reproduction of Diaphanosoma celebensis (Crustacea: Cladocera): influence of salinity on feeding, survival, growth and neonate production

 Effect of salinity on the feeding rate and parthenogenetic reproduction of asexual females of the cladoceran Diaphanosoma celebensis Stingelin was studied. Short-term (10 h) grazing experiments were conducted using Isochrysis galbana as feed at 5, 17, 25 and 30 psu salinity. Gut pigment concentration showed a significantly higher rate of feeding at lower salinities. Survival, growth, maturity attainment and neonate production of asexual females reared in the above four test salinities indicated preference for lower salinities (5 and 17 psu). The mean size of adult females decreased from 909 to 593 μm, mean life span from 24 to 5 d, mean neonate production from 12 to 2 and mean size of neonates from 434 to 400 μm as the salinity increased from 5 to 30 psu. Salinity variations also affected the size and age of primiparous females. Resting egg formation and sexual reproduction did not occur at the tested salinities. The results indicate that D. celebensis is adapted to low saline, estuarine environments. Received: 14 January 2000 / Accepted: 24 March 2000     

Genetic differentiation in a pelagic crustacean (Meganyctiphanes norvegica: Euphausiacea) from the North East Atlantic and the Mediterranean Sea

Meganyctiphanes norvegica (M. Sars)is a pelagic crustacean that plays a key role in marine food webs of North Atlantic Ocean and marginal seas. We studied eight population samples collected in the European Atlantic and Mediterranean Sea. By means of single strand conformation polymorphism analysis (SSCP) and direct sequencing, we investigated a segment of 158 base pairs of the mitochondrial gene coding for the subunit 1 of NADH dehydrogenase. We found 12 sequence variants among the 385 individuals studied. Analysis of molecular variance (AMOVA) showed that 14.75% of the total genetic variability was explained by differences between populations, thus indicating absence of panmixia for these populations. Pairwise comparisons revealed three distinct genetic pools: the first one represented by Cadiz Bay, the second one by the Ligurian Sea, and the third one included all the NE Atlantic samples. We also investigated one population from the Alboran Sea (within the Mediterranean basin, east of the Strait of Gibraltar). This population was found to be genetically intermediate between the NE Atlantic samples and the Ligurian sample, suggesting that the restriction to the gene flow is not associated with the Strait of Gibraltar, but possibly with the Oran–Almeria oceanographic front. The present work indicates that M. norvegica, although endowed with a high dispersal capacity because of its pelagic habit, can develop separate breeding units inside the same oceanic basin (the Atlantic). Furthermore, the Ligurian sample should be considered as a distinct evolutionary entity, separated from the Atlantic population. Received: 2 May 1999 / Accepted: 26 November 1999     

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     

Interannual, seasonal, local and body size variations in reproduction of the prawn Penaeus (Marsupenaeus) japonicus (Crustacea: Decapoda: Penaeidae) in the Ariake Sea and Tachibana Bay, Japan

 The reproductive biology of female Penaeus japonicus Bate was investigated in the Ariake Sea and Tachibana Bay (located outside the Ariake Sea), Japan from 1994 to 1996. Interannual, seasonal, individual female body size and spatial influences on the incidence of spawning were examined. The proportion of inseminated females, increased with increasing body size up to 170 mm body length (BL) and decreased thereafter. The minimum size at maturity was similar between years; 130 to 140 mm BL. The minimum size of individuals with developing ovaries or spermatophores differed from that of ripe females by 20 to 25 mm. Spawning occurred in the central part of the Ariake Sea and Tachibana Bay but rarely in the inner part. P. japonicus had a clear reproductive cycle. Spawning started earlier and ended later, occurring from mid-May to mid-October, in the eastern central part of and outside of the Ariake Sea compared with the western central part, where spawning occurred from mid-June to mid-September. Spawning dynamics differed across seasons, body sizes and areas but not across years. The seasonal peak in the proportion of ripe females varied with body size. In small individuals (130 to 169 mm BL), no peak was observed, whereas in large individuals (>170 mm BL) the proportion of ripe individuals peaked in June. The proportion of ripe individuals increased with increasing body size and was high outside the Ariake Sea. Factors causing the variation in spawning dynamics are discussed. Received: 30 November 1998 / Accepted: 12 October 1999     

Predation on meiofaunal and macrofaunal invertebrates by western sandpipers (Calidris mauri): evidence for dual foraging modes

Western sandpiper (Calidris mauri) predation was examined by concurrent experiments and direct observations of foraging behaviour on high intertidal mudflats of the Fraser River estuary, British Columbia. Western sandpipers foraged by either “pecking” on the surface (64% of observational time) or probing into sediment (29%). The first experiment (probe-mark method) consisted of collecting small-volume cores (21.2 cm³) of probed (experimental) and non-probed (control) sediment on the tidal flat, following a 22.5-min feeding period. The second experiment (exclosure method) involved deploying exclosures immediately prior to the feeding period and subsequent collection of cores from inside (control) and outside (experimental) the exclosures. Sediment cores were analysed for both macrofaunal and meiofaunal size fractions. Comparisons between macro- and meiofaunal invertebrate densities in experimental and control sediments revealed significant differences, attributed to shorebird predation, for both experiments. The probe-mark experiment detected the removal of large infaunal polychaetes (∼ 20 mm), while the exclosure experiment showed depletion of epifaunal harpacticoid copepods (0.063–0.5 mm). Predation on macrofaunal cumaceans was detected in both experiments. Invertebrates selected by western sandpipers neither fell within traditional infaunal size classifications (macro- vs. meiofauna; 500 μm delineation) nor corresponded to the highest densities of taxa. Rather, inference from experimental results and observations is that western sandpipers forage in two modes, by: (1) surface gleaning of epibenthic copepods and cumaceans in the macro- and meiofaunal size ranges and (2) selective probing for larger infauna, such as polychaetes. These findings were facilitated by the combination of methodologies employed. Received: 29 December 1999 / Accepted: 11 September 2000