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.     

On the potential use of magnesium and strontium concentrations as ecological indicators in the calcite skeleton of the red coral (Corallium rubrum)

 The variability of magnesium, strontium and calcium concentrations in the two skeleton types (sclerites and axis) of the red coral (Corallium rubrum) was assessed by using X-ray fluorescence spectroscopy and microprobe analysis as a prerequisite for their use as ecological indicators. Axis cross sections showed light and dark circular growth bands corresponding to fast and slow growth, respectively. Using microprobe analyses the Mg and Sr concentrations ranged from 2.8 to 3.0% and from 0.21 to 0.29%, respectively, in the sclerites and from 2.4 to 3.0% and from 0.1 to 0.28%, respectively, in the axis. Mg/Ca and Sr/Ca ratios varied within sclerites (average, 12% for both ratios) and within the axis of single colonies (average, 20% for Mg/Ca and 48% for Sr/Ca). Sr/Ca ratios in the axis were lower in fast-growing branch tips than in older, more basal parts of the colonies, whereas the Mg/Ca ratio did not differ significantly between colony regions. Mg/Ca and Sr/Ca ratios in the axis decreased significantly with depth, and we estimated an increase of the Mg/Ca ratio of 0.004–0.006 per degree Celsius. In all depth layers, Sr/Ca ratios showed a significant direct relationship with skeleton density in axis cross sections, whereas there was no significant relationship for Mg/Ca. Overall, our data indicate that temperature promotes the incorporation of Mg in C. rubrum as in other calcite skeletons, whereas Sr concentrations are inversely related to growth rate. This preliminary study suggests that Mg and Sr concentrations in the axis of the red coral have a strong potential as ecological indicator for temperature and growth rate. Received: 3 March 2000 / Accepted: 30 August 2000     

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     

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     

DNA sequence analysis identifies genetically distinguishable populations of harp seals (Pagophilus groenlandicus) in the Northwest and Northeast Atlantic

Harp seals (Pagophilus groenlandicus Erxleben, 1777) comprise three populations based upon whelping areas in the Greenland Sea, White Sea, and Northwest Atlantic. The last comprises two subpopulations, one whelping in the Gulf of St. Lawrence (“Gulf ”) and one on the pack ice of the southern Labrador/northern Newfoundland coastal shelf (“Front”). A total of 40 female seals from the four whelping areas were collected during the 1990 and 1992 whelping seasons. DNA sequence variation was examined in a 307 bp region of the mitochondrial cytochrome b gene. Eleven variable nucleotide positions defined 13 genotypes: a significant fraction of the genotypic variance (F _ST=0.12, or 0.09 as measured by Weir's coancestry coefficient θ) is attributable to differentiation between Northwest and Northeast Atlantic populations. There was no significant differentiation between the two whelping areas in the Northwest Atlantic, or between the Greenland Sea and White Sea. These findings suggest significant reproductive isolation exists between trans-Atlantic breeding populations. Received: 18 January 1999 / Accepted: 22 February 2000     

Can synchronous spawning be predicted from environmental parameters? A case study of the lugworm Arenicola marina

The annual epidemic spawning period of a Scottish population of Arenicola marina (L.) has been recorded over a period of 13 yr. This population spawns between mid-October and mid-November in a discrete spawning event over a period of 4 to 5 d. Endocrine manipulation experiments showed that spawning is induced in females only if sufficient titres of PMH (prostomial maturation hormone) are present in the prostomia. These levels are attained during the 2 to 3 wk prior to the natural spawning date. The East Sands, St. Andrews population always spawns during periods of spring tides regardless of tidal amplitude or whether they are full- or new-moon tides. Meteorological data, including sea-temperature data were collected for each year, and correlation of the environmental data with spawning time was attempted. Correlation of spawning times with weather patterns showed that mean daily air pressures were significantly higher during the spawning period than from September to November as a whole. Evidence also suggests that a reduction in sea temperature is required prior to spawning. A significant moderate negative correlation was found between May to July air temperatures and spawning date, suggesting that higher May to July temperatures may induce early spawning. Daily rainfall and wind speed were also lower during the spawning period, but not significantly so. These results indicate that air pressure (or changes therein) may act as a final spawning cue, and the advantages of this are discussed in relation to fertilization success. A model of the interplay between environmental parameters and the endocrine mechanisms controlling the induction of spawning is proposed. Higher than average summer temperatures may advance gametogenesis to bring the population into a state of maturity (full-size oocytes, well-developed sperm morulae), and may also advance spawning time. Once the population has completed gametogenesis, a drop in sea temperature is then required to trigger an increase in endocrine titres within the prostomium, without which spawning cannot be induced by prostomial injection. The population spawns on spring tides; however a lack of clement weather coinciding with the spring tide will result in population-wide spawning being aborted, as in 1996. Clement weather (high pressure, low rainfall and wind speed) in conjunction with spring tides permits spawning to proceed to completion. Received: 21 June 1999 / Accepted: 25 January 2000     

Modulation of the free amino acid pool and protein content in populations of the brine shrimp Artemia spp.

 Free amino acid (FAA) and protein content were measured in various developmental stages of Artemia franciscana, from cysts to Instar III metanauplii. In addition, decapsulated cysts of 15 Artemia populations from different localities were compared with respect to their FAA and protein content. Furthermore, the content and composition of the FAA pool were modulated by hatching the cysts at various salinities, and by enriching the nauplii with algae or a lipid-enrichment emulsion. The FAA content increased threefold from cysts to nauplii, and Instar III metanauplii contained nearly 50% taurine of total FAA. Cysts of A. franciscana were found to contain one-third the amount of FAA compared to the other Artemia species investigated. The content and pool composition of FAA was successfully modulated in 11 of 13 populations, where by the content of FAA was significantly increased when hatched at high salinity. Finally, enrichment elevated the content of FAA and changed the pool composition, thereby showing a dietary effect. Algal enrichment also increased the protein content. Received: 27 September 1999 / Accepted: 17 July 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     

The effect of pup vocalisations on food allocation in a cooperative mammal, the meerkat (Suricata suricatta)

In the meerkat (Suricata suricatta), a cooperative mongoose, pups follow potential feeders while the group is foraging and emit incessant calls when soliciting food from them. In contrast to a ’stationary’ brood of chicks, in which nestlings are fed at a fixed location, meerkat pups are ’mobile’ and become spread out. The question arises whether meerkat pups that experience different constraints to those facing chicks have evolved similar begging strategies. This paper describes the vocalisations that meerkat pups emit in the context of begging and investigates the influence of these calls on food allocation by older group members and on the behaviour of littermates. Meerkat pups use two types of calls when soliciting food from a potential feeder. The most common is a ’repeat’ call, which pups emit continuously when following an older forager over several hours a day. In addition, when a potential feeder finds a prey item, the pups next to it emit a bout of calls with increased calling rate, amplitude and fundamental frequency, termed ’high-pitched’ calls. Observations, together with playback experiments, showed that more prey was allocated to pups that called longer and more intensely. The pup closest to a feeder was almost always fed. The probability of emitting high-pitched calls did not depend on the time since a pup had received food, and the change from repeat to high-pitched calls occurred suddenly. The main function of the high-pitched call, therefore, does not appear to be to signal a pup’s hunger state. More likely, the two calls, in the context of begging, may be an adaptation to energetic constraints in a mobile feeding system. Pups, which are dispersed during foraging, may emit repeat calls over long periods to prevent potential feeders from eating all the prey themselves. At the moment a potential feeder finds prey, pups may give the more intense high-pitched calls to direct feeders to bring the food item to them and not to a littermate. Therefore, unlike the stationary feeding system where chicks emit one type of begging call when the feeder approaches the nest, meerkats, with a mobile feeding system, have evolved two discrete types of vocalisations in the context of begging. Received: 22 November 1999 / Revised: 1 July 2000 / Accepted: 17 July 2000     

Utility of North Atlantic Right Whale Museum Specimens for Assessing Changes in Genetic Diversity

Abstract: We examined six historical specimens of the endangered North Atlantic right whale ( Eubalaena glacialis ) using DNA isolated from documented baleen plates from the late nineteenth and early twentieth centuries. Sequences from the mitochondrial DNA (mtDNA) control region from these samples were compared with those from a near-exhaustive survey (269 of approximately 320 individuals) of the remaining right whales in the western North Atlantic Ocean. Our results suggest that there has been only relatively modest change in maternal lineage diversity over the past century in the North Atlantic right whale population. Any significant reduction in genetic variation in the species most likely occurred prior to the late nineteenth century. One historical specimen was from the last documented female capable of propagating one of the maternal lineages in the population today; no females in the existing population have been found to carry this mtDNA haplotype. Analysis of the only specimens from the eastern North Atlantic right whale population ever to be examined revealed that eastern and western North Atlantic right whales may not have been genetically differentiated populations. Loss of gene diversity experienced by North Atlantic right whales over the last century has been modest, and the six decades of protection have been successful in maintaining much of the maternal lineage diversity that was present in the late nineteenth century.