Escape responses of four Chilean intertidal limpets to seastars

The Chilean intertidal Lottiidae limpet fauna consists of at least nine species, grouped into the monophyletic taxon Scurria and a single taxon preliminarily designated as `Lottia' orbignyi (Dall). Within the Scurria clade, the most basal species are Scurria variabilis (Sowerby), S. viridula (Lamarck), and S.zebrina (Lesson), and among the most crown groups are S. ceciliana (Orbigny) and S. araucana (Orbigny). This phylogenetic information, obtained by the analysis of molecular characters (16S mtDNA), provides a framework for a comparative study of behavioral characters. In this study, we analyze the escape responses of two basal limpet species (`L.' orbignyi and S. viridula) and two derived limpet species (S. araucana and S. ceciliana) in the presence of the limpet predatory seastar Heliaster helianthus (Lamarck) and the limpet non-predatory seastars Stichaster striatus Muller and Troschel and Patiria chilensis Lutken. Neither P.chilensis nor Stichaster striatus induced escape responses such as those observed with the predatory H. helianthus. Moreover, in the presence of H. helianthus, basal and derived limpet species differed significantly in the percentage of individuals responding, reaction time, and duration of the response. The basal species `L.' orbignyi and Scurriaviridula exhibited instantaneous and vigorous locomotor responses, whereas the derived species S. araucana and S. ceciliana displayed no locomotor responses. The same pattern was found in responses to H. helianthus homogenates. The results strongly suggest a correlation between phylogeny and anti-predator escape responses to the seastar H. helianthus. Alternative escape responses have evolved for basal and derived species within the Chilean clade. Received: 27 April 2000 / Accepted: 16 August 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     

Size-related aspects of arm damage, tissue mechanics, and autotomy in the starfish Asterias rubens

 Arm damage is a widely reported but superficially investigated aspect of the biology of the starfish Asterias rubens L. In the present study, the incidence of arm damage was surveyed in populations of A.  rubens at two sites in the Firth of Clyde, Scotland, and three sites in Gullmarsfjorden, Sweden. The mean (±SD) incidence across all sites of individuals with basal arm damage (resulting from detachment at the basal autotomy plane) was 19.69 ± 8.86%, the incidence of those with distal arm damage (resulting from amputation at more distal levels) was 7.74 ± 10.01%. The mean incidence of arms with basal damage was 5.28 ± 4.12%, of those with distal damage 1.83 ± 2.45%. There was a significant negative correlation between size and the incidence of basal damage at all but one site, but no significant correlation between size and distal damage at any site. Mechanical tests on specimens of the aboral body wall from the basal region of the arm (which included the autotomy plane) and from a more distal region revealed that with increasing body size there was a significant increase in yield stress, ultimate stress and Young's modulus (stiffness) but no significant change in yield strain and ultimate strain. There was no significant difference between the relationships for basal and distal specimens. It is hypothesised that in larger individuals increased mechanical toughness replaces autotomy as an effective antipredator strategy. Using two methods to induce autotomy, a significant positive correlation between size and the delay between the onset of stimulation and arm detachment was found; this may represent a size-related decline in the efficiency of the autotomy mechanism through the relaxation of selection pressure. Since size is an unreliable indicator of age in A.  rubens, the trends identified herein can be interpreted only tentatively as age-associated phenomena. Received: 4 September 1999 / Accepted: 17 February 2000     

Allozyme electrophoresis indicates high gene flow between populations of Holothuria (Microthele) nobilis (Holothuroidea: Aspidochirotida) on the Great Barrier Reef

 Genetic variation in 15 Holothuria (Microthele) nobilis (Selenka, 1867) populations on the Great Barrier Reef was studied at seven polymorphic allozyme loci. Although populations were separated by distances up to 1300 km, there were no apparent restrictions to gene flow (F _ST values were not significantly different from 0) and the maximum Nei's unbiased genetic distance was 0.003. Populations were in Hardy–Weinberg equilibrium at all loci. The estimated maximum sexual input [this is the ratio of the number of sexually produced individuals (N*) to the sample size (N _i )] and the minimum sexual input [this is the ratio of the number of genotypes (N _go ) over the sample size (N _i )] were used as estimators for the amount of asexual reproduction. Both parameters suggested that H. nobilis reproduces solely by sexual means (N*:N _i : = 1; N _go :N _i  = 0.74 to 1). The allozyme data indicated high gene flow between populations, but the possibility that allozyme frequencies may not be at equilibrium means that it was not possible to distinguish whether the patterns reflect present-day dispersal or dispersal that occurred in the past. Received: 27 March 2000 / Accepted: 29 June 2000     

The nature of heat coma in Littorina littorea (Mollusca: Gastropoda)

 The nature of heat coma was examined in the edible periwinkle Littorina littorea (L.). Duration of acclimation did not influence heat-coma temperature at 12 °C, although other acclimation temperatures were important in influencing thermal tolerance, with positive shifts in coma temperature observed in response to elevated temperatures. Previous thermal history also influenced heat-coma temperatures. Individuals subjected to repeat heat-coma events on a daily basis showed significant declines (P < 0.05) in coma-temperature; in contrast individuals exposed to repeat heat-coma events on a weekly basis showed no decline in thermal tolerance. Size-effects occurred at selected sites, where decreased heat-coma temperatures were recorded in large individuals. Received: 16 August 1999 / Accepted: 13 June 2000     

Patterns of time and space utilisation in coelacanths (Latimeria chalumnae), determined by ultrasonic telemetry

Acoustic telemetry was used to examine patterns of activity and space utilisation of coelacanths, nocturnal predators which spend the day in submarine caves. Nine coelacanths (Latimeria chalumnae) were tracked, each for a period of 1 to 16 nights at Grande Comore, West Indian Ocean. Activities lasted on average 9 h, usually starting shortly after sunset and ending before sunrise. Vertically, coelacanths moved up and down at and below cave level by following the bottom contour, mainly between 180 and 400 m depth. The deepest record was 698 m, the shallowest 133 m. Most time was spent between 200 and 300 m depth. Large individuals performed deep excursions to depths below 400 m, usually once per night. The fish spent most time in water temperatures of 15 to 19 °C; they rarely ventured into waters warmer than 22 °C measured at depths shallower than 160 m depth. Horizontally, coelacanths stayed in narrow areas ranging from <1 to 10 km of coastline. Coelacanths are extremely slow drift-hunters with an estimated average swimming speed of 3.2 m min⁻¹, often travelling not more than 3 km per night. They probably take advantage of local upwelling and downwelling and slow currents occurring parallel to the steep slopes. This study shows that coelacanths are inhabitants of the subphotic zone, where they are active mainly below the depth of their daytime refuges. Received: 7 July 1999 / Accepted: 11 February 2000     

Short retention times of stomach temperature loggers in free-living seabirds: is there hope in the spring?

Stomach temperature loggers have proved useful for the study of feeding activity in free-living seabirds, but their usage has been restricted because they are frequently dislodged and lost during the seabirds' regular regurgitation of indigestible prey remains. In the present study we examine the incidence of spontaneous regurgitation (pellet production) in free-living seabirds, consider the effect this has on the likely retrieval of stomach temperature loggers and present a structural modification of the logger housings which leads to a much lower incidence of regurgitation. Systems were tested on albatrosses, cormorants, gannets and penguins. Received: 15 July 1997 / Accepted: 31 October 1997     

Simultaneous hermaphroditism in the marine shrimp Lysmata wurdemanni (Caridea: Hippolytidae): an undescribed sexual system in the decapod Crustacea

Simultaneous hermaphroditism with outcrossing, a previously unconfirmed sexual system in decapod crustaceans, is documented in the shrimp Lysmata wurdemanni (Gibbes), using time-lapse video observations on mating in pairs of “female-phase” (FP) individuals. Copulations between FPs resulted in successful spawning and development of embryos. However, female-phase hermaphrodites maintained in isolation were unable to self-fertilize spawned eggs. All smaller individuals possessed characters typical of caridean males, including male gonopores, appendices masculinae, ejaculatory ducts, and cincinnuli on the first pleopods. However, the gonad of these male-phase (MP) individuals was an ovotestis with an undeveloped ovarian portion. FPs, which spawn eggs and incubate embryos, also had male gonopores and an ovotestis terminating in ejaculatory ducts containing sperm. In FPs, male pleopod characters were absent or reduced, and a female incubatory character, expanded pleopod flanges, showed a greater relative size. Smaller size classes of the population were composed only of MPs while larger size classes consisted of both MPs and FPs. “Transitionals,” individuals with MP characters but an ovotestis containing vitellogenic oocytes, were rare and overlapped completely in size with FP and larger MP individuals. It is suggested that not all MPs transform into FP hermaphrodites. Social interactions may mediate MP change into FP hermaphrodites in L. wurdemanni. The social system of L. wurdemanni and other Lysmata spp. may determine the exact form of the sexual system in these species. Received: 26 October 1997 / Accepted: 21 April 1998     

Male characteristics, sperm traits, and reproductive success in winter-spawning Celtic Sea Atlantic herring, Clupea harengus

The relationship between sperm characteristics and reproductive success was examined in male herring, Clupea harengus L. Males were categorised as being first-time or repeat spawners on the basis of their age; they were also grouped according to whether their sperm were immediately active and exhibited forward motion on contact with seawater (FM) or had little or only vibratory motion (VM). Unlike the Pacific herring C. pallasii Valencienes, Atlantic herring sperm is usually motile on contact with seawater. The age, weight and gonadosomatic index (testes mass as a percentage of somatic mass = GSI) were measured and used as characteristics for individual fish. Sperm traits measured were (1) adenosine triphosphate (ATP) concentration, (2) sperm count, (3) duration of sperm motility. Reproductive success for each male was estimated from the fertilisation rate and from the length of larvae at hatching. Fertilisation rates for all fish were generally >80%. The ATP concentration of non-activated spermatozoa was negatively correlated with fertilisation rate. Among repeat spawners, fish with higher GSIs produced larvae that were larger at hatching. Although VM sperm fertilised eggs at rates equivalent to fertilisation by FM sperm, the larvae produced by VM sperm were significantly smaller at hatching. Larval length tended to increase in parallel with the duration of sperm motility, but the relationship was not significant in these tests. The results did not indicate any age or size pattern to spawning readiness in male herring. Sperm that are not yet ready to be shed are not fully motile on contact with seawater, but are still capable of fertilising eggs that hatch successfully. There is likely to be a progression of males which come into spawning readiness within a spawning shoal; therefore it is possible that paternal influences would result in a progressive decrease in larval size over the spawning period in winter-spawning Celtic Sea herring. Received: 22 November 1997 / Accepted: 8 June 1998     

Effects of Trichodesmium spp. blooms on penaeid prawn larvae

The effects of blooms of the cyanobacterium Trichodesmium spp. on penaeid prawn larvae were examined using in situ and laboratory rearing experiments and plankton surveys in Albatross Bay, Gulf of Carpentaria, Australia. The in situ experiments demonstrated that, during a bloom of Trichodesmium spp., larvae of the prawn Penaeus merguiensis did not develop beyond the first protozoea stage, and survival was low compared with times when diatoms were dominant in the same study area. Laboratory experiments confirmed the in situ results. None of the prawn larvae fed Trichodesmium sp. in laboratory experiments developed beyond the first protozoeal stage. In contrast, 94% of prawn larvae fed the green flagellate Tetraselmis suecica successfully developed to the second protozoea stage. Electron microscopy of larvae gut-contents revealed that Trichodesmium spp. were ingested by larvae but were of no nutritional value, resulting in starvation. A 7 yr plankton survey, from 1985 to 1992, showed that minimum abundance of prawn larvae occurs during the annual summer blooms of Trichodesmium spp. and that maximum abundance of prawn larvae generally occurs just after the bloom. There was a negative correlation between the abundance of larvae and the abundance of Trichodesmium at individual sites, one offshore and one inshore, indicating that the blooms affect the survival of larvae. We conclude that variations in both timing and magnitude of Trichodesmium blooms are important determinants of prawn larvae abundance in Albatross Bay. Received: 28 April 1997 / Accepted: 2 April 1998