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     

Spatial and temporal patterns of predation on seeds of the surfgrass Phyllospadix torreyi

Spatial and temporal patterns of predation on seeds of the seagrass Phyllospadix torreyi S. Watson were quantified at four sites near Santa Barbara, California, USA. Over a period of four flowering seasons during 1995 to 1998, monthly patterns of seed fall and intensity of seed predation were similar among sites, but were temporally quite variable. Abundance of dispersed seeds varied greatly both among seasons and years. Within any one year, seeds were present in the environment every month, but they peaked in abundance during the fall months following the annual flowering period. Seeds were more abundant during the earlier years of the study. The intensity of predation steadily increased throughout the study period, from a low of ≤10% seeds consumed during 1995 to ∼50% consumption by 1997, and it was not correlated with abundance of seeds in the environment. Pre-dispersal seed loss also was estimated in two flowering seasons by counting the numbers of seeds consumed prior to release from the plant, and was relatively low (<15%). No differences were detected among the four study sites in patterns of pre-dispersal loss. Field surveys were done at two sites to identify potential seed predators. Three of the most abundant species identified in these surveys were tested in the laboratory to determine size-specific patterns of feeding activity and to assess which early life stages of P. torreyi (fruits within spadices, dehisced fruits, seedlings) were most vulnerable to predation. All three of the early life stages were consumed. The crabs Pugettiaproducta (Randall) and Pachygrapsus crassipes Randall were much more voracious predators than the isopod Idotea resecata Stimpson. Received: 8 June 1999 / Accepted: 2 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     

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     

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     

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     

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     

Characteristics that influence male reproductive success on a lek of Lethrinops c.f. parvidens (Teleostei: Cichlidae)

Sexual selection has long been proposed as a mechanism leading to the diverse cichlid (Teleostei: Cichlidae) fauna of Lake Malawi, Africa. Many of the shallow-water, sand-dwelling, bower-building cichlid species are particularly well suited for studies of sexual selection because they participate in leks. Since females in lekking systems appear to acquire only genetic material from their mates, it has been suggested that leks are ideal systems to study female mate choice. The objectives of the investigation were to examine Lethrinops c.f. parvidens male bower characteristics (i.e., bower size and location) as well as other male characteristics (i.e., length, gular color, and duration on the lek) for their influence on male mating success as measured by the number of visits, circles, and eggs laid by females. These measures are nested in that a visit by a female may or may not lead to circling, and circling by a female may or may not lead to egg-laying. We found increased bower height and higher numbers of conspecific neighbors (analogous to shallow-water, near-shore bower positions) to be positively, significantly associated with the number of visits by females. The only significant correlate with the number of circles was visits, and similarly circles was the only significant correlate with the number of eggs laid. The R ² value for the egg-laying regression was quite low (19.8%) compared with visits (54.3%) and circling (78.9%), suggesting that females may be using additional cues, that we failed to measure, when in close proximity to males or simply that a small proportion of the females were ready to spawn. Both indirect selection and direct selection pressure due to egg predation may have influenced female choice on the lek. Received: 10 April 1999 / Received in revised form: 26 July 1999 / Accepted: 18 September 1999     

Multiple song repertoire characteristics in the willow warbler (Phylloscopus trochilus): correlations with female choice and offspring viability

Male willow warblers have song repertoires which vary in complexity along several dimensions. We examined whether female choice, as measured by date of pairing, was based on these song characteristics in 4 different years. Pairing date was negatively correlated with song repertoire size in 1 year, and with song versatility in another year, but there was no consistent effect of any song characteristic on pairing throughout the years or in the pooled sample. The variable that best explained how soon a male pairs is male arrival date (only males that had settled territories before the first female arrived were considered in the analysis). This correlation is consistently significant in all years. This is most parsimoniously interpreted as females choosing some habitat characteristic in the same way that males do. A small percentage of males (8.3%) attracted and paired with a second female. The likelihood of becoming polygynous was not explained by any measured song characteristic, but it was related to arrival date: early males were more likely to pair with two females. Males with large repertoires fledged more young in their primary nests, and there was a trend for the offspring of these males to have a greater probability of being recruited into the population. In conclusion, the results show that in most years there is no sexual selection by female preference on song characteristics, although the data on reproductive success is consistent with the idea of repertoire size being an indicator of male quality. Received: 4 June 1999 / Received in revised form: 1 December 1999 / Accepted: 31 December 1999