Differences in morphology and habitat use among the native mussel <Emphasis Type="Italic">Mytilus trossulus</Emphasis>, the non-native <Emphasis Type="Italic">M. galloprovincialis,</Emphasis> and their hybrids in Puget Sound,Washington

Mytilus galloprovincialis (Mg), the Mediterranean blue mussel, is sympatric with the native M. trossulus (Mt) throughout much of the north Pacific, likely as the result of human introduction. We investigated the distribution of the two species and their hybrids (Mgt) in Puget Sound, Washington, to determine whether differences occur in habitat preference between the two species and hybrids. In addition, we investigated whether there were consistent size and shape differences between the native and introduced mussels and hybrids. Measurements of over 6,000 mussels from 30 sites, of which 1,460 were genotyped for a species-specific genetic marker, revealed that Mg and Mgt can be found throughout Puget Sound. Mg and Mgt were larger and exhibited a greater height:length ratio than Mt. Frequencies of Mg and Mgt were higher in subtidal habitats, such as docks, than on intertidal rocks, walls or pilings. Within intertidal habitats, Mg and Mgt were more frequent than Mt in the lower reaches of the intertidal. At slightly more than half the sites the frequency of the three genotypes accorded with random mating expectations suggesting no consistent barriers to gene flow between species. The standardized random sampling methods and simple morphometric identification techniques described here can be used to test whether the frequency of invasive mussels changes over time and space in Puget Sound.     

Morphological and molecular data suggest a cosmopolitan distribution of the polychaete <Emphasis Type="Italic">Proscoloplos cygnochaetus</Emphasis> Day, 1954 (Annelida,Orbiniidae)

The discovery of a possibly invasive Proscoloplos species on the French Atlantic coast led to a detailed morphological and molecular investigation. Proscoloplos consists of three nominal species, but molecular analyses of the variable ITS1 and ITS2 region revealed no clade support for samples from South Africa, France and Australia, representing at least two of these nominal species. We found no unambiguous diagnostic characters for the three different species even with scanning electron microscopy. Two main characters—first appearance of branchiae and hooks—that were used for species delineation show considerable intrapopulational variation and do not withstand critical evaluation. The obtained data and the observed regenerative capabilities point to a cautious use of the position of branchiae and hooked chaetae as taxonomic marker in Proscoloplos. After experimental bisection of animals, the recovery led to a phenotype without recognizable signs of regeneration, but neither branchiae nor hooked chaetae reoccurred at the same initial segmental position. Summarising the molecular and morphological data we suggest the synonymy of the known Proscoloplos species under the senior synonym Proscoloplos cygnochaetus Day, 1967. The enormous geographic range of this species comprises the temperate waters of the whole southern hemisphere and includes the French population. We discuss a dispersal enabled through the effective anchorage on vessels using mucous glands and chaetae possibly combined with architomic reproduction.     

Trophic variation with length in two ommastrephid squids, <Emphasis Type="Italic">Ommastrephes bartramii</Emphasis> and <Emphasis Type="Italic">Sthenoteuthis oualaniensis</Emphasis>

From 1998 to 2001 a total of 200 Ommastrephes bartramii (27 paralarvae) and 170 Sthenoteuthis oualaniensis (14 paralarvae) were sampled from the Central North Pacific. One group of non-paralarval O. bartramii (n = 30) was sampled from farther northwest in 1996. The δ¹⁵N of mantle muscle of non-paralarval O. bartramii ( = 12.4‰) was significantly greater than that of non-parlarval S. oualaniensis ( = 8.1‰) (P < 0.001). The δ¹⁵N of whole paralarvae of O. bartramii ( = 6.4‰) was not significantly different than parlarvalae of S. oualaniensis ( = 6.1‰) (P = 0.528). There was no significant difference between the mantle muscle δ¹⁵N values of male (n = 95, = 13.3‰) and female (n = 18, = 12.9 ‰) O. bartramii greater than 300 mm mantle length (ML) (P = 0.15). There was also no significant difference between the mantle muscle δ¹⁵N values of male (n = 15, = 7.2‰) and female (n = 26, = 7.3 ‰) S. oualaniensis in the same size range (P = 0.41). Overall there was a distinct logistic increase in δ¹⁵N with mantle length for O. bartramii, whereas S. oualaniensis showed an exponential increase in δ¹⁵N with mantle length that was stronger within individual years than with all samples combined. In general, adult O. bartramii are more than a trophic level above S. oualaniensis (4.3‰, 1.3 TLs). Because of the nature of the sampling protocol, this study could not separate spatial and temporal effects on the δ¹⁵N signals from each squid species. This study demonstrates the ability of stable isotope analyses to differentiate trophic levels between squid species as well as track trophic changes across size ranges from paralarvae to adults. Additional research is needed to validate these trophic changes across size within individuals.     

Animals on marine flowers: does the presence of flowering shoots affect mobile epifaunal assemblage in an eelgrass meadow?

Eelgrass, Zostera marina, produces two types of shoots: morphologically simple vegetative shoots and highly branched flowering (reproductive) shoots, the latter found only in summer months. We examined whether the abundance and diversity of mobile epifaunal assemblage are affected by the presence of flowering shoots in an eelgrass meadow of Otsuchi Bay, northeastern Japan. Comparisons of epifauna in natural vegetation revealed that density and species richness did not differ significantly between sites consisting of both flowering and vegetative shoots, and those only of vegetative shoots. A transplant experiment, conducted to examine the colonization rates of epifauna to defaunated eelgrass planted with different combination of vegetative and flowering shoots, showed no obvious variation in abundance and species richness. At species level, the density of some species such as a tanaid Zeuxo sp. and a polychaete Platynereis sp. was higher at sites and/or treatments with flowering shoots, whereas that of some gastropods, such as Lirularia iridescens and Siphonacmea oblongata was higher at sites without flowering shoots. The species-specific response led to dissimilarity of epifaunal assemblage between sites and among treatments with different densities of vegetative and flowering shoots. Similar patterns observed for natural vegetation and the transplant experiment suggest that the variation in assemblage structure is caused by habitat selection of each species, for example, the utilization of flowering shoots as feeding ground and nursery by Zeuxo sp.     

Annual population development and production by small copepods in Disko Bay,western Greenland

The population of small copepod species (approximately <1 mm) were investigated during an annual cycle in Disko Bay, western Greenland. The small species considered were Acartia longiremis, Pseudocalanus spp., Oithona spp., Oncaea spp., Microsetella spp., and Microcalanus spp. Most of the small species were present in the surface waters year round and numerically dominated the community, and in biomass from late summer and throughout winter. Oithona spp. was numerically the main contributor, while Pseudocalanus spp. dominated in terms of biomass. In the uppermost 50 m, maximum abundance, biomass and secondary production were observed in late September after the phytoplankton production practically had terminated and the winter initiated. The free spawning Acartia longiremis showed a strong seasonal fluctuation in biomass and egg production, in contrast to the egg carrying species Pseudocalanus spp. and Oithona spp. These had a long spawning season and maintained a more stable biomass year round. Secondary production was estimated by three different ways: (1) based on the obtained specific egg production rates, (2) a temperature dependent equation, and (3) a multilinear regression taking temperature, body weight and chlorophyll into consideration. The contribution of the small species was insignificant when compared to the large Calanus species during the spring- and post-bloom. However, during late summer and winter, where Calanus had left the upper water strata for hibernation, the small species played a crucial role in the pelagic carbon cycling.     

Transient ontogenetic expression of hermaphroditic gonad morphology within the Gobiosoma group of the Neotropical seven-spined gobies (Teleostei: Gobiidae)

The Neotropical seven-spined gobies (tribe Gobiosomatini), including the Gobiosoma and Microgobius groups, constitute a speciose, monophyletic gobiid taxon. In particular, member species of the Gobiosoma group exhibit a combination of behavioral diversification and microhabitat specialization that may have played a major role in their collective rapid evolution and high rate of speciation. Functional hermaphroditism, which is common among gobiids, can promote exploitation of patchily distributed micro-niches by countering potential reductions in reproductive opportunities. However, the possible role of functional hermaphroditism in the exploitation of microhabitats and rapid speciation in the Gobiosoma group is unknown. One member species, Tigrigobius multifasciatus, is a functional hermaphrodite. Two other Gobiosoma group species are functionally gonochoric (i.e., constant-sexed), but exhibit transient hermaphroditic ovarian structure among immatures. In this study, ovarian morphology among immature and adult females of several Gobiosoma group species was examined to see if hermaphroditic gonadal features are present. Although no evidence of functional hermaphroditism in the form of precursive accessory gonadal structures (pAGS) associated with the adult ovary was found among newly examined species, all species exhibited the transient expression of hermaphroditic gonadal features associated with the immature ovary. In contrast, among six species of non-Gobiosomatini genera having no record of hermaphroditism, none exhibited similar transient hermaphroditic features associated with the immature ovary. These findings suggest that hermaphroditism may have been an ancestral trait which has been secondarily lost within some Gobiosoma group species. This study also shows that ontogenetic and morphogenic processes within the gobiid reproductive system may provide new insights into the evolution of life history traits and significantly further our understanding of the extraordinary evolutionary success of this group.     

Rapid tissue reduction and recovery in the sponge <Emphasis Type="Italic">Aplysinella</Emphasis> sp.

We observed a pronounced, yet reversible tissue reduction in the tropical sponge Aplysinella sp. under non-experimental conditions in its natural habitat, after transfer into seawater tanks, as well as after transplantation from deep to shallow water in the field. Tissue reduction resulted in the formation of small “reduction bodies” tightly attached to the sponge skeleton. Although volume loss and gain were substantial, both tissue reduction and regeneration were often remarkably rapid, occurring within few hours. Microscopic analysis of the reduction bodies revealed morphological similarities to previously described sponge primmorphs, with densely packed archaeocytes and spherulous cells enclosed by a thin layer of epithelial-like cells. Denaturing gradient gel electrophoresis (DGGE) revealed pronounced changes in the sponge-associated microbial community upon tissue reduction during laboratory and field experiments and following changes in ambient conditions after transplantation in the field. Generally, the microbial community associated with this sponge proved less stable, less abundant, and less diverse than those of other, previously investigated Verongid sponges. However, one single phylotype was consistently present in DGGE profiles of Aplysinella sp. This phylotype clustered with γ-proteobacterial sequences found previously in other sponge species of different taxonomic affiliations and geographic provenances, as well as in sponge larvae. No apparent changes in the total secondary metabolite content (per dry weight) occurred in Aplysinella sp. upon tissue reduction; however, comparative analysis of intact and reduced tissue suggested changes in the concentrations of two minor compounds. Besides being ecologically interesting, the tissue reduction phenomenon in Aplysinella sp. provides an experimentally manipulable system for studies on sponge/microbe symbioses. Moreover, it may prove useful as a model system to investigate molecular mechanisms of basic Metazoan traits in vivo, complementing the in vitro sponge primmorph system currently used in this context.     

Distribution-free two-sample comparisons in the case of heterogeneous variances

Behavioral ecologists are often faced with a situation where they need to compare the central tendencies of two samples. The standard tools of the t test and Mann–Whitney U test (equivalent to the Wilcoxon rank-sum test) are unreliable when the variances of the groups are different. The problem is particularly severe when sample sizes are different between groups. The unequal-variance t test (Welch test) may not be suitable for nonnormal data. Here, we propose the use of Brunner and Munzel’s generalized Wilcoxon test followed by randomization to allow for small sample sizes. This tests whether the probability of an individual from one population being bigger than an individual from the other deviates from random expectation. This probability may sometimes be a more clear and informative measure of difference between the groups than a difference in more commonly used measures of central tendency (such as the mean). We provide a recipe for carrying out a statistical test of the null hypothesis that this probability is 50% and demonstrate the effectiveness of this technique for sample sizes typical in behavioral ecology. Although the test is not available in any commercial software package, it is relatively straightforward to implement for anyone with some programming ability. Furthermore, implementations in R and SAS are freely available on the internet.     

Territorial counter-singing in male sac-winged bats (<Emphasis Type="Italic">Saccopteryx bilineata</Emphasis>): low-frequency songs trigger a stronger response

In many animal species, individuals compete for resources but avoid escalated conflicts by threat displays, i.e. a mutual signalling behaviour that enables the opponents to predict the outcome of the conflict without the necessity of actual fighting. For example, territory holders may use acoustic signals to communicate not only their own identity and the borders of their territory but also their competitive quality, fighting ability and motivation. Here, we show that male sac-winged bats, Saccopteryx bilineata, adjust their vocal territorial displays according to the fundamental frequency of territorial songs of their opponents. In playback experiments with territorial males, low-frequency stimuli elicited a higher territorial song rate and length than high-frequency stimuli. Male S. bilineata that sing more often and with lower fundamental frequencies have been shown to sire more offspring than their competitors. Fundamental frequency of territorial songs, hence, may reveal male quality and, consequently, the resulting threat posed to competing males. We argue that this is reflected in the increased response of competitors to low-frequency territorial songs shown here. Such competitive signalling behaviour has been shown in a few mammal species like red deer and baboons but, thus far, not in bats. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The downfall of mating: the effect of mate-carrying and flight muscle ratio on the escape ability of a pierid butterfly

In many species, the ability to evade predators is known to be periodically impaired by increased weight loads due to feeding and reproduction. Not only may extra weight reduce escape speed, but feeding and mating can also make the prey more noticeable to visually hunting predators. A number of butterfly species mate for hours, and if a mating couple is disturbed, one of the butterflies is responsible for flying, whereas its partner remains still. This study investigated the ability of male Pieris napi butterflies to fly while mating, with the prediction that mate carrying impairs flight ability compared to single flying males and that males with relatively high flight muscle ratios (FMR; male thorax mass/male + female body mass) will have better flight performance in copula. Our results clearly show that whereas single males always take off at steep angles and fly upwards, couples invariably have a negative take-off angle and rarely gain height. Moreover, landing height of the couples is positively associated with higher FMR. Hence, male flight ability when in copula is positively associated with a high relative thorax mass. Butterfly pairs may thus be at greater risk of predation as a consequence of their impaired flight ability, especially couples with critically low FMRs (<16%).