The demographic consequences of the cost of reproduction in ungulates

The cost of reproduction can generate covariation between demographic rates that can potentially influence demography and population dynamics in long-lived iteroparous species. However, there has been relatively little work linking the survival cost of reproduction and population dynamics. The apparent scarcity of information on this important link is potentially due to covariation between vital rates, which can substantially influence fluctuations in population size. In this paper we examine the opportunity for survival costs of reproduction to leave a dynamic signature using a simulation model based broadly on an ungulate life history. We find that an increase in the cost delays the onset of reproduction and reduces reproductive rates of young, but not of prime-age, females. Accordingly, the number of offspring produced declines and the interval between reproductive events increases among young females experiencing high cost. These effects are translated to an age structure skewed toward young ages and reduced population density. These results suggest that, by delaying reproduction when conditions deteriorate, females protect their survival during the critical first three years of life, after which the negative effect of reproduction on survival declines. Unless conditions for reproduction are severe, it is not profitable to delay reproduction beyond age 3 years due to the high risk of death before having a chance to reproduce. We also demonstrate that lack of adjustment of reproductive strategies to elevated levels of the cost of reproduction, for example due to rapid changes in environmental conditions, results in lower average density and longevity compared to females that have sufficient time to adjust to changes in the cost. This suggests that even moderate costs of reproduction may have a major negative effect on population dynamics of ungulates.     

Structure of a Mediterranean cryptobenthic fish community and its relationships with habitat characteristics

In the last 10 years, several studies have been carried out on the fish fauna of the Ustica Island marine reserve, yet no investigation was specifically addressed to the cryptobenthic fish assemblage. The first task of this study, conducted along the shallow rocky reefs of Ustica, was to determine the species composition, diversity and relative density of the resident cryptobenthic fishes. Furthermore, we aimed to assess the effects of some macro- and microscale habitat characteristics on the distribution pattern of fishes. In particular, the effect of predator density was indirectly evaluated by comparing density data collected within and outside the integral reserve zone. Overall, 20 species belonging to Blenniidae, Gobiidae, Tripterygiidae, Scorpaenidae and Gobiesocidae were recorded. Gobius bucchichi, Scorpaena maderensis, Tripterygion delaisi, T. melanurus and T. tripteronotus were the numerically dominant and most common species. The effects of zone (i.e. of predator density), bottom type and depth on species richness, diversity and evenness were not significant. A greater total fish density was observed on stones compared with rocky cliff and plateau, but only in the shallowest depth range. At level of single species, G. bucchichi was more abundant inside than outside the integral reserve, but only on stones and at 0–2 m depth range. Density of G. bucchichi was generally higher on stones than on rocky cliffs or plateau and between 0 and 5 m depth, although these differences were not always significant. T. delaisi was conversely more abundant in the deepest stratum (7–10 m). Canonical analyses demonstrated that bottom type and depth influenced significantly the fish assemblage structure. The observed differences in the assemblage structure relied mainly upon the dominant species. T. tripteronotus was mainly associated with rocky plateau and the intermediate depth range (3–5 m), whereas S. maderensis, T. melanurus and Lipophrys trigloides inhabited preferentially the rocky cliffs. At microscale level, the habitat choice of the investigated species was almost entirely based on whether the substrate was either vegetated or composed of bare rock. T. delaisi and T. tripteronotus were associated with substrata covered by algae, whilst G. bucchichi, S. maderensis and T. melanurus preferred bare rock bottoms. In some species, the electivity indices for the less abundant type of cover, measured at different spatial scale, changed accordingly. For instance, the smaller the size of the sampled area, the higher was the intensity of the association between G. bucchichi and Anemonia viridis.     

Seasonal rhythm in a Mediterranean coastal fish community as monitored by a cabled observatory

To adequately describe seasonal rhythms in habitat utilization by fish communities, observations are needed that occur at a relatively high frequency and over large temporal windows. For the first time, images collected from a cabled video-observatory (the western Mediterranean expandable SEAfloor OBservatory; OBSEA) were utilized to examine habitat utilization by coastal fishes. Over the course of a year, hourly digital images of the water column and an artificial reef were obtained during daylight hours. For each image, the total number of species, an estimate of the abundance of individuals, and the Shannon Diversity Index were quantified. A total of 22 fish species commonly associated with nearshore western Mediterranean habitats were identified, and significant spatial (water column vs. artificial reef), seasonal (spring, summer, autumn, and winter), and daily (morning, midday, and sunset) differences in habitat utilization occurred. Four species (Diplodus vulgaris, Diplodus cervinus, Diplodus sargus, and Scorpaena porcus) were associated with the artificial reef, while one species (Pagrus pagrus) chiefly occurred in the water column. Chromis chromis and Diplodus annularis occurred at the site more frequently in the winter and autumn, respectively, while 14 other species utilized the site more frequently in either the spring (8 spp) or summer (6 spp). In addition, Dentex dentex and Spicara maena occurred more frequently at sunset, while D. cervinus displayed a crepuscular rhythm (occurring more frequently in the morning and at sunset). Species diversity was highest in the summer and lowest in the winter. If not taken into account when planning in situ sampling, such seasonal and/or diel differences may lead to spurious estimates of population sizes and biodiversity. We suggest that cabled video-observatories offer a non-invasive and reliable technology for faunistic sampling and population assessment in coastal water of the Mediterranean and likely elsewhere.     

Year-round reproduction in a seasonal sea: biological cycle of the introduced ascidian Styela plicata in the Western Mediterranean

The widely introduced ascidian Styela plicata is very common in the Western Mediterranean, an area that can act as a source for secondary introductions due to its high shipping activity. In order to understand the potential of this species to colonize new habitats, its reproductive features were assessed in the Western Mediterranean by means of monthly monitoring of two populations (Vilanova i la Geltrú 41°12′53″N, 1°44′11″E; Blanes 41°40′29″N, 2°47′56″E) from January 2009 to December 2010. The reproductive activity of this species was assessed through gonad histology and a gonad index. Population size-structure was measured monthly in order to study recruitment dynamics. No clear seasonal pattern was observed, and mature gametes and recruits were present all year long. Spawning was potentially continuous, although it seemed punctuated with pulses of gamete release, particularly in spring. A prolonged reproductive period is likely to confer a competitive advantage on S. plicata in temperate seas, where most species reproduce seasonally, and may promote recurrent introductions as larvae are available for settlement on transport vectors over much of the year.     

High self-recruitment levels in a Mediterranean littoral fish population revealed by microsatellite markers

Self-recruitment rates are essential parameters in the estimation of connectivity among populations, having important consequences in marine conservation biology. Using ten highly polymorphic microsatellite loci, we estimate, over 3 years, the self-recruitment in a population of Tripterygion delaisi in the NW Mediterranean. Six previously described source populations were used for the assignment (Costa Brava, Columbretes, Formentera, Cabo de Palos, Cabo de Gata and Tarifa). Even though this species has a 16–21 day larval duration, a mean of 66.4 ± 1.4% of the recruits settled in their natal population. When refining in a more local scale the origin of individuals self-recruited to Costa Brava, using as source the three sampling localities that conform this population (Cap de Creus, Tossa and Blanes), the highest percentage (40.6 ± 8.9%) was self-assigned to the adult source locality (Blanes) where recruits were sampled each year. Our results suggest that a high proportion of the larvae of T. delaisi remained close to, or never leave, their natal spawning area. This observation can be extrapolated to other species with similar early life-history traits and low adult mobility and can have important implications for the conservation and management of Mediterranean littoral fishes.     

Predation on meiofauna by juvenile fish in a Western Mediterranean flatfish nursery ground

The authors investigated the gut contents of juveniles of three flatfish species (Buglossidium luteum, Arnoglossus thori and A. laterna) collected from Banyuls Bay, Western Mediterranean, from January 1981 through June 1982. Young solenettes, B. luteum, appear to be dependent on meiofauna, particularly harpacticoid copepods, as prey during their first year of benthic life. By comparison, the scaldfish, especially A. laterna, prey primarily upon peracarids from the beginning of their benthic life. Evidence of prey selection exists for the genera Photis (amphipod) and Longipedia (harpacticoid copepod). By regrouping these results with those of Bodiou and Villiers (1978/1979) on the goby Deltentosteus quadrimaculatus, and comparing them with other data from the literature, it appears that the impact of predation by juvenile fish is significant for some meiofaunal species, providing that they are actively selected by the predators, as is Longipedia scotti. Generally, however, predation impact on the meiofauna as a whole is weak.     

Parental care and sexual size dimorphism in wasps and bees

Sexual size dimorphism, in which one sex is larger than the other, occurs when body size has differential effects on the fitness of males and females. Mammals and birds usually have male-biased size dimorphism, probably because of strong sexual competition among males. Invertebrates usually have female-biased size dimorphism, perhaps because their inflexible exoskeletons limit ovary size, leading to a strong correlation between female body size and fecundity. In this paper, we test whether an additional factor, the type of parental care provided, affects the degree of sexual size dimorphism. Among wasps and bees, there is a contrast between provisioning taxa, in which females must gather and transport heavy loads of provisions to nests they have constructed, and non-provisioning taxa, in which females lay eggs but do not construct nests or transport provisions. Males have no role in parental care in either case. An analysis of British wasps and bees shows that provisioning taxa have significantly more female-biased size dimorphism than non-provisioning taxa. This is true for simple cross‑species comparisons and after controlling for phylogeny. Our data imply that the demands of carrying provision loads are at least part of the explanation for this pattern. Thus, sexual size dimorphism is greatest in pompilid wasps, which carry the heaviest prey items. Bees, which transport minute pollen grains, exhibit the least dimorphism. We also find that cavity‑nesting species, in which nest construction costs may be minimized, exhibit reduced dimorphism, but this was not significant after controlling for phylogeny.     

Food resource partitioning in a Mediterranean demersal fish assemblage: the effect of body size and niche width

We investigated the effects of body size, feeding strategy and depth distribution on the trophic resource partitioning among the 26 dominant fish consumers in a fish assemblage on the central Mediterranean shelf-break. The fish assemblage was structured in two major trophic guilds: epibenthic and zooplanktonic feeders, according to the position of each predator along the benthos–plankton gradient. Within each main guild, the species were segregated along a prey-size or fish-size gradient into five further guilds. Fish size and prey size were strongly correlated, indicating that the prey-size niche can be well explained by predator size. Fish consumers showed a significant negative correlation between the similarity in prey type and the similarity in depth distribution; most species with similar trophic preferences segregated along the depth dimension. The only predators overlapping in both food and depth preferences were those with a more specialist trophic behavior. These results suggest that fish body size and depth preferences are the two main niche dimensions, explaining a large part of the coexistence between the Mediterranean shelf-break fish consumers.     

Temporal patterns of settlement, recruitment and post-settlement losses in a rocky reef fish assemblage in the South-Western Mediterranean Sea

Post-settlement events can significantly alter the density distribution of settlers and subsequently the adult population structure. The temporal and inter-annual variability of settlement and the effects of mortality on recruitment were investigated across 2 years in the vicinities of Cabo de Palos–Islas Hormigas Marine Reserve by visual census and light trap sampling. Settlement was seasonal with greater species richness and abundance in summer. Although temporal synchronization was observed between larval supply and settlement, densities of settlers could not be predicted from post-larval abundances. Timing of settlement was consistent between years but with high inter-annual variation in abundance. High mortality (~80 %) and general decoupling between post-larval and settlement stages suggest that early mortality is driving such patterns. Nevertheless, indications of habitat-mediated mortality were found for benthic species such as Symphodus and Diplodus, highlighting the importance of habitat in shaping population demography.     

Sexual and asexual reproduction in two species of Tethya (Porifera: Demospongiae) from a Mediterranean coastal lagoon

The sexual and asexual phases of reproductive cycles of two sponges, Tethya citrina and T. aurantium, living sympatrically in a Mediterranean coastal lagoon (Stagnone di Marsala, NW Sicily) were studied from samples collected over an 18-mo period. Both species are oviparous and gonochoric. They have a summer, partially overlapping, period of oocyte production, although T. citrina appear to mature earlier. No males were found, possibly due to the very short period of spermatogenesis. Both species produce asexual buds during the autumn/winter months. However, they seem to follow different reproductive strategies, with T. citrina showing a significantly lower production of buds than T. aurantium; by contrast, egg production is significantly lower in the latter species. The difference in the reproductive resource allocation is consistent with data reported in the literature on the anatomy features, genetic population structure and ecological distribution.     

Algal blooms decrease care but increase egg survival in a fish with paternal care

Human-induced eutrophication, resulting in algal blooms and increased water turbidity, is an alarming problem in aquatic systems. Here, we experimentally tested the impact of algal turbidity on parental care, egg fanning, and time in the nest, in the sand goby, Pomatoschistus minutus, a fish with uniparental male care. We allowed males to care for their eggs in either clear water or water made turbid by planktonic algae. In the early brood cycle, males fanned their eggs less in turbid than in clear water, but this difference disappeared later. Despite decreased care, egg survival was higher in turbid conditions, indicating that early fanning may partly be redundant for egg survival and perhaps used more as courtship. Males also spent more time out of their nest in turbid water, perhaps as a means to encounter additional females under conditions of low visibility.     

Ventilation or nest defense—parental care trade-offs in a fish with male care

Brood guarding animals face many critical trade-offs. Sand goby males (Pomatoschistus minutus) build nests with larger openings during low oxygen conditions, presumably to enhance ventilation. However, this may make the nest easier for egg predators to detect and harder for guarding males to defend. Manipulating oxygen level and predator presence (a small crab) for small and large males, we found support for a parental trade-off between fanning and nest defense. An increased fanning activity resulted in less time for guarding. Small males and males in low oxygen showed a higher fanning expenditure than large males and males in high oxygen, but surprisingly, filial cannibalism did not differ between these groups. Males built larger nest openings in low than high oxygen. However, males in both high and low oxygen treatments reduced their nest opening size in the presence of a predator, again indicating an important trade-off between ventilation and nest defense.     

Effects of fish feeding by snorkellers on the density and size distribution of fishes in a Mediterranean marine protected area

Although there is a great deal of evidence to show that supplementary feeding by humans in terrestrial environments causes pronounced changes in the distribution and behaviour of wild animals, at present very little is known about the potential for such effects on marine fish. This study evaluated the consequences of feeding by snorkellers on fish assemblages in the no-take area of the Ustica Island marine protected area (MPA; western Mediterranean) by (1) determining if reef fish assemblage structure is affected in space and time by tourists feeding the fish; (2) assessing the effects of feeding on the abundance of the most common fish species; and (3) assessing the effects of feeding on the size structure of the two most numerically dominant ones. In particular, we hypothesised that both the abundance and the size structure of some fish species would increase at the study site following supplementary feeding, since the additional food provided by humans would make the site more appealing to them. Fish feeding influenced the fish assemblages within the Ustica MPA, and significant spatio-temporal changes occurred. While fish feeding appeared to have no effect on the ornate wrasse Thalassoma pavo, there was a noticeable increase in the number of Oblada melanura and Epinephelus marginatus in the impacted location after feeding. It is very likely that aggregations of fishes that evolve as a result of fish feeding by the public may have negative effects on local populations of fishes and invertebrates that make up their prey. Recreational use of coastal areas and MPAs is increasing elsewhere, making fish feeding a generalised human activity. Accurate information about its effect on the fish assemblage is essential to make responsible management decisions.Communicated by R. Cattaneo-Vietti, Genova     

Parental care of nestlings by male red-winged blackbirds

Summary Nestling feeding by males is less common among birds with polygynous mating systems than in monogamous species, because of the pronounced trade-off between parental behavior and the attraction of additional mates. In this study, however, we found that male red-winged blackbirds (Agelaius phoeniceus) commonly assisted females in feeding nestlings in several Ontario marshes. Male parental care was additional to that provided by females, and it significantly enhanced the fledging success of nests (Table 2). Male redwings did not help to feed all nests on their territories: primary and secondary nests were much more likely to receive male parental care than tertiary and later nests. Contrary to expectation, male parental care was not restricted to the nests of primary females: a greater proportion of secondary than primary nests were assisted (Tables 4a and b). The presence of new females settling on the territory at the same time that a resident female had nestlings, resulted in males deferring parental care until a later brood. This suggests that males trade off the recruitment of females against parental care to an existing brood. Although the number of nestlings fledging from a male's territory was strongly influenced by the number of females in the harem, males could additionally increase their reproductive success by feeding nestlings in one or more nests on their territories (Fig. 2). The reproductive success of females was significantly enhanced by male assistance in feeding nestlings (Table 3). However, those females not receiving male assistance on territories of feeding males did not suffer a significantly reduced reproductive success in comparison to females on territories of non-feeding males (Table 2). Males varied considerably in the quality of brood care given. We therefore suggest that the quality of male parental care may be a factor considered by females in choosing a breeding situation.     

Rapid changes in growth,condition, size and age of small pelagic fish in the Mediterranean

Since 2007, the ecosystem of the Gulf of Lions has shifted to a different regime, characterised by a low anchovy (Engraulis encrasicolus) and sardine (Sardina pilchardus) biomass and a remarkably high sprat (Sprattus sprattus) biomass. Surprisingly, the abundance and recruitment of anchovy and sardine remained high. To understand which processes (bottom-up or top-down control, etc.) could have caused this shift, we studied the changes in body condition, growth and size and age of anchovy, sardine and sprat over 1984–1985 and 1992–2012, using data from scientific surveys. The annual age structure of anchovy and sardine was estimated using Bayesian mixture models based on size frequency data with priors on the age–length relationship derived from independent otolith readings. The results indicated periods during which anchovy and sardine were in an average (1992–2004), good (2005–2007) or poor (2008–2012) overall state of condition. For sardine, the shift towards smaller fish observed during these past 4 years was explained by a combination of slower growth and the disappearance of older individuals (ages 2+). Despite the increase in biomass of sprat since 2008, indications were found that sprat was also smaller than in the past. As growth and condition decreased and overexploitation has not been documented or suspected for those three species in this area, we propose that the current decline in sardine and anchovy biomass could be due to qualitative and/or quantitative modifications in the planktonic production (i.e. a bottom-up control) or mass mortalities of adults due to an epidemic disease.     

Consequences of paternal care on pectoral fin allometry in a desert-dwelling fish

Positive static allometry is a scaling relationship where the relative size of traits covaries with adult body size. Traditionally, positive allometry is thought to result from either altered physiological requirements at larger body size or from strongly condition-dependent allocation under sexual selection. Yet, there are no theoretical reasons why positive allometry cannot evolve in fitness-related traits that are solely under the influence of natural selection. We investigated scaling and sexual dimorphism of a naturally selected trait, pectoral fin size, in comparison to a trait important in male–male combat, head width in natural populations of a fish, the desert goby Chlamydogobius eremius. Male desert gobies provide uniparental care and use their pectoral fins to fan the brood (often under hypoxic conditions); hence, larger fins are expected to be more efficient. Male pectoral fins do not appear to fulfil a signalling function in this species. We found that, for both pectoral fin size and head width, males exhibited positive allometric slopes and greater relative trait size (allometric elevation) than females. However, for head width, females also showed positive allometry, albeit to a lesser degree than males. Because fin locomotory function typically does not result in positive allometry, our findings indicate that other naturally selected uses, such as paternal care, can exaggerate trait scaling relationships.