Research Outputs
Presence of the tunicate Asterocarpa humilis on ship hulls and aquaculture facilities in the coast of the Biobio Region, south central Chile
Non-native ascidians are important members of the fouling community associated with artificial substrata and man-made structures. Being efficient fouling species, they are easily spread by human-mediated transports (e.g., with aquaculture trade and maritime transports). This is exemplified by the ascidian Asterocarpa humilis which displays a wide distribution in the Southern Hemisphere and has been recently reported in the Northern Hemisphere (NW Europe). In continental Chile, its first report dates back from 2000 for the locality of Antofagasta (23_x000E_S). Although there was no evidence about the vectors of introduction and spread, nor the source, some authors suggested maritime transport by ship hulls and aquaculture devices as putative introduction pathways and vectors. In the present study, we report for the first time the presence of A. humilis on the hull of an international ship in a commercial port in Concepción bay (36_x000E_S), south central Chile. We also found one individual associated to a seashell farm, 70 km far from Concepción bay. Further individuals were subsequently identified within Concepción bay: one juvenile settled upon international harbor pilings and a dozen individuals along aquaculture seashell longlines. For the first specimens sampled, species identification was ascertained using both morphological criteria and molecular barcoding, using the mitochondrial gene cytochrome c oxidase subunit I (COI) and a nuclear gene (ribosomal RNA 18S). The nuclear 18S gene and the mitochondrial gene COI clearly assigned the specimens to A. humilis, confirming our morphological identification. Two haplotypes were obtained with COI corresponding to haplotypes previously obtained with European and Northern Chilean specimens. The present study thus reports for the first time the presence of A. humilis in the Araucanian ecoregion, documenting the apparent expansion of this non-native tunicate in Chile over 2,000 km, spanning over three ecoregions. In addition we reveal the potential implication of the international maritime transport as a vector of spread of this species along the Eastern Pacific coast, and the putative role of aquaculture facilities in promoting local establishments of non-native tunicates.
Non-indigenous species contribute equally to biofouling communities in international vs local ports in the BiobÃo region, Chile
Growing coastal urbanization together with the intensification of maritime traffic are major processes explaining the increasing rate of biological introductions in marine environments. To investigate the link between international maritime traffic and the establishment of non-indigenous species (NIS) in coastal areas, biofouling communities in three international and three nearby local ports along 100 km of coastline in south-central Chile were compared using settlement panels and rapid assessment surveys. A larger number of NIS was observed in international ports, as expected in these ‘invasion hubs’. However, despite a few environmental differences between international and local ports, the two port categories did not display significant differences regarding NIS establishment and contribution to community structure over the studied period (1.5 years). In international ports, the free space could be a limiting factor for NIS establishment. The results also suggest that local ports should be considered in NIS surveillance programs in Chile.
Spread of the non-native anemone Anemonia alicemartinae Haussermann & Forsterra, 2001 along the Humboldt-current large marine ecosystem: An ecological niche model approach
The geographical expansion of invasive species depends mainly on its dispersal potential, and the abiotic and biotic factors affecting it. Knowing the invasive dynamic of non-native species, as well as its behavior at different natural or anthropogenic scenarios, is fundamental for planning conservation management policies and control plans. The invasive sea anemone Anemonia alicemartinae in habits from the north (18 S) to the south-central (36 S) coast of Chile and its distribution range has expanded by approximately 1,928 km in the last 50 years. Previous works have proposed that human-mediated southward transport associated with regional-scale maritime activities could explain its rapid spread. To evaluate this hypothesis, we used ecological niche models (ENM) to evaluate the potential colonization of the southernmost area of South America. Additionally, we conducted a post hoc analysis to evaluate the relationship between the prediction of the ENM and human activity measured as the number of landings of ships in ports. The models were built based on presence records of A. alicemartinae, and oceanographic variables. Results showed that sea surface salinity and annual sea surface temperature (variance) are the best predictor variables to explain the distribution of A. alicemartinae. There was a positive and significant relationship between the geographical distribution of the sea anemone predicted by the ENM and the number of landings, as a proxy of anthropogenic activity. The most susceptible areas to invasion were those that showed the highest variability in both oceanographic predictors. These areas included the BiobÃo region, Chiloé´s inland sea, Aysén, and Chacabuco regions, which together comprise two biogeographical provinces. These results sustain the proposed hypothesis and, overall, the results suggest that along with the characteristics of the life history of A. alicemartinae, oceanographic conditions and How to cite this article Pinochet J, Rivera R, Neill PE, Brante A, Hernández CE. 2019. Spread of the non-native anemone Anemonia alicemartinae Häussermann & Försterra, 2001 along the Humboldt-current large marine ecosystem: an ecological niche model approach. maritime transport as vector contribute to the southern range expansion of this invasive cryptogenic species in the Humboldt-current large marine ecosystem.
Aggressive interactions between the invasive anemone Anemonia alicemartinae and the native anemone Phymactis papillosa
The distribution range expansion of species mediated by natural or anthropic mechanisms is one of the main causes of changes in biodiversity patterns. Anemonia alicemartinae is a cryptogenic species found along the coasts of the Southeast Pacific Ocean. This species has expanded its range by >1900 km along the Chilean coast throughout the last 50 yr. A. alicemartinae cohabits with the native anemone Phymactis papillosa in the low intertidal zone, and given the limited mobility of both species, limited space could encourage aggressive behavior between them. P. papillosa shows different color phenotypes, and, as in other anemone species, color is associated with its level of aggressiveness. Here, we evaluated the aggressive behavior of A. alicemartinae on 2 color morphotypes of P. papillosa. Also, intraspecific agonistic interactions were considered in individuals of A. alicemartinae from 2 localities. Four experiments were conducted: (1) individual vs. individual, (2) individual vs. group forming a frontal line, (3) individual vs. group surrounding the individual and (4) group vs. group. Results showed A. alicemartinae to be a weak competitor against P. papillosa. The cryptogenic species lost more contests with green P. papillosa than with the red phenotype. Few aggressive interactions between conspecifics of A. alicemartinae were found. Results suggest that the expansion success of A. alicemartinae could be explained by alternative strategies, such as escape behavior, asexual reproduction and high dispersal potential.
Contrasting metatrochal behavior of mollusc and annelid larvae and the regulation of feeding while swimming
Molluscan veliger larvae and some annelid larvae capture particulate food between a preoral prototrochal band of long cilia that create a current for both swimming and feeding and a postoral metatrochal band of shorter cilia that beat toward the prototroch. Larvae encountering satiating or noxious particles must somehow swim without capturing particles or else reject large numbers of captured particles. Because high rates of particle capture are inferred to depend on the beat of both ciliary bands, arrest of the metatroch could be one way to swim while reducing captures. Larvae in eight families of annelids arrest metatrochal cilia frequently during prototrochal beat, often over a large part of the metatrochal band and with the arrested cilia aligned near the beginning of the effective stroke. In contrast, metatrochs of veligers of gastropods and bivalves rarely arrested while the prototroch beat, and those arrests were more localized and variable in position. This difference in metatrochal arrest was unexpected under hypotheses of either a single origin of this feeding mechanism or multiple origins within each phylum. Although different in metatrochal arrests, larvae of both phyla can separate swimming from feeding while both prototroch and metatroch beat. One hypothesis explaining low rates of capture per encounter, without metatrochal arrest, is a change in adhesion of prototrochal cilia with algae. In a few observations, part of the velar edge was retained within the veliger's shell so that exposed prototrochal cilia contributed to swimming while the adjacent metatroch and food groove were sequestered.
A new case of poecilogony from South America and the implications of nurse eggs, capsule structure, and maternal brooding behavior on the development of different larval types
Poecilogony is the production of different larval types within the same species. Although rare, poecilogonous species are ideal systems for testing the evolutionary and ecological implication of different developmental modes in marine invertebrates. Here, we described a new case of poecilogony, the Southern Hemisphere spionid Boccardia wellingtonensis. We used a combination of common-garden experiments, video recordings, and in vitro manipulations of individuals from three sites to (1) document the type of poecilogony, the brooding behavior of the mother, and the hatching process; (2) experimentally measure the effect of nurse eggs on the growth and type of larvae produced; and (3) document variation in the length of the brooding period, number of capsules, larvae, and nurse eggs of mothers from three sites to explore the potential for plasticity in reproductive traits. These results were compared to the previously reported poecilogonous species B. proboscidea, which resembles B. wellingtonensis in size, morphology, ecology, and reproductive strategy but differs in capsule structure. We found that in contrast to B. proboscidea, B. wellingtonensis produced larvae that, in isolation and in the presence of nurse eggs, developed into a wide range of offspring sizes. Mothers brood and hatch the larvae with frequent partial hatching of the brood during the brooding period. Although larvae could not liberate themselves, larvae crossed to other capsules as interconnections between capsules broke during the developmental period, potentially affecting food availability, sibling competition for nurse eggs, and cannibalism. Variation in brooding time and number of capsules deposited among sites suggest local adaptations.
Post-settlement movement as response to interspecific competition between the bioengineer mussels Semimytilus algosus and Perumytilus purpuratus
Post-settlement movement has been reported mainly in marine species with low or reduced adult mobility, where distribution varies in accordance with ontogeny, thus avoiding or reducing environmental stress or biological interactions. Mussels show high dispersal at the larval stage, and settlement is a highly complex process in which larvae must choose an appropriate site to attach. Although adults are mainly sedentary, it has been shown that they move on a local scale during the benthic phase in response to physical and biological factors. Semimytilus algosus and Perumytilus purpuratus are two bioengineer mussel species cohabiting most of the Chilean rocky shores. While S. algosus occurs in the low intertidal zone, P. purpuratus dominates the mid and mid-high zones. Field and laboratory experiments have shown that S. algosus is a weak competitor with respect to P. purpuratus, and post-settlers present high mobility to relocate in the intertidal. Under this scenario, we evaluated the dispersal behavior of juveniles and adults of S. algosus as a potential response to competition with P. purpuratus. We also measured the attachment strength of S. algosus in the presence of its competitor, as a measure of its escape response ability. Our results showed that the presence of P. purpuratus increased the movement activity of juveniles and adults of S. algosus and decreased their attachment strength. Field experiments carried out with marked individuals in a Chilean rocky shore, showed that S. algosus exhibits higher local dispersion in the zone where P. purpuratus is present. Mussels' high dispersal ability throughout the whole benthic phase may not only serve to reach the optimal physiological position in the intertidal, but also to reduce competition interaction.
Tidal height and sand as potential drivers of the ecological interaction of the two intertidal mussels Perumytilus purpuratus and Semimytilus algosus
The mussels Perumytilus purpuratus and Semimytilus algosus are two dominant species of intertidal rocky shores of central Chile. These species have marked differences in their distribution patterns with P. purpuratus dominating the mid-intertidal zone and S. algosus dominating the lower intertidal zone usually in habitats influenced by sand. Although it has been suggested that differences in tolerance to environmental conditions, such as air exposure and presence of sand, can explain the distribution of these species, there are currently no experimental studies to support such hypotheses. Here, we evaluated the growth and survival rate in the field of both mussel species at four different tidal heights: 25, 75, 135, and 175 cm above the zero tide. In addition, filtration rates were estimated for both species in the presence and absence of sand in laboratory conditions. The results showed that shell and wet weight growth rates of P. purpuratus were highest in mid- and medium-high tidal heights, whereas the growth rate of S. algosus was highest in the medium-low level. Similar pattern was observed for survival percentage. Furthermore, small S. algosus individuals cultivated with sand in suspension had significantly higher filtration rates than P. purpuratus. In large individuals, no differences were observed between the two species nor between treatments (presence and absence of sand). These results indicate that the differences in the distribution patterns of P. purpuratus and S. algosus in the intertidal can be explained by differences in physiological tolerances to both air exposure and to the presence of sand.
Intracapsular development in the freshwater gastropod Chilina dombeiana (Bruguiere, 1789). (Gastropoda: Hygrophila: Chilinidae)
Chilina dombeiana (Bruguière, 1789) is a native Chilean species inhabiting freshwater and estuarine environments. In the present study, a series of stages for embryonic development of the embryo is described. The snails lay gelatinous and transparent zig-zag-like string egg masses, ranging between 10 and 130 mm in length. Each egg mass contains a variable number of embryos ranging from 60 to 298 eggs with a mean density of 2.9 mm (+ 0.7 SD). Embryos inside egg masses are individually encapsulated and embedded in a jelly matrix. Hatching as crawling juveniles took place after 28 days, indicating that direct development occurs in this species. In contrast to other fresh water pulmonates, the well-developed operculum observed in C. dombeiana from the veliger stage, suggest a marine ancestry for this species.
The relationship between sex change and reproductive success in a protandric marine gastropod
Protandric species switch sex during their lifetime. According to theory, the time (body size) at which sex change occurs is determined by the reproductive success of individuals affected by social interactions as well as by post-copulatory factors. Experimental evidence is biased to few social systems making the exploration of general patterns difficult. We used the protandric marine gastropod Crepidula coquimbensis that partakes in intrabrood sibling cannibalism to test the following hypotheses: 1. Male-male competition for access to females and sibling cannibalism determine male reproductive success; 2. Males with greater access to females and with higher reproductive success will have reduced growth rates and will delay sex change. Artificial aggregations with different social structures were constructed and male reproductive success was estimated by paternity analysis. The results supported our expectations showing that male competitive ability for access to the female, time spent by males in the copulatory position, and sibling cannibalism affect reproductive success and influence time to sex change, with less successful males hastening sex change. Also, males that spent more time in the copulatory position had reduced growth rates. Comparing these results with those reported for other sequential hermaphrodites provides evidence supporting general patterns of sex change in nature.