Testing the hypothesis of greater eurythermality in invasive than in native ladybird species: from physiological performance to life‐history strategies

1. Global warming and biological invasions are important threats to biodiversity. Nonetheless, there is little information on how these factors influence performance or life‐history traits of invasive and native species. 2. The effects of temperature on physiological and fitness traits of two invasive alien species (Harmonia axyridis and Hippodamia variegata) and one native species (Eriopis chilensis) of coccinellid were evaluated, testing a model of eurythermality. Eggs of all species were exposed to four temperature treatments (20, 24, 30 and 33 °C). In adult F₂ we measured fecundity, locomotor performance, development time (total and per life stage), survival, and preferred body temperature in a thermal gradient. 3. It was found that H. axyridis had comparatively better performance at low temperatures (i.e. 20 °C), while the performance of H. variegata and E. chilensis did not change with temperature or was better at higher temperatures (30 °C). The standardised Levins index showed that all species are eurythermic. E. chilensis had a high niche overlap with the invasive alien ladybird species, rejecting the hypothesis of greater eurythermality of invasive species than native species. 4. Although there were differences in the temperature preferences and in the response of some physiological and life‐history traits of ladybirds to temperature, both the native and invasive alien species are eurythermic, contrary to the prediction. The better performance of H. axyridis at lower temperatures may result in displacement of its current distribution, and thus not all invasive species will respond favourably to global warming.     

Reproductive cycle of the sea cucumber Psolus patagonicus Ekman 1925, off Mar del Plata,Buenos Aires,Argentina

The reproductive cycle of Psolus patagonicus Ekman 1925 was studied. The species exhibits an annual reproductive cycle with one reproductive period in austral summer (February–March). During spring and summer, females showed the highest gonad-index (GI) values, while the GI values of males did not vary significantly throughout the year. Males had higher GI values than females. Maximum oocyte diameters ranged from less than 300?µm in spring to 900?µm in summer. In February, there is an increase in phytoplankton bloom for the bottom community in which P. patagonicus inhabits. The annual range in water temperature is small (6–7°C). Spawning coincided with increase in available food rather than the increase in water temperature.     

Legacy introductions and climatic variation explain spatiotemporal patterns of invasive hybridization in a native trout

Hybridization between invasive and native species, a significant threat to worldwide biodiversity, is predicted to increase due to climate‐induced expansions of invasive species. Long‐term research and monitoring are crucial for understanding the ecological and evolutionary processes that modulate the effects of invasive species. Using a large, multidecade genetics dataset (N = 582 sites, 12,878 individuals) with high‐resolution climate predictions and extensive stocking records, we evaluate the spatiotemporal dynamics of hybridization between native cutthroat trout and invasive rainbow trout, the world's most widely introduced invasive fish, across the Northern Rocky Mountains of the United States. Historical effects of stocking and contemporary patterns of climatic variation were strongly related to the spread of hybridization across space and time. The probability of occurrence, extent of, and temporal changes in hybridization increased at sites in close proximity to historical stocking locations with greater rainbow trout propagule pressure, warmer water temperatures, and lower spring precipitation. Although locations with warmer water temperatures were more prone to hybridization, cold sites were not protected from invasion; 58% of hybridized sites had cold mean summer water temperatures (<11°C). Despite cessation of stocking over 40 years ago, hybridization increased over time at half (50%) of the locations with long‐term data, the vast majority of which (74%) were initially nonhybridized, emphasizing the chronic, negative impacts of human‐mediated hybridization. These results show that effects of climate change on biodiversity must be analyzed in the context of historical human impacts that set ecological and evolutionary trajectories.     

Temperature‐dependent development of Aproceros leucopoda (Hymenoptera: Argidae), an invasive pest of elms in Europe

The temperature‐dependent development of Aproceros leucopoda Takeuchi, 1939 (Hymenoptera: Argidae), an invasive pest of elms in Europe, was studied in the laboratory on Siberian elm (Ulmus pumila) at six constant temperatures (10.9, 15.0, 19.5, 23.0, 24.3 and 27.0°C) and at a photoperiod of 16L:8D. The larvae of the species developed through 4–7 instars, and it was the 6th instar individuals of which continued their development at the highest rate. The developmental threshold (T_min) and the thermal constant (K) were determined by the linear model and Lactin‐2 model for egg, larval, prepupal and pupal stages and one whole generation. The estimated values of T_min and K by the two models for one generation were 7.3 and 7.1°C, and 426.5 and 432.7 degree‐days, respectively. Based on the survival and fecundity rates, the optimal temperature range for the species may be expected to be between 15.0 and 19.5°C. Allowing for the period of diapause, it was estimated that A. leucopoda might potentially develop through up to four or five generations per year in Hungary. These results may contribute to the better understanding of the biology of an invasive alien species in Europe.     

Climate‐induced warming imposes a threat to north European spring ecosystems

Interest in climate change effects on groundwater has increased dramatically during the last decade. The mechanisms of climate‐related groundwater depletion have been thoroughly reviewed, but the influence of global warming on groundwater‐dependent ecosystems (GDEs) remains poorly known. Here we report long‐term water temperature trends in 66 northern European cold‐water springs. A vast majority of the springs (82%) exhibited a significant increase in water temperature during 1968–2012. Mean spring water temperatures were closely related to regional air temperature and global radiative forcing of the corresponding year. Based on three alternative climate scenarios representing low (RCP2.6), intermediate (RCP6) and high‐emission scenarios (RCP8.5), we estimate that increase in mean spring water temperature in the region is likely to range from 0.67 °C (RCP2.6) to 5.94 °C (RCP8.5) by 2086. According to the worst‐case scenario, water temperature of these originally cold‐water ecosystems (regional mean in the late 1970s: 4.7 °C) may exceed 12 °C by the end of this century. We used bryophyte and macroinvertebrate species data from Finnish springs and spring‐fed streams to assess ecological impacts of the predicted warming. An increase in spring water temperature by several degrees will likely have substantial biodiversity impacts, causing regional extinction of native, cold‐stenothermal spring specialists, whereas species diversity of headwater generalists is likely to increase. Even a slight (by 1 °C) increase in water temperature may eliminate endemic spring species, thus altering bryophyte and macroinvertebrate assemblages of spring‐fed streams. Climate change‐induced warming of northern regions may thus alter species composition of the spring biota and cause regional homogenization of biodiversity in headwater ecosystems.     

Sexual reproduction and diapause of Hexarthra sp. (Rotifera) in short-lived ponds in the Chihuahuan Desert

1. In the life cycle of monogonont rotifers it is generally assumed that diapausing eggs invariably hatch into amictic stem females which produce female offspring parthenogenetically. Diapausing eggs are only produced by later generations after sexual reproduction has been induced by environmental cues. 2. We show that populations of an undescribed Hexarthra species inhabiting small temporary ponds in the Chihuahuan Desert deviate from this life cycle pattern. These ponds may dry within days and up to 85% of females were mictic. Females producing male offspring and diapausing eggs were observed 1 or 2 days, respectively, after ponds had filled with water. 3. Under laboratory conditions, 7–46% of females hatching from re‐hydrated sediments were sexual. Male offspring of these females can fertilise other mictic stem females leading to diapausing egg formation. In laboratory experiments, females produced fully developed diapausing eggs within 1.9 days at 20 °C and 1.2 days at 30 °C. 4. In addition, embryonic development time (1.1–0.3 days at temperatures between 12 and 30 °C) and juvenile period (2.1–0.5 days for the same temperature range) are shorter than those of other rotifer species. In short‐lived habitats, the potential for rapid population development and production of new diapausing eggs may be crucial in the long‐term survival of populations.     

Phenotypic variation in invasive and biocontrol populations of the harlequin ladybird, Harmonia axyridis

Despite numerous releases for biological control purposes during more than 20 years in Europe, Harmonia axyridis failed to become established until the beginning of the 21st century. Its status as invasive alien species is now widely recognised. Theory suggests that invasive populations should evolve toward greater phenotypic plasticity because they encounter differing environments during the invasion process. On the contrary, populations used for biological control have been maintained under artificial rearing conditions for many generations; they are hence expected to become specialised on a narrow range of environments and show lower phenotypic plasticity. Here we compared phenotypic traits and the extent of adaptive phenotypic plasticity in two invasive populations and two populations commercialized for biological control by (i) measuring six phenotypic traits related to fitness (eggs hatching rate, larval survival rate, development time, sex ratio, fecundity over 6 weeks and survival time of starving adults) at three temperatures (18, 24 and 30°C), (ii) recording the survival rate and quiescence aggregation behaviour when exposed to low temperatures (5, 10 and 15°C), and (iii) studying the cannibalistic behaviour of populations in the absence of food. Invasive and biocontrol populations displayed significantly different responses to temperature variation for a composite fitness index computed from the traits measured at 18, 24 and 30°C, but not for any of those traits considered independently. The plasticity measured on the same fitness index was higher in the two invasive populations, but this difference was not statistically significant. On the other hand, invasive populations displayed significantly higher survival and higher phenotypic plasticity when entering into quiescence at low temperatures. In addition, one invasive population displayed a singular cannibalistic behaviour. Our results hence only partly support the expectation of increased adaptive phenotypic plasticity of European invasive populations of H. axyridis, and stress the importance of the choice of the environmental parameters to be manipulated for assessing phenotypic plasticity variation among populations.     

Nutritional ecology of the invasive freshwater mysid Limnomysis benedeni: field data and laboratory experiments on food choice and juvenile growth

The introduction of alien species leads to changes in species composition and therefore the trophic structure. The highly invasive freshwater mysid Limnomysis benedeni is established and abundant in Lake Constance since 2006. Our aim was to confirm the assumption that L. benedeni feds on small particles and to identify their food web position. We examined the feeding mode and food preference of adult L. benedeni collected from the field and from laboratory feeding experiments by analysing their stomach contents. Stable isotope analyses (SIA) confirm an omnivorous feeding of L. benedeni. In laboratory growth experiments, we determined the growth of juveniles by feeding juveniles with different natural food sources. Growth was supported by natural biofilm, dreissenid biodeposited material and an algal suspension. Stomach analyses showed clearly that L. benedeni fed on small particles of both benthic and pelagic origin and fed by both filtering and grazing. Absence of high numbers of animal structures in the stomachs and SIA supported the lack of predatory impact in the field, but the mysid probably affects the nutrient cycles in the littoral. L. benedeni seems to play an important role in the trophic relations in Lake Constance and all other water bodies they invaded.     

Communal nesting under climate change: fitness consequences of higher incubation temperatures for a nocturnal lizard

Communal nesting lizards may be vulnerable to climate warming, particularly if air temperatures regulate nest temperatures. In southeastern Australia, velvet geckos Oedura lesueurii lay eggs communally inside rock crevices. We investigated whether increases in air temperatures could elevate nest temperatures, and if so, how this could influence hatching phenotypes, survival, and population dynamics. In natural nests, maximum daily air temperature influenced mean and maximum daily nest temperatures, implying that nest temperatures will increase under climate warming. To determine whether hotter nests influence hatchling phenotypes, we incubated eggs under two fluctuating temperature regimes to mimic current ‘cold’ nests (mean = 23.2 °C, range 10–33 °C) and future ‘hot’ nests (27.0 °C, 14–37 °C). ‘Hot’ incubation temperatures produced smaller hatchlings than did cold temperature incubation. We released individually marked hatchlings into the wild in 2014 and 2015, and monitored their survival over 10 months. In 2014 and 2015, hot‐incubated hatchlings had higher annual mortality (99%, 97%) than cold‐incubated (11%, 58%) or wild‐born hatchlings (78%, 22%). To determine future trajectories of velvet gecko populations under climate warming, we ran population viability analyses in Vortex and varied annual rates of hatchling mortality within the range 78– 96%. Hatchling mortality strongly influenced the probability of extinction and the mean time to extinction. When hatchling mortality was >86%, populations had a higher probability of extinction (PE: range 0.52– 1.0) with mean times to extinction of 18–44 years. Whether future changes in hatchling survival translate into reduced population viability will depend on the ability of females to modify their nest‐site choices. Over the period 1992–2015, females used the same communal nests annually, suggesting that there may be little plasticity in maternal nest‐site selection. The impacts of climate change may therefore be especially severe on communal nesting species, particularly if such species occupy thermally challenging environments.     

Thermal limits for the establishment and growth of populations of the invasive apple snail <Emphasis Type="Italic">Pomacea canaliculata</Emphasis>

Pomacea canaliculata is a South American freshwater snail considered as one of the world’s worst invasive alien species. A temperature of around 25 °C has usually been considered to be optimal for rearing P. canaliculata. Nevertheless, snails have not been reared under a wide range of temperatures to reveal the optimum for performance in terms of population increase. We investigated the effect of temperature on growth, survival and reproduction, estimating demographic parameters for P. canaliculata in the wide range of temperatures at which these snails are active (15–35 °C). No reproductive activity was evidenced for the snails reared at 15 °C, probably explained by the small sizes attained at this temperature. Temperatures above 25 °C did not promote a significant acceleration in growth so higher temperatures will not result in a reduction in time to reach maturity. In fact, snails from 25 and 30 °C began reproduction at the same age. We report here for the first time a detrimental effect of high temperatures that provoked a significant decrease in the contribution of snails to the next generation: the viability of eggs from the snails reared at 30 °C was very low and the snails exposed to a constant water temperature of 35 °C were unable to produce eggs. Our findings reveal a new environmental constraint that could be a determinant of the range limits of this species in invaded regions, especially during the coming decades, anticipating the scenario predicted from global warming.     

Fecundity and fertility of ladybird beetle Harmonia axyridis after prolonged cold storage

Some species of ladybird beetles (Coleoptera: Coccinellidae) mate both before and after overwintering. The purpose of the pre-diapause mating was studied in the alien invasive ladybird Harmonia axyridis (Pallas 1773). Our study demonstrates the persistence of high fecundity (daily oviposition rate of 21 eggs per fertilized female during the first month of reproduction) and fertility (85 % of eggs hatching) of females of H. axyridis after long storage (up to eight months) at low temperature (6 °C). The females were not mated after activation in spring and had to rely on the sperm supply maintained from the pre-winter period (58 % of females were fertilized). Unfertilized females also laid eggs but in low numbers (an average of 345 eggs by virgin females during an individual’s lifetime, 1,174 eggs by females fertilized before winter) and after a longer pre-oviposition period (2–5 weeks in comparison to 7–8 days for fertilized females). We show that the unfertilised eggs were not trophic eggs. The high sperm survival ability observed questions the need for the high levels of sexual activity generally observed in Coccinellidae. Fertilized females of H. axyridis may found large colonies after dispersal to new areas even without males, which contributes to the striking invasive ability of this species.     

The reproductive ecology of the invasive Asian paddle crab,Charybdis japonica (Brachyura: Portunidae), in northeastern New Zealand

The Asian paddle crab, Charybdis japonica, native to the northwest Pacific, is an invasive species that has established populations within northeastern New Zealand. Here, we provide a detailed examination of C. japonica's reproductive seasonality and gametogenesis, sizes at physiological maturity, and sex ratios outside its native range. Trapping in the Weiti River Estuary, Whangaparoa (36°38.4′S, 174°43.6′E) from February 2010 to May 2012 indicated a male‐biased population. Reproductive seasonality largely depended on female condition, since males contained ripe spermatozoa year‐round. Female gametogenesis began during late autumn, with oocyte development occurring throughout winter to early spring. Based on the presence of mature gametes during late winter and the appearance of gravid females in early November, mating began during early spring, with spawning in mid‐spring. Spawning coincided with a large drop in the gonad index, and occurred between sea surface temperatures of 17–22°C and 11.75–13.5 h day length. Females remained reproductive for up to 5 months of the year. Histological observation suggested that CW50 (the carapace width at which 50% of the population is mature) for females was 46.39 mm, while CW50 for males could not be determined due to a paucity of immature males. Histological analysis also suggested that females of C. japonica could produce store sperm and multiple broods annually.     

Stochastic models for the Trojan Y-Chromosome eradication strategy of an invasive species

The Trojan Y-Chromosome (TYC) strategy, an autocidal genetic biocontrol method, has been proposed to eliminate invasive alien species. In this work, we develop a Markov jump process model for this strategy, and we verify that there is a positive probability for wild-type females going extinct within a finite time. Moreover, when sex-reversed Trojan females are introduced at a constant population size, we formulate a stochastic differential equation (SDE) model as an approximation to the proposed Markov jump process model. Using the SDE model, we investigate the probability distribution and expectation of the extinction time of wild-type females by solving Kolmogorov equations associated with these statistics. The results indicate how the probability distribution and expectation of the extinction time are shaped by the initial conditions and the model parameters.     

Moving south: effects of water temperatures on the larval development of invasive cane toads (Rhinella marina) in cool‐temperate Australia

The distributional limits of many ectothermic species are set by thermal tolerances of early‐developmental stages in the life history; embryos and larvae often are less able to buffer environmental variation than are conspecific adults. In pond‐breeding amphibians, for example, cold water may constrain viability of eggs and larvae, even if adults can find suitable thermal conditions in terrestrial niches. Invasive species provide robust model systems for exploring these questions, because we can quantify thermal challenges at the expanding range edge (from field surveys) and larval responses to thermal conditions (in the laboratory). Our studies on invasive cane toads (Rhinella marina) at the southern (cool‐climate) edge of their expanding range in Australia show that available ponds often average around 20°C during the breeding period, 10°C lower than in many areas of the toads’ native range, or in the Australian tropics. Our laboratory experiments showed that cane toad eggs and larvae cannot develop successfully at 16°C, but hatching success and larval survival rates were higher at 20°C than in warmer conditions. Lower temperatures slowed growth rates, increasing the duration of tadpole life, but also increased metamorph body mass. Water temperature also influenced metamorph body shape (high temperatures reduced relative limb length, head width, and body mass) and locomotor performance (increased speed from intermediate temperatures, longer hops from high temperatures). In combination with previous studies, our data suggest that lower water temperatures may enhance rather than reduce recruitment of cane toads, at least in areas where pond temperatures reach or exceed 20°C. That condition is fulfilled over a wide area of southern Australia, suggesting that the continuing expansion of this invasive species is unlikely to be curtailed by the impacts of relatively low water temperatures on the viability of early life‐history stages.     

The relationship between invasive alien species and main climatic zones

Analysis of numbers of invasive species (insects, plants, plant diseases) in each of Chinese provinces and the world’s climatic zones respectively, demonstrated positive correlations between diversity of bioinvasion and air temperature and precipitation (the two main criteria for classifying world climate types). When the air temperature is within the range of 1–25°C, the degree of diversity of invasive alien species increases exponentially with air temperature. However, when the air temperature falls outside the range of 1–25°C, the diversity of harmful invasive alien species changes with air temperature in a parabolic pattern, showing a “mid-latitude bulge”. Namely, when air temperatures are too high or too low, a lower average invasion frequency of harmful alien species is observed. Invasion by harmful invasive alien species is more weakly related to precipitation. Different climatic zones showed dramatic differences in resistance to the invasion of invasive alien species due to their different characteristic climatic factors, mainly including air temperature and precipitation.     

Prunus serotina in Italy: a challenging candidate for the national list of priority invasive alien species

Abstract

In accordance with the European regulation on Invasive Alien Species (IAS), the black cherry tree (Prunus serotina Ehrh.) has recently been indicated as one of the 96 species proposed for the development of the national list of priority invasive alien species in Italy. The species, native to North America, is recognised as one of the most harmful IAS in Europe, given its high spread potential and the associated ecological and economic impacts reported in its alien range. Although P. serotina is recognised as a pest within the EU, plants are still available on the market, confirming intentional introduction as a current potential pathway for this species arrival into new areas. Since a comprehensive overview of the main features characterizing the invasive potential of this species is still lacking, we aim to underpin the high priority status of P. serotina as a IAS of main concern in Italy by outlining the main biological features, pathways and impacts of the black cherry in its secondary range. Management measures to be potentially included in specific action plans are also summarised.     

The Life Histories of Two Temporarily Coexisting,Pond Dwelling Cladocerans

Investigations on the life histories of two cladocerans, Moina brachiata and Daphnia obtusa, in a small, nearly temporary pond in South Germany revealed that M. brachiata is better adapted to fluctuating environmental conditions; the species dominated from May to October. D. obtusa was present in spring and autumn/winter but disappeared completely during the summer months. Both species coexisted for extended periods in spring and autumn; abundance of D. obtusa was generally by an order of magnitute lower. Four periods of low water level were slightly preceded by or coincided with a decrease of clutch size, a decrease of the proportion of egg bearing females indicating that both species suffered from food shortage. Laboratory investigations on life history parameters showed that the two species have different temperature tolerances and preferences. M. brachiata showed its highest reproductive success at 25 and 30°C but died at temperatures <15°C and ≥ 35°C. D. obtusa experienced a broader temperature range (2 to 25°C) but could not withstand temperatures ≥ 30°C. Short term starvation periods (3d) caused the death of M. brachiata females, while D. obtusa soon recovered and reproduced when being refed. M. brachiata is a typical r-species with early reproduction, rapid development, high population growth rates and a high tendency to produce resting eggs; D. obtusa pursues more the concept of k-selection.     

Temperature-dependent development of the predatory mite <Emphasis Type="Italic">Cheyletus malaccensis</Emphasis> (Acari: Cheyletidae)

The effect of temperature on the development of immature stages of the predator Cheyletus malaccensis Oudemans, produced by either fertilized or virgin females, was studied at 17.5, 20, 25, 30, 32.5, and 35°C, 80 ± 5% relative humidity, in complete darkness, and fed Tyrophagus putrescentiae (Schrank). The population maintained at 15°C failed to reproduce. The thermal data obtained were used for the estimation of the thermal requirements (developmental thresholds, thermal constant, optimum temperature) of this predator by a linear and nonlinear model (Logan type I model). Upper and lower developmental thresholds ranged between 37.4–37.8 and 11.6–12.0°C, respectively. Optimum temperature for development was estimated at between 33.1 and 33.5°C. The thermal constant ranged between 238.1 and 312.5 degree-days. Based on the data of the total pre-imaginal period, immatures’ survival peaked at 25°C. Arrhenotokous parthenogenesis (haplodiploidy) is confirmed in the species: virgin females always produce males, whereas fertilized females give rise to offspring of both sexes. Survival of the immature stages and offspring sex ratio were not significantly influenced by temperature.     

Biological studies and phenology of the slug Deroceras reticulatum (Müller, 1774) (Pulmonata: Stylommatophora)

Summary

In the southeast of Buenos Aires Province (Argentina), slugs cause reductions in crop yield by killing seeds and seedlings, by destroying stems and growing points, and by reducing leaf area. Deroceras reticulatum (Müller, 1774) was introduced into Argentina a long time ago, but the biology and ecology of this species in this country are still unknown. The aim of this study was to determine how temperature affects growth rate, reproduction and survival of D. reticulatum and also to assess the seasonal phenology of this species. The life cycle of D. reticulatum was investigated at 12°C, 20°C and at a temperature which alternated between 12°C for 16 h and 20°C for 8 h. The light/dark regime during the experiment was LD: 8/16 h. Slug populations were sampled from grassland located at the Experimental Station of INTA Balcarce. Eggs hatched after 16.3 to 39.8 days and fertility varied between 83.9% and 91.1%. The mean time from hatching to oviposition was longer and fecundity lower at 20°C than 12°C and 12/20°C (p <0.05). The net reproductive rate was 1.49 to 70.53 and the mean generation time varied between the 27 and 30 weeks. In natural conditions, eggs of D. reticulatum were found from early winter until the end of the spring, when soil temperature was 8–15°C and humidity was above 15%. These results indicate that D. reticulatum has one generation per year, univoltine phenology, with peak densities of slugs from the beginning of winter to the end of spring, which coincides with the sowing and emergence of winter and summer crops.     

Growth and survival of the invasive alga, <Emphasis Type="Italic">Caulerpa taxifolia</Emphasis>, in different salinities and temperatures: implications for coastal lake management

The alga Caulerpa taxifolia is an invasive pest species in many parts of the world and has recently become established in several estuaries in south eastern Australia. A major infestation has occurred in Lake Conjola, an intermittently open and closed coastal lagoon in southern NSW. Short term (1 week) laboratory experiments were carried out to investigate growth and survival of fragments of C. taxifolia collected from this outbreak, under a range of salinities (15–30 ppt) and water temperatures (15–30°C). Fronds, stolons and thalli of the alga all displayed similar responses. Many of the algal fragments doubled in size over the week and a maximum growth rate of 174 mm/week was recorded. Fragments showed good growth (> 20 mm/week) at salinities > 20 ppt and temperatures > 20°C. Almost total mortality occurred at salinities lower than 20 ppt and temperatures less than 20°C. Historical records of water quality demonstrate that prior to entrance manipulation in 2001, salinities in Lake Conjola had often dropped to below 17 ppt for extended periods (up to 2 years). This suggests that management of the alga may be improved if the lake was allowed to undergo its normal cycles of opening and closing to the ocean, and that entrance manipulation may be one factor that has influenced the success of this invasive species.