Effects of substrata and conspecific odour on the metamorphosis of mangrove crab megalopae, Ucides cordatus (Ocypodidae)

Environmental cues associated with the adult habitat induce metamorphosis in many marine invertebrate larvae and thus settlement in a place appropriate for completing their life cycle. Ucides cordatus is a semi-terrestrial crab that exemplifies organisms living in mangrove estuaries as adults, but whose larvae develop offshore and return to the estuarine environment prior to metamorphosis. We investigated the influence of substrata (burrow mud, channel-bank mud, dead wood, sand, nylon mesh) and conspecific adult cues (crab-conditioned seawater) on metamorphosis and the duration of the megalopal phase. Our results suggest that U. cordatus megalopae settle in areas populated by conspecific crabs and/or muddy habitats. High moulting rates were observed in the treatments with mud and/or adult odour (67–91%), whereas in the other substrata treatments and in the seawater control less than 20% and 5% of the megalopae metamorphosed, respectively. Megalopae are also capable of delaying metamorphosis. Over channel-bank mud they moulted 8 days later at an average, compared to the treatment with crab-conditioned seawater. Such a delay could result in a reduced juvenile fitness. In seawater control almost all megalopae died prior to metamorphosis, without moulting spontaneously as observed in other species. This reflects the crucial importance of habitat cues for the settlement and recruitment of this ecologically and economically important species. We outline the necessity of future research concerning post-metamorphic carry-over effects, the possible role of biofilms in absorbing and/or producing settlement cues, as well as the question whether the megalopae react specifically to the odour of U. cordatus, or also to the one of other crab other species. The latter is of particular interest against the background of the spreading lethargic crab disease that results in massive mortalities of U. cordatus, but not of co-occurring-species

Salinity tolerance of northern Brazilian mangrove crab larvae, Ucides cordatus (Ocypodidae): Necessity for larval export?

The life cycle of the semiterrestrial mangrove crab Ucides cordatus includes pelagic larvae that are released into estuarine waters during the wet season and who may thus encounter potentially stressful low and variable salinity conditions. The effect of salinity on the survival of the zoea larvae, the number of zoeal stages and the duration of development from hatching to megalopa was experimentally studied by rearing larvae from the Caeté estuary, Northern Brazil, in seven salinity treatments (0, 5, 10, 15, 20, 25, and 30). For a better interpretation of the laboratory results, estuarine salinities were measured over five consecutive years during the species' reproductive season. The survival of the zoea larvae varied significantly with salinity, while the number of stages and the duration of their development remained constant. Development to megalopa took 20.77 ± 1.57 days and comprised five zoeal stages with ZI and ZII being euryhaline and later stages stenohaline. The newly hatched larvae stayed alive for up to 6 days in freshwater (average 4.32 ± 0.82 days), but did not moult to the second zoeal stage. ZII larvae first occurred from salinity 5 onwards and later zoeal stages at all tested salinities ≥10. However, the larvae only survived to megalopa at salinities ≥15, with highest numbers at salinity 30 (72%) and lowest at 15 (16%). Lethal salinities ≤10 occurred frequently in the estuary during the reproductive season. This suggests a need for larval export to offshore and thus more saline waters to allow for significant larval survival and maintenance of viable populations of this commercially important species. A regional rather than local approach for management is suggested due to the likelihood of long distance larval dispersal by offshore currents

O efeito da salinidade no desenvolvimento larval do caranguejo - uçá, Ucides cordatus (Linnaeus, 1763) (Decapoda: Ocypodidae) no Norte do Brasil

The present work studied the effect of salinity on the survival and duration of larval development of the mangrove crab, Ucides cordatus (from the Caeté River estuary, North of Brazil) until the megalopal phase in seven salinity treatments (0, 5, 10, 15, 20, 25 e 30). Salinity significantly affected the survival of the zoea larvae, however it did not affect the duration of the larval development (20.77 ± 1.56 days). In salinity 0, 5 and 10 all zoea larvae died. Only from off salinity 15, complete development until the megalopal phase occurred. The survival rate was highest in salinity 30 (72%) and lowest in 15 (16%). The reduced survival rate of the U. cordatus zoea larvae, in low salinities, indicates the necessity of larval dispersion from the estuary to coastal waters, where salinity conditions for larval development are more favorable. Otherwise, if there was no dispersion, the reduced salinity of estuarine waters in the rainy season would cause a high mortality thereby affecting the recruitment, maintenance and growth of the U. cordatus population in the mangroves

The effect of salinity on the larval development of the uca-crab, Ucides cordatus (Linnaeus, 1763) (Decapoda: Ocypodidae) in northern Brazil.

The present work studied the effect of salinity on the survival and duration of larval development of the mangrove crab, Ucides cordatus (from the Caeté River estuary, North of Brazil) until the megalopal phase in seven salinity treatments (0, 5, 10, 15, 20, 25 e 30). Salinity significantly affected the survival of the zoea larvae, however it did not affect the duration of the larval development (20.77 ± 1.56 days). In salinity 0, 5 and 10 all zoea larvae died. Only from off salinity 15, complete development until the megalopal phase occurred. The survival rate was highest in salinity 30 (72%) and lowest in 15 (16%). The reduced survival rate of the U. cordatus zoea larvae, in low salinities, indicates the necessity of larval dispersion from the estuary to coastal waters, where salinity conditions for larval development are more favorable. Otherwise, if there was no dispersion, the reduced salinity of estuarine waters in the rainy season would cause a high mortality thereby affecting the recruitment, maintenance and growth of the U. cordatus population in the mangroves

Metamorphosis of mangrove crab megalopae, Ucides cordatus (Ocypodidae): effects of interspecific versus intraspecific settlement cues

It has recently been shown that metamorphosis of Ucides cordatus megalopae is triggered by substrata from the mangrove forest habitat, and, in particular, adult conspecific odours. Here we demonstrate that the gender of the odour-emitting crabs is insignificant for the metamorphic response in this species. We further investigate whether other estuarine crabs (Goniopsis cruentata, Uca spp., and Callinectes danae) also induce settlement and metamorphosis of U. cordatus megalopae. This is of special interest for population recovery in areas hit by lethargic crab disease (LCD), a fungus that selectively kills U. cordatus but not co-occurring species. Ucides megalopae were reared in four treatments with interspecific-conditioned seawater and tested against the effects of conspecific-conditioned seawater (positive control) and pure seawater (negative control). All megalopae in the positive control metamorphosed successfully, while only one (2%) moulted in the negative control, with a delay of 10 days compared with the latest metamorphosis in the former treatment. In seawater conditioned with U. maracoani and C. danae, which occur on sediment banks and in tidal creeks respectively, all larvae died before reaching the juvenile stage. In the treatments with odours of species that share the same mangrove forest microhabitat as U. cordatus, i.e. G. cruentata and a group of five fiddler crab species (mixed-odour treatment), 20 and 10% respectively of the megalopae moulted with a delay of up to 11 days. No specimens metamorphosed after day 39, but megalopae lived up to 93 days. Since only the conspecific- and coexisting-species treatments stimulated development, we hypothesize that Ucides megalopae are able to precisely identify species-habitat-specific settlement cues. This will be investigated in more detail in future studies, which will also test the effects of the odours of the five forest fiddler crab species separately. The impact of the interspecific odour treatments was much smaller than that of the conspecific odours, nevertheless elevated moulting rates of up to 18% relative to seawater may still significantly accelerate the repopulation of U. cordatus in areas lacking conspecifics, e.g. after massive crab mortalities or at first colonization

Chemical induction in mangrove crab megalopae. Ucides cordatus (Ucididae): Do young recruits emit metamorphosis-triggering odours as do conspecific adults?

In many brachyuran species, including the mangrove crab Ucides cordatus, water-soluble chemicals (odours) emitted by adult residents trigger metamorphosis of megalopae, probably facilitating habitat selection and settlement near conspecific crab population. New field findings revealed that early benthic crab stages co-inhabit burrows of both juveniles and adults of U. cordatus which raised the question whether megalopae are also stimulated by sexually immature juveniles. Therefore, we tested in an experimental laboratory study the hypothesis that small benthic recruits and older juveniles also emit metamorphosis-stimulating odours as do conspecific adult crabs. U. cordatus megalopae were cultivated in eight conspecific odour-treatments containing seawater previously conditioned with crabs of different carapace widths (CW 0.15–5.0 cm) and in a control treatment with filtered seawater not conditioned with crabs. In all odour-treatments, including those with small immature crabs, the percentage of metamorphosed larvae was significantly higher (≥74%) and the average development was shorter (15.8–19.3 days) than in the control group, where only 30% moulted after 25.6 ± 6.6 days of megalopal development. In addition, megalopae developed 2.7 days faster when exposed to odours from young and older juveniles compared to those larvae kept in contact with odours from conspecific adults. Our results clearly demonstrate that the emission of metamorphic odours in U. cordatus is independent of size/age or sexual maturity. The responsiveness of megalopae to chemicals emitted by resident crabs of varying ages should aid the natural recovery of U. cordatus populations in areas significantly affected by size-selective fishery where only large conspecific adults are harvested

Metamorphosis of the edible mangrove crab Ucides cordatus (Ucididae) in response to benthic microbial biofilms

Benthic microbial communities can play an important role in the induction of larval metamorphosis in marine invertebrates. The role of biofilms in recognizing the parental habitat is unknown for the ecologically and economically important mangrove crab Ucides cordatus, a species suffering from environmental pressures such as habitat degradation and disease. In the laboratory, we experimentally tested the influence of biofilms grown in offshore and estuarine waters on the moulting rates and developmental time to metamorphosis (TTM) of the last larval stage (megalopa) of U. cordatus. Here, we specifically studied whether: (i) megalopae of this larval-exporting species respond differently to marine (MB) and estuarine biofilms (EB), (ii) biofilms previously immersed (24 h) in adult crab-conditioned seawater (ACSW; i.e. conspecific chemical cues) exhibit a greater effectiveness (i.e. by inducing higher moulting rates and shorter TTM) on larval metamorphosis than non-immersed ones and (iii) biofilms pre-immersed (24 h) in ACSW decrease their effectiveness when incubated (24 h) in pure filtered seawater (FSW) before offering them to megalopae. U. cordatus megalopae metamorphosed to the first juvenile crab stage (JI) in response to both MB and EB, however the moulting rates were significantly higher and development TTM shorter in the presence of EB compared to MB. The metamorphosis-inducing effects were significantly enhanced when both types of biofilms were pre-immersed in ACSW, suggesting that conspecific cues can be absorbed and stored by microbial matrix. The higher inductiveness of EB compared to MB persisted after previous contact with ACSW. Furthermore, EB exhibited a significantly greater biomass production (measured as dry weight) than MB, and when immersed in ACSW, both groups of biofilms significantly increased in biomass (maintaining the hierarchy EB > MB), suggesting that water-soluble chemical substances emitted by the adult crabs may have been absorbed and metabolized by the biofilms. The metamorphic inductiveness and biomass production decreased when both groups of biofilms that were previously kept (24 h) in ACSW were thereafter immersed (24 h) in FSW. All megalopae successfully moulted to JI after accelerated TTM when reared in ACSW, regardless of presence or absence of biofilms, corroborating that conspecific stimuli are the most effective metamorphosis-stimulating cues tested so far in U. cordatus. The fact that EB improved the moulting rates and shortened the development TTM indicates that megalopae can recognize and respond to microbial assemblages typical for specific environments. This should be ecologically important during larval recruitment by facilitating settlement in appropriate habitats for the post-metamorphic development of early benthic recruits. In addition, the influence of EB could encourage colonization of new estuarine areas and aid natural recovery of U. cordatus in mangrove habitats where crab populations have suffered significant reduction due to deforestation, fishing pressure or diseases

Carry-over effects of delayed larval metamorphosis on early juvenile performance in the mangrove crab Ucides cordatus (Ucididae)

Competent larvae of most benthic marine invertebrates including brachyuran decapod crustaceans can delay their settlement and metamorphosis in the absence of suitable environmental cues. This however may carry significant costs for the post-larval performance and fitness of early benthic juveniles. In this paper, we examined in the laboratory whether delayed larval metamorphosis affects early juvenile performance of the mangrove crab Ucides cordatus, measured as survival and growth through the first five juvenile crab stages (JI–JV). We further investigated whether ‘carry-over’ effects of an extended larval period vary intraspecifically among juveniles from three different hatches (F1–F3). Ucides megalopae were cultivated in two treatments. In the first one, the control, megalopae were cultivated continuously in exposure to conspecific crab odours (adult-conditioned seawater = ACSW) until moulting to the first juvenile crab stage. In the second, the delay-treatment, settlement-competent megalopae were kept for 20 days in pure filtered seawater without metamorphosis-triggering odours. From day 21 onwards, megalopae were transferred to ACSW to induce larval moult. The results showed that delayed metamorphosis affects early juvenile survival and growth of U. cordatus. In the delay-treatment, survival rates of juveniles (JI–JV) from F1–F3 were 11–31% lower and intermoult development periods 1.5–4.2 days longer than in the control. Delayed juveniles (F1–F3) exhibited 9.2–14.1% smaller body sizes (carapace width) and 1.4–3.9% lower growth rates mostly in the first two moulting events compared to specimens derived from non-delayed megalopae. Most of the detrimental effects were observed in almost all five crab stages studied, indicating that the costs of delayed metamorphosis may persist throughout early juvenile development. Juvenile morphology, however, was not affected. Our study also showed that carry-over effects of delayed larval moult vary intraspecifically among offspring produced by different females. This plasticity in phenotypic effects could relate to differences in the nutritional state, health or age of the females, to stressful environments during embryogenesis or to genetic differences. The observed carry-over effects of delayed settlement on early juvenile performance may affect the overall recruitment success and population dynamics of U. cordatus. In the field, such a situation would appear when new mangrove areas void of conspecific crabs are colonized or when degraded sites are re-populated by larvae originating from elsewhere populations