The influence of pre‐settlement and early post‐settlement processes on the adult distribution and relative dominance of two invasive mussel species

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Habitat selection and juvenile persistence control the distribution of two closely related Caribbean damselfishes

Summary On many Caribbean fringing coral reefs, two closely related and ecologically similar damselfishes, the beaugregory (Stegastes leucostictus Müller and Troschel) and the cocoa damselfish (S. variabilis Castelnau), occupy nonoverlapping vertical distributions. In St. Croix (USVI), beaugregory are very abundant in shallow water back reef habitats (1–2 m depth) while cocoa damselfish are restricted to the base of the forereef (10–15 m depth).In this study, the roles of habitat selection at settlement and juvenile persistence were investigated to determine their influence on this pattern of zonation. Settlement events observed at intervals over a two-year period revealed that habitat selection occurred at settlement and was confined to habitats occupied by adults. In addition, differences in juvenile persistence (due to mortality and/or emigration) were found when species were translocated between depths. Over a period of 100-days, juvenile beaugregory moved from 1 m to 12 m depth suffered four-fold greater losses at the deeper sites than shallow water controls, while translocated cocoa damselfish suffered twice as many losses in shallow water than controls at 12 m depth. Despite these differences in persistence, growth rates of the two species were similar and independent of depth. These results indicate that preferential habitat selection at settlement, perhaps an evolutionary response to differential juvenile mortality, may play a deciding role in determining distributions of ecologically similar species of coral reef fishes.     

The influence of resident adults on recruitment: a comparison to settlement

For species recruiting into established sessile communities, the adult colonies and individuals already present form a significant part of the environment and have the potential to alter both larval settlement rates and post-settlement mortality. Settlement rates can be reduced by predation on larvae, by the removal or addition of substratum space, or by stimulation or prohibition of larvae from settling on adjacent substratum. Once attached, the recruiting individual can still be influenced by predation or overgrowth by residents, by the added physical structure for firmer attachment, or by being camouflaged from motile predators. To examine those processes by which residents affect recruitment we exposed experimental substrata with three densities of adults of a single species at a site in eastern Long Island Sound, USA for a 1-wk period. Seven different species of common invertebrates were used in nine separate experiments. The major effect of most resident species was the usurpation of space and the restricting of recruitment to adjacent unoccupied areas. This was particularly true for resident ascidians and bryozoans, but less so for barnacles and oysters. In fact several species recruited in higher densities on or next to oysters and barnacles. Comparison to 1-day settlement experiments indicated that the encrusting ascidian species Diplosoma and possibly Botryllus reduced recruitment relative to settlement, probably by overgrowing newly-settled individuals. However, in the presence of most resident species, recruitment patterns were not greatly different from settlement patterns, indicating that the effects of the attached community on recruitment may result from influences on settlement.     

The Compatibility of Control Agents used for the Control of the South American Leafminer, Liriomyza huidobrensis

The compatibility of infective juveniles (IJs) of the entomopathogenic nematode, Steinernema feltiae, the parasitic wasps Diglyphus isaea and Dacnusa sibirica, and chemical insecticides to control larval stages of the South American leafminer, Liriomyza huidobrensis, on leafy salad crops was investigated. In intact leaf mines, leafminer larvae already parasitized by D. isaea which had developed to the larval or pupal stage, and D. sibirica in the larval stage were subsequently also infected by S. feltiae following a foliar application. This reduced the potential of the wasps to survive until the adult stage. In similar tests foliar applied chemical insecticides reduced the survival to adulthood of D. sibirica and their host larvae. However, a soil drench of imidaclorpid did not cause a significant reduction in the number of D. sibirica which survived the treatment and developed to adult emergence. Tests to investigate the utilisation of parasitoids following foliar applications of nematodes found that adult D. isaea did not discriminate between the healthy and nematode-infected leafminer larvae for host feeding, although 98% of eggs laid by the female wasps were deposited alongside healthy larvae.     

Production and supply of larvae as determinants of zonation in a brooding tropical coral

Production and settlement of planktonic larvae of the coral Favia fragum (Esper) were studied. The species is restricted to shallow back- and fore-reef habitats throughout the Caribbean Sea. Adults are in their greatest abundance on the reef-crest and shallow reef slope (<3 m) at Tague Bay, St. Croix, in the US Virgin Islands. Because F. fragum broods larvae that are capable of immediate settlement, this distribution pattern may be due to variation in fecundity among depths. Corals were collected from shallow (1.0 m) and deep depths (10-13 m) and cultured in individual containers exposed to shaded ambient light. Corals from shallow depths had greater fecundity (polyp⁻¹ lunar cycle⁻¹) and were larger than deep corals. To test the hypothesis that fecundity was related to successful fertilization, corals were kept in different densities in an area with sea-grass, where there were no natural adults. Production of larvae 6 months later was not affected by density of adults, possibly due to self-fertilization. Larval choice of habitat was also examined. In the laboratory, twice as many larvae settled on coral rubble fragments collected from depths where adults were common (1.5 and 3 m) than on those from depths where adults were rare (10 m). Larval supply may establish the vertical distribution of adults on St. Croix.     

Anti-predatory chemical defenses of ascidians: secondary metabolites or inorganic acids?

Both secondary metabolites and inorganic acids have been hypothesized to protect adult ascidians from predation, raising the possibility of alternative defensive strategies in these sessile, soft-bodied, benthic invertebrates. The objective of this investigation was to determine if ascidian species from the Western Atlantic have these chemical defenses against fish predators, and if so, to determine their location within the body of the ascidian. The palatability of crude organic extracts of whole ascidians, as well as the dissected tunics, viscera, and gonads (when possible) were determined at natural volumetric concentrations using laboratory feeding assays with the bluehead wrasse, Thalassoma bifasciatum. Acidified food pellets were also assayed to determine the effect of lowered pH on predation. Sixteen of the 17 species tested had deterrent organic extracts from some region of the body (Aplidium constellatum, Aplidium stellatum, Ascidia interrupta, Ascidia nigra, Botrylloides sp., Clavellina picta, Didemnum candidum, Didemnum vanderhosti, Diplosoma listerianum, Ecteinascidia turbinata, Eudistoma capsulatum, Eudistoma hepaticum, Rhopalaea abdominalis, Styela plicata, Symplegma rubra, and Trididemnum solidum). The location of the deterrent secondary metabolites was isolated in the gonad in all three solitary species, raising the possibility that these defenses are passed on to eggs or larvae. Nine ascidian species sequestered acid in their tunics (A. interrupta, A. nigra, A. stellatum, D. candidum, D. vanderhosti, E. capsulatum, E. hepaticum, R. abdominalis, and T. solidum) at levels that were effective in deterring fish predation (pH≤3.0). Only one species (Botrylloides nigrum) had neither chemical defense. Results of this study indicate that there is not a clear trade-off between the presence of secondary metabolites and inorganic acid defenses in ascidians, suggesting that these defenses are redundant, or that alternative chemical defenses may have evolved for different predators or for different stages in the life history of the ascidians producing them.     

Comparative predation rates on larval snappers (Lutjanidae) in oceanic, reef, and nearshore waters

The life cycle of most reef fishes involves pelagic larvae entering the nearshore environment to settle to benthic substrates. Settlement is considered to be highly risky as larvae encounter high rates of predation mortality associated with shallow nearshore habitats. This potential bottleneck may be particularly significant for many tropical snapper (Lutjanidae) species which bypass the reef to settle to very nearshore seagrass areas. To test the hypothesis that predation-related mortality increases as naïve late-stage fish larvae leave the offshore oceanic environment and enter the nearshore to settle, relative nocturnal predation rates on tethered late-stage snapper larvae were measured in oceanic, coral reef, and nearshore surface waters of the lower Florida Keys, USA. Both relative predation rate and probability of predation in oceanic areas seaward of the reef was significantly greater than over reef or nearshore seagrass/hardbottom habitats. This surprising result may be due to differences in the density or spatial distribution of potential predators between deep offshore (near flotsam at the surface) and shallow nearshore environments (demersal). These findings suggest that successful late-stage snapper larvae should avoid surface waters in deep oceanic areas and move upward in the water column as they pass over the reef and other shallow nearshore environments prior to settlement.     

In situ settlement behaviour of damselfish (Pomacentridae) larvae

Settlement‐stage damselfish (Pomacentridae) larvae of 13 species in seven genera were obtained from light traps at Lizard Island, Great Barrier Reef, Australia. Behaviour, observed in situ by SCUBA divers, of 245 larvae (6–13 mm, L_S; 5–60 individuals per species) released individually within a few m of reefs during the day differed markedly among species. From 0–28% (range among 13 species) of individuals of each species swam away from the adjacent reefs without swimming to the reefs. Of those that swam to a reef, 0–75% settled. For three species, sufficient data were available to test the hypothesis that these percentages did not differ amongst reefs: the hypothesis was rejected in one species. From 0–75% of larvae that reached the reef were eaten, 0–63% subsequently left the reef and 0–60% were still swimming over the reef at the end of the observation period. Swimming speeds of all but one species were greater when swimming away from the reef than toward it. Most species exceeded average current speeds when swimming away from reefs, but not when swimming toward and over them. Average swimming depths were in the upper half of the water column for most species, and were somewhat greater where the water depths were greater. The time the larvae swam over the reef before settling and the distance swum varied greatly among species from 0 to a mean of 5.5 min and 43 m. Settlement habitats chosen differed amongst species, and in some species, they were very specific. Average settlement depth varied among species from 6–13.5 m. In one species, settlement depth varied between reefs. About half of the 53 observed interactions between larvae and reef resident fishes were predation attempts: fishes of eight species (six families) attacked larvae. The other interactions were aggressive approaches by 11 species of resident fishes, all but one of which were pomacentrids. Many of these aggressive interactions discouraged settlement attempts. Larvae of some species experienced no predatory or aggressive interactions, whereas in other species interactions averaged >0.6 per released larva. Species that swam more‐or‐less directly to settlement sites near the reef edge experienced more interactions. Even within the same family, settlement behaviour differed among species in nearly all measures.     

Effects of sedimentation on settlement of Acropora millepora

Sedimentation is assumed to affect communities of reef-building corals by inhibiting larval settlement, however there has been no quantitative evidence to support this supposition. The effects of varying rates of sedimentation (0.5 to 325 mg cm^-2d^-1) on settlement rates of Acropora millepora larvae were examined experimentally, in aquaria. Settlement and juvenile orientation onto conditioned cut coral plates were recorded after two days. Higher sedimentation rates reduced the number of larvae settling on uppersurfaces, but total numbers of settled larvae were not significantly affected by sedimentary regime. While total settlement was unaffected by sediment under the confined experimental conditions, it is likely to be reduced under field conditions, since accumulation of sediment on upward-facing surfaces will greatly reduce the overall amount of suitable substratum available.     

Depth distributions of coral reef fishes: the influence of microhabitat structure,settlement, and post-settlement processes

Many coral reef fishes have restricted depth ranges that are established at settlement or soon after, but the factors limiting these distributions are largely unknown. This study examines whether the availability of microhabitats (reef substrata) explains depth limits, and evaluates whether juvenile growth and survival are lower beyond these limits. Depth-stratified surveys of reef fishes at Kimbe Bay (Papua New Guinea) showed that the abundance of new settlers and the cover of coral substrata differed significantly among depths. A field experiment investigated whether settling coral reef fishes preferred particular depths, and whether these depth preferences were dependent on microhabitat. Small patch reefs composed of identical coral substrata were set up at five depths (3, 6, 10, 15 and 20 m), and settlement patterns were compared to those on unmanipulated reef habitat at the same five depths. For all species, settlement on patch reefs differed significantly among depths despite uniform substratum composition. For four of the six species tested, depth-related settlement patterns on unmanipulated habitat and on patch reefs did not differ, while for the other two, depth ranges were greater on the patch reefs than on unmanipulated habitat. A second experiment examined whether depth preferences reflected variation in growth and survival when microhabitat was similar. Newly settled individuals of Chrysiptera parasema and Dascyllus melanurus were placed, separately, on patch reefs at five depths (as above) and their survival and growth monitored. D. melanurus, which is restricted to shallow depths, had highest survival and growth at the shallowest depth. Depth did not affect either survival or growth of C. parasema, which has a broader depth range than D. melanurus (between 6 and 15 m). This suggests that the fitness costs potentially incurred by settling outside a preferred depth range may depend on the strength of the depth preference.     

Dendrosoter protuberans (Hymenoptera: Braconidae), An Important Elm Bark Beetle Parasitoid

The most favourable period for Dendrosoter protuberans development occurred when wasps were introduced 16 days after the beginning of Scolytus multistriatus development (the greatest number of emerged wasps per a single parasitoid female with a very positive sexual index and high percentage of parasitised elm bark beetle larvae). No significant differences were observed in parasitoid efficiency between the 11^th and the 21^st day. The most unfavourable period for elm bark beetle parasitisation was six days after the beginning of S. multistriatus development (a low reproduction of D. protuberans, a negative sexual index and a low number of destroyed S. multistriatus larvae). The reproduction of both the elm bark beetle and D. protuberans was greater in the first, than in the second generation. The developmental period of D. protuberans in the first generation was approximately two days longer than in the second generation. Furthermore, the development of females was 1-2 days longer than the development of males. D. protuberans developed most successfully on S. multistriatus (the most abundant parasitoid reproduction with the longest developmental period and a very positive sexual index that resulted in a high percentage of parasitised larvae) and then on S. ensifer, S. pygmaeus and Pteleobius kraatzi. On the other hand, S. scolytus larvae were the poorest host of all elm species - the reproduction of D. protuberans in this species was the scarcest with the shortest period of development and a negative sexual index, hence larval parasitisation was the lowest.     

A semi-synthetic diet for rearing Dipha aphidivora (Lepidoptera: Pyralidae), a promising predator of woolly aphid in sugarcane

Dipha aphidivora (Meyrick), a lepidopteran predator of sugarcane woolly aphid (SWA) Ceratovacuna lanigera Zehntner was successfully reared for three successive generations, for the first time on a freeze-dried beef liver-based larval semi-synthetic diet. We compared biological parameters viz., larval survival, adult weight, pre-and oviposition period, fertility, fecundity and female longevity of D. aphidivora reared on the semi-synthetic diet with the predators reared on SWA. Development time of larvae reared from first instar to pupation was 20.6 days on the semi-synthetic diet and 12 days on SWA, while survival of the larvae to adults was 61.8 and 91.8% on larval semi-synthetic diet and SWA, respectively. Fecundity recorded from semi-synthetic diet (41 eggs/female) was significantly less than those produced on SWA (58 eggs/female). However, fertility and longevity of the predators reared on SWA and semi-synthetic diet did not differ significantly. The study revealed the possibility of rearing D. aphidivora larvae using synthetic diet.     

大型枝角类蚤状溞对小型热带湖泊浮游植物群落影响的研究

2006年4月10日至2006年4月22日,采用原位吊瓶实验的方法,在暨南大学校园明湖中进行了大型枝角类蚤状溞(Daphnia pulex)对浮游植物种群变化和群落结构影响的实验。实验初期,明湖浮游植物群落的组成是以飞燕角甲藻(Ceratium hirundinella)和单角盘星藻(Pediastrum simplex)为优势种。在室内将蚤状溞培养至体长2mm以上,将蚤状溞加入到装满湖水的4.5L的透明瓶中,设置4个梯度:0ind(A,对照),10ind.(B),20ind(C)和30ind(D),每个梯度有3个平行,将实验瓶置于水表层50cm处。12d后实验结束,不同处理之间的浮游植物种群数量和组成有了明显的差异。与对照组相比,飞燕角甲藻和颗粒直链藻(Meclosira granulata)密度在实验组B、C和D中下降明显;小环藻(Cyclotella)和栅藻(Scenedesmus)等小型浮游植物在牧食压力比较大的C和D瓶中密度低于对照组,而在牧食压力相对较低的B瓶中,它们的密度高于对照组,这可能是由于蚤状溞加速了水体的营养盐循环反过来促进这些小型浮游植物的生长。处理组中绿藻门的盘星藻数量没有明显的下降,这与盘星藻不能被浮游动物直接滤食有关。实验结果表明蚤状溞对浮游植物的群落数量和组成的影响程度与其自身的种群密度密切相关,也与浮游植物群落结构有关。由于飞燕角甲藻是一种细胞较大的种类,蚤状溞对它的抑制作用主要是通过机械伤害作用实现的。     

Bathymetric Variation in Recruitment and Relative Importance of Pre- and Post-Settlement Processes in Coral Assemblages at Lyudao (Green Island), Taiwan

Studies on coral communities have typically been conducted in shallow waters (∼5 m). However, in the face of climate change, and as shallow coral communities become degraded, a greater understanding of deeper coral communities is needed as they become the main reef remnants, playing a central role in the future of coral reefs. To understand the dynamics of deeper coral assemblages, the recruitment and taxonomic composition of different life-stages at 5 and 15 m depths were compared at three locations in Lyudao, southeastern Taiwan in 2010. Coral recruits (<1 cm diameter, <4 months old) were examined using settlement plates. Juvenile corals (1–5 cm, several years old) were examined with quadrats, and adult corals (>5 cm, several years to decades old) were examined using transect lines. Pocilloporid and poritid corals had similar and higher numbers of recruits at 5 m compared to 15 m, whereas acroporid recruits were more abundant at 15 m. The primary cause for the former may be larval behavior, such that they position themselves in shallow waters, while that for the latter may be the dominance of brooding acroporid species (Isopora spp.) at 15 m. The taxonomic composition, especially between recruits and juveniles/adults, was more similar at 15 m than at 5 m. These results suggest a change in the relative importance of pre- and post-settlement processes in assemblage determinants with depth; coral assemblages in shallow habitats (more disturbed) are more influenced by post-settlement processes (mortality events), while those in deeper habitats (more protected) are more influenced by pre-settlement processes (larval supply).     

Daphnia development and reproduction: Responses to temperature

1. 1.|The development times and reproduction were measured for Daphnia pulex and Daphnia magna from 5 to 30°C at 5°C increments.

2. 2.|The general trends for D. pulex and D. magna were for the duration of all juvenile instars to be less than that of adults and for the last juvenile (or adolescent) instar to be longer than all previous juvenile instars.

3. 3.|The number of juvenile instars both species pass through before adulthood is influenced by temperature with increasing numbers occurring at temperature extremes.

4. 4.|Duration of development time decreased over the entire range of increasing temperatures measured for D. pulex but increased for D. magna at 30°C in relation to 25°C.

5. 5.|Quadratic models were less desirable than simple linear logarithmic transformations of the form ln Y = ln a + b ln X for describing the temperature/development relationship.

6. 6.|The greatest young production occurs at 15 and 20°C with significant decreases occurring at temperatures above and below these.

7. 7.|The observed temperature-dependent phenomena an the ecological relationships for the two species are discussed.

Algal colonization in urchin barrens: defense by association during recruitment of the brown alga Agarum cribrosum

We investigated the process whereby juveniles of the kelp Agarum cribrosum escape grazing by the green sea urchin, Strongylocentrotus droebachiensis, on urchin barrens in the rocky subtidal zone in the Mingan Islands, northern Gulf of St. Lawrence. The highest recruitment of juvenile A. cribrosum occurred under the canopy of the large filamentous phaeophyte Desmarestia viridis, where urchin densities were markedly reduced, compared to the surrounding area. This pattern of distribution appeared to be related to the wave-induced sweeping motion of D. viridis, although currents may modify the back and forth motion of the alga by pushing the canopy towards a specific direction, thereby allowing urchins to invade the non-swept areas. The density of juveniles under D. viridis plants increased with plant size and increasing proximity to the holdfast. Living under D. viridis slightly reduced the growth rate of the A. cribrosum juveniles, but this loss in growth was clearly outweighed by the gain in protection from sea urchin grazing. The time scale over which D. viridis provides protection is in the order of months, as D. viridis is an annual alga that disappears in early autumn. This defensive association of juvenile A. cribrosum with D. viridis is possibly a successional step leading to the formation of mature stands of A. cribrosum.     

Adding coral rubble to substrata enhances settlement of Pocillopora?damicornis larvae

Settlement preferences of Pocillopora damicornis larvae were examined on artificial substrata. Planulation of P. damicornis followed a lunar cycle and the release of larvae occurred after new moon. P. damicornis larvae had the highest rates of settlement within 3 days of being presented settlement substrata. Cumulative settlement gradually increased from 3 to 8 days, and post-settlement mortality was most frequent after 8 days. Settlement experiments showed greatest settlement preference to cement tiles containing 10% coral rubble. This study suggests that physical cues are important in the settlement process, which may be useful for coral reef rehabilitation projects.     

Effect of ultrasound on cyprids and juvenile barnacles

Settlement inhibition of barnacle (Amphibalanus amphitrite) cypris larvae resulting from exposure to ultrasound was measured at three frequencies (23, 63, and 102 kHz), applied at three acoustic pressure levels (9, 15, and 22 kPa) for exposure times of 30, 150, and 300?s. The lowest settlement was observed for 23?kHz, which also induced the highest cyprid mortality. Cyprid settlement following exposure to 23?kHz at 22?kPa for 30?s was reduced by a factor of two. Observing surface exploration by the cyprids revealed an altered behaviour following exposure to ultrasound: step length was increased, while step duration, walking pace, and the fraction of cyprids exploring the surface were significantly reduced with respect to control cyprids. The basal area of juvenile barnacles, metamorphosed from ultrasound-treated cyprids was initially smaller than unexposed individuals, but normalised over two weeks' growth. Thus, ultrasound exposure effectively reduced cyprid settlement, yet metamorphosed barnacles grew normally.     

Cloning and preliminary characterization of a novel cuticular antigen from the filarial parasite Dirofilaria immitis

We have described here the cloning and partial characterization of a cDNA encoding a cuticular antigen of Dirofilaria immitis. A 48-h third-stage larval D. immitis cDNA library was immunoscreened with sera raised in mice against third-stage larval cuticles (mouse anti-L3 cuticle antisera). A strongly immunoreactive clone (L3MC4) was isolated. Sequence analysis of L3MC4 showed that it was a partial length cDNA. The missing 5′ end of the clone was amplified by PCR from D. immitis adult female first-strand cDNA using the nematode 22-base splice leader sequence and a L3MC4-specific antisense primer. The composite cDNA sequence comprised 616 bases (nDiL3MC4) encoding a full-length protein of 146 amino acids (DiL3MC4). GenBank analysis showed that DiL3MC4 shared some homology to an unknown C. elegans gene product (31%) at the amino acid level. However, there were no related filarial expressed sequence tags in the current GenBank™ database. Antibodies to recombinant DiL3MC4 (rDiL3MC4) identified a 19-kDa native antigen in the adults and in the L3 and L4 larval stages of D. immitis. In addition, the antibodies bound to the cortical layers of the L3 cuticle, as revealed by immuno-gold electron microscopy. The native protein was not detected in larval and adult excretory–secretory products. Immunoblot analysis showed that serum from a rabbit that was repeatedly injected with a small number of D. immitis third stage larvae reacted with rDiL3MC4. Thus, DiL3MC4 is a novel cuticular antigen of a filarial parasite.     

Genetic analysis of mortality in murine angiostrongyliasis costaricensis using SMXA recombinant inbred mouse strains

The SMXA recombinant inbred mouse strain set was produced by systematic inbreeding from the F2 generation of a cross between two progenitor inbred strains, A/J and SM/J, which differed markedly with respect to the patterns of infection with Angiostrongylus costaricensis. We have applied this set to genetic analysis of mouse susceptibility to this nematode infection. The mortality was variable among substrains of the SMXA RI strains, indicating the involvement of multiple genes. Linkage analysis showed several chromosomal regions closely linked to mortality; chromosome 6 (D6Rik86, 87; P0.001), 10 (D10Rik66–D10Mit12; P=0.0058), 13 (D13Rik79, 80; P=0.0096) and 17 (D17Mit28–D17Rik76; P=0.0088).