Settlement behavior of cyphonautes larvae of the bryozoan Membranipora membranacea in response to two algal substrata

Abstract. Settlement is an important process in the biphasic life histories of many marine invertebrates. Little is known regarding the fine-scale behavioral mechanisms for finding and attaching to a suitable substratum, particularly under conditions that may impose a potential challenge, such as flow. In this study, we examined the settlement behavior of cyphonautes larvae of the bryozoan, Membranipora membranacea, in response to two different algal substrata. Larvae showed a strong preference for settling on the kelp Nereocystis luetkeana over the red alga Mazzaella splendens. We then tested whether the behavioral mechanisms used by larvae to attach to these algae differed when presented with the challenge of flowing water during settlement. We found that larvae exhibited different behaviors on the two species of algae in flowing water. Larvae were more often observed in direct contact with the preferred alga (N. luetkeana) exhibiting fine-scale active search behaviors. On the less preferred alga (M. splendens), larvae were less frequently observed in direct contact with the alga, and appeared to be exhibiting broad-scale passive search behaviors along the surface of the blade. Our results suggest that cyphonautes larvae alter their behavior in response to their preferred settlement habitat.     

NOTES ON THE CAPRIMULGIDAE OF THE KAFUE BASIN

During 2002–2007, we assessed the status of the Purple Heron Ardea purpurea in Numidia, Algeria by surveying all the major wetlands in the region. We located six distinct breeding sites; four of these were not previously known. We also investigated nest site selection and determined the species’ reproductive success at Dakhla, a dunary pond during two successive years (2006 and 2007). The egg laying period was comparable to that reported for southern Europe (March-May) and a seasonal change of breeding success was recorded. A growth curve was derived for developing nestlings and the impact of an ectoparasite (mite) infestation upon nestling's growth was considered. We also examined 73 food boluses regurgitated by nestlings and identified 329 prey items. Fish, mainly Gambusia holbrookii and Cyprinus carpio, dominated by mass, whereas insects, mainly aquatic Coleopteran larvae and aeshnids (Odonata), were the most frequent prey (67.1%). Loss of habitat is identified as a major threat to the future of colonial herons in Algeria.     

Possible latitudinal clines in Antarctic intertidal and subtidal zone communities encrusting ephemeral hard substrata

Aim Encrusting faunal communities on rocks were examined from Southern Ocean intertidal and subtidal (6 m) zones to investigate potential change with latitude. Location The site locations were South Georgia (54°S [sub-Antarctic]), Signy Island (60.5°S [maritime Antarctic]) and Adelaide Island (68°S [high Antarctic]). Methods The number of taxa, degree of colonization and bryozoan growth and mortality were measured. Results The communities present were relatively simple in terms of composition and interactions. Spirorbid polychaetes (Annelida) and cheilostomatid bryozoans were the principal components. The community simplicity and dominance of unitary (compared to colonial) fauna increased with latitude. Mean annual mortality of subtidal cheilostomatids also significantly increased with latitude. The number of bryozoan species and potential aspects of community complexity increased with latitude. The growth of the bryozoan Inversiula nutrix increased with latitude reversing the general trend of decreased growth rate. Conclusions These findings suggest that disturbance may significantly increase with latitude within the Southern Ocean. The apparent reversal of typical latitudinal growth trends is probably a result of classical island biogeography, with species decreasing away from a centre of high diversity.     

Reproductive biology of viviparous and oviparous species of the leaf beetle genus Oreina

In five species of the genus Oreina Chevrolat (Coleoptera, Chrysomelidae) we compared the size of offspring, the fecundity of the females, the timing of offspring production and female investment over the season. Two of the species, O. elongata and O. luctuosa, laid eggs, while O. cacaliae, O. gloriosa and O. variabilis gave birth to larvae. Offspring size corrected for female size was similar in the two oviparous species and in the viviparous O. cacaliae. In the two other viviparous species the larvae were two to three times bigger in relation to the female. The greater size of the offspring was not traded off for lower fecundity in these latter two species, yet the production of bigger larvae was associated with a longer laying period and thereby a spreading of reproductive investment over the season. The prediction of life history theory that higher investment in individual offspring should be traded off for lower fecundity could not be confirmed. The investigation of egg and larval development showed that in one of the oviparous species, O. luctuosa, the length of the egg stage was more variable. This corroborates the view that in this species the eggs can be retained for varying times before being laid. Greater size at birth does not necessarily lead to shortened developmental times: the larval periods of O. cacaliae, O. elongata, O. gloriosa and O. variabilis were all comparable although the larvae of the first two species were relatively smaller when laid; only the small larvae of O. luctuosa needed significantly longer for their development. For all growth parameters examined the differences between species were larger than the differences between populations. A comparison of larval growth of the oligophagous species O. cacaliae on three plant genera showed that larval growth rate is influenced by the food plant. However, the plant on which the larvae grew worst is apparently not chosen for oviposition in the field. A comparison with a phylogeny of the species based on allozymes suggests that species with similar reproductive parameters are closely related, yet that viviparity evolved independently in O. cacaliae on one hand and O. variabilis and O. gloriosa on the other.     

Reproductive traits explain contrasting ecological features in sponges: the sympatric poecilosclerids Hemimycale columella and Crella elegans as examples

Our study focused on the Mediterranean species Hemimycale columella and Crella elegans, which have overlapping ecological distributions but contrasting population densities and resilience. We formulated the hypothesis that differential reproductive traits were the main cause underlying these ecological differences in the study area. The issues addressed were whether recruits compete for the substrate, either because their respective reproductive cycles overlap or/and larval performance differs between species, and whether a contrasting investment in reproduction contributed to explain their diverse densities. Both species were simultaneous hermaphrodites and incubated their larvae. The reproductive period was notably shorter in C. elegans than in H. columella, while investment in reproductive tissue was higher in individuals of H. columella than in C. elegans. In contrast, C. elegans larvae harbored larger amounts of lipids and yolk inclusions than H. columella larvae. Moreover, the former contained amazing collagen masses densely packed among their inner cells. H. columella is likely to produce a higher number of poorer equipped larvae while C. elegans seems to produce a lower number of more resistant, better fitted with energy reserves, and thus potentially more successful larvae. These reproductive patterns agree with a higher and stable density of C. elegans in the study site compared with a more variable abundance of H. columella.     

Experimental cultivation of the invasive freshwater bryozoan <Emphasis Type="Italic">Pectinatella magnifica</Emphasis>

The freshwater bryozoan Pectinatella magnifica (Leidy, 1851) is an invasive species in many countries all over the world. Although native to North America, it has been found in many countries of Europe and Asia. Pectinatella magnifica forms the largest colonial masses from all recently known bryozoan species. Culturing this organism in an aquarium has never been achieved for more than few days so far. Colonies from laboratory culture are important for various studies on its biology and life cycle of this species in experimental conditions. Young colonies successfully hatched from germinating statoblasts of P. magnifica in the laboratory and were maintained over eight weeks. Moreover, this was the first time when the compound colonies of this species were carried from its natural habitat to the laboratory, into a special aquarium system, and kept alive for more than three weeks. In both experiments the physicochemical parameters of the water (temperature, concentration of dissolved oxygen, electrolytic conductivity and pH) and changes in weight of compound colonies of P. magnifica in laboratory conditions were checked. The results found in this study are essential for understanding the invasiveness of this species and identifying methods for elimination of its ecological risks because these are closely resembling those of other invasive species.     

The effects of infection by Tetracapsuloides bryosalmonae (Myxozoa) and temperature on Fredericella sultana (Bryozoa)

The myxozoan, Tetracapsuloides bryosalmonae, exploits freshwater bryozoans as definitive hosts, occurring as cryptic stages in bryozoan colonies during covert infections and as spore-forming sacs during overt infections. Spores released from sacs are infective to salmonid fish, causing the devastating Proliferative Kidney Disease (PKD). We undertook laboratory studies using mesocosm systems running at 10, 14 and 20 °C to determine how infection by T. bryosalmonae and water temperature influence fitness of one of its most important bryozoan hosts, Fredericella sultana, over a period of 4 weeks. The effects of infection were context-dependent and often undetectable. Covert infections appear to pose very low energetic costs. Thus, we found that growth of covertly infected F. sultana colonies was similar to that of uninfected colonies regardless of temperature, as was the propensity to produce dormant resting stages (statoblasts). Production of statoblasts, however, was associated with decreased growth. Overt infections imposed greater effects on correlates of host fitness by: (i) reducing growth rates at the two higher temperatures; (ii) increasing mortality rates at the highest temperature; (iii) inhibiting statoblast production. Our results indicate that parasitism should have a relatively small effect on host fitness in the field as the negative effects of infection were mainly expressed in environmentally extreme conditions (20 °C for 4 weeks). The generally low virulence of T. bryosalmonae is similar to that recently demonstrated for another myxozoan endoparasite of freshwater bryozoans. The unique opportunity for extensive vertical transmission in these colonial invertebrate hosts couples the reproductive interests of host and parasite and may well give rise to the low virulence that characterises these systems. Our study implies that climate change can be expected to exacerbate PKD outbreaks and increase the geographic range of PKD as a result of the combined responses of T. bryosalmonae and its bryozoan hosts to higher temperatures.     

Epizoic and parasitic rotifers

Many rotifer species live in close association with plants or other animals. Most of these associations are of a commensal or synoecious nature, some rotifer species having lost the ability to live independently. Few rotifers are true parasites, actually harming their hosts.The Seisonidae, Monogononta and Bdelloidea include epizoic and parasitic species. The most widely known are probably the parasites of colonial and filamentous algae (e.g. Volvox, Vaucheria). However, rotifers are also found on a wide range of invertebrates: colonial, sessile Protozoa; Porifera; Rotifera; Annelida; Bryozoa; Echinodermata; Mollusca, especially on the shells and egg masses of aquatic gastropods; Crustacea, including the lower forms (e.g. Daphnia, Asellus, Gammarus) and in the gill chambers of Astacus and Chasmagnathus; the aquatic larvae of insects. There appear to be few records of epizoic or parasitic rotifers among vertebrates, apart from Encentrum kozminskii on carp, Limnias ceratophylli on the Amazonian crocodile, Melanosuchus niger, and an unidentified Bdelloid apparently living as a pathogenic rotifer in Man.     

Host-Specific Myrmecophily and Myrmecophagy in the Tropical Coccinellid Diomus thoracicus in French Guiana

A variety of arthropods, particularly insects, have developed myrmecophilous interactions with ants to gain access to resources and/or for protection. Among these myrmecophiles, only a few examples have been documented in the Coccinellidae, most of them involving species able to feed on ant-tended Hemiptera. We report here a new case of obligate myrmecophily in the coccinellid Diomus thoracicus. Larvae are invariably and exclusively found in the nests of the ant Wasmannia auropunctata and seem to rely on ant brood as their only food source. Not only do ant workers show no aggressiveness toward the D. thoracicus larvae in their behavioral interactions at the colonial level, but also at the species level; while coccinellid adults are always attacked. The integration of the larvae inside of the ant nests is based on their chemical mimicry of the host's cuticular cues. Therefore, given the presence of the D. thoracicus larvae inside of the ant's nest, their predation on Wasmannia brood and their chemical mimicry, this species can be considered a specific parasite of W. auropunctata. Overall, this new case of myrmecophily not only specifically involves a highly invasive ant species, but also provides insights into the evolution of myrmecophily and myrmecophagy in coccinellids.     

Persistence and space preemption explain species‐specific founder effects on the organization of marine sessile communities

Community assembly may not follow predictable successional stages, with a large fraction of the species pool constituted by potential pioneering species and successful founders defined through lottery. In such systems, priority effects may be relevant in the determination of trajectories of developing communities and hence diversity and assemblage structure at later advanced states. In order to assess how different founder species may trigger variable community trajectories and structures, we conducted an experimental study using subtidal sessile assemblages as model. We manipulated the identity of functionally different founders and initial colony size (a proxy of the time lag before the arrival of later species), and followed trajectories. We did not observe any effects of colony size on response variables, suggesting that priority effects take place even when the time lag between the establishment of pioneering species and late colonizers is very short. Late community structure at experimental panels that started either with the colonial ascidian Botrylloides nigrum, or the arborescent bryozoan Bugula neritina, was similar to control panels allowed natural assembling. In spite of high potential for fast space domination, and hence negative priority effects, B. nigrum suffered high mortality and did not persist throughout succession. Bugula neritina provided complex physical microhabitats through conspecific clustering that have enhanced larval settlement of late species arrivals, but no apparent facilitation was observed. Differently, panels founded by the encrusting bryozoan Schizoporella errata led to different and less diverse communities compared to naturally assembled panels, evidencing strong negative priority effects through higher persistence and space preemption. Schizoporella errata founder colonies inhibited further conspecific settlement, which may greatly relax intraspecific competition, allowing resource allocation to colony growth and space domination, thus reducing the chances for the establishment of other species.     

Egg Recognition and Social Parasitism in Formica Ants

Nests of social insects are an attractive resource in terms of nutrition and shelter and therefore targeted by a variety of pathogens and parasites that harness the resources of a host colony in their own reproductive interests. Colonies of the ants Formica fusca and F. lemani serve as hosts for mound‐building Formica species, the queens of which use host colonies during colony founding. Here, we investigate whether workers of the host species can mitigate the costs imposed on them by invading parasite queens by recognizing and selectively removing eggs laid by these queens. We used behavioural assays, allowing host workers to choose between con‐colonial eggs and eggs laid by the parasite species F. truncorum. We show that workers of both host species discriminate between the two types of eggs in favour of con‐colonial eggs. Moreover, workers of F. fusca rejected more con‐colonial eggs than F. lemani. This higher rate of error in F. fusca may reflect a greater selectivity or a greater difficulty in discriminating between the two egg types. Nevertheless, both host species removed parasite eggs at a similar rate, when these were artificially introduced into the colonies, although some eggs remained after 10 d. In addition, upon receiving parasite eggs, host workers started to lay unfertilized male‐destined eggs within 6 d, thus employing an alternative pathway to gain direct fitness when the resident queen is no longer present and the colony is parasitized.     

Ethnobotany of Horseradish (<Emphasis Type="Italic">Armoracia rusticana</Emphasis>, Brassicaceae) and Its Wild Relatives (<Emphasis Type="Italic">Armoracia</Emphasis> spp.): Reproductive Biology and Local Uses in Their Native Ranges

Ethnobotany of Horseradish ( Armoracia rusticana , Brassicaceae) and Its Wild Relatives ( Armoracia spp.): Reproductive Biology and Local Uses in Their Native Ranges. Horseradish (Armoracia rusticana) has been cultivated for its root for over 2,000 years. Today, commercial production takes place primarily in North America and Europe where A. rusticana is propagated exclusively vegetatively. Like many vegetatively-propagated crops, cultivated A. rusticana plants are generally sterile. Armoracia rusticana is one of three species in the genus: A. macrocarpa is native to the Central Danube River Basin, A. rusticana is putatively native to Eastern Europe, and A. sisymbrioides is native to Siberia. Documenting the ways in which the three Armoracia species are used in their putative native ranges and the reproductive biology of Armoracia in these regions will advance understanding of the role of humans in the evolution of sterility in A. rusticana. In this broad-scale ethnobotanical analysis, we document 1) which Armoracia species are used in different geographic regions, 2) the reproductive biology of cultivated and natural Armoracia populations, and 3) the uses of Armoracia species in their native areas. Fieldwork was conducted in Austria, Bulgaria, Romania, and Russia. Ethnobotanical data were collected through written questionnaires. In all areas visited, A. rusticana is the primary Armoracia species; however, there is regional variation in the way the plant is used. Armoracia rusticana is propagated vegetatively and is not found in the wild. Armoracia macrocarpa and A. sisymbrioides reproduce by seed in nature. These data provide an understanding of contemporary uses and management strategies of Armoracia species in their native ranges, demonstrate the differential reproductive biology of a clonally-propagated crop relative to its wild progenitors, and emphasize the importance of ethnobotanical data collection for crops and their wild relatives from a broad geographic range.     

Near reef abundance and on–offshore distribution of tuna larvae around Minicoy Island in Lakshadweep Sea,India

ABSTRACT

Near reef abundance and coast–offshore distribution of tuna larvae around Minicoy Island were studied based on three cruises carried out on board FORV Sagar Sampada during 2014–15. The samples were collected by oblique tows using bongo twin nets (300 micron) for three seasons (Fall inter-monsoon (FIM), Spring inter-monsoon (SIM) and Summer monsoon (SM)) in six on–offshore stations ranging from 1 to 20?km off-shore of the Minicoy. Results indicated that the tuna larvae aggregate in the near reef waters and total tuna larvae abundance (all the species combined) shows a decreasing trend towards off-shore. Tuna larvae are found to be more abundant during SM and FIM. Comparison of the species-wise variation in tuna larvae for one season (SM) indicated that the larvae of Euthunnus affinis (32.5%) dominated among the tuna species, followed by Auxis spp. (31.6%), Thunnus spp. (3.9%) and Katsuwonus pelamis (2.7%). A pattern of highest larvae abundance in the near reef areas was observed except for Thunnus larvae. GAM indicated that mean seawater temperature and mean seawater salinity are the major environmental factors that significantly influence the larvae abundance, particularly for Thunnus followed by Katsuwonus pelamis, E. affinis and Auxis spp. The study confirms that the environmental parameters show high variation between stations and among transects as a result of the complex oceanographic characteristics around the islands. Further studies with better understanding of the larvae dynamics and their association with the environmental parameters including physical processes are essential to elucidate these relationships more clearly.     

Clinal variation, intraspecific heterochrony, and microevolution in the Permian bryozoan Tabulipora carbonaria

Clinal geographic variation across an onshore-to-offshore environmental gradient occurs in the Lower Permian trepostome bryozoan Tabulipora carbonaria collected from three widespread calcareous shales of the Wreford Megacyclothem of Kansas. Separate multivariate statistical tests dclineate significant differences between populations within each shale unit, and between onshore, intermediate, and offshore populations pooled across all three shales, suggesting that even weakly-developed gradients may initiate different intraspecific morphological responses. Growth trajectories for populations along the cline also differ significantly, indicating astogenetic (developmental) heterochrony. Populations from onshore habitats are generally paedomorphic relative to those in more offshore settings, and exhibit pre- and postdisplacement and hypermorphosis in zooecial and acanthostyle characteristics. These heterochronic processes may have increased colonial reproductive potential and the efficiency of water flow for feeding and waste disposal in colonies from onshore habitats. Variation from tightly constrained development (astogenetic plasticity) decreased monotonically in an onshore-to-offshore direction; canalized growth may characterize colonies from more stable offshore habitats, whereas greater flexibility during the growth of colonies from unstable onshore biotopes may have increased their rate of survival. Populations of colonies from stratigraphically successive calcareous shales of the Wreford display patterns of growth that are nearly identical to those found in an offshore-to-onshore direction along the cline. Both clinal and temporal patterns probably resulted from selection for more paedomorphic morphologies in onshore, perhaps unstable, habitats and represent microevolution in T. carbonaria. These local adjustments to environmental conditions may produce variation that affects the rate of macroevolutionary change. □Bryozoa, clines, heterochrony, microevolution, Permian, variability.     

Seasonality and life histories of two endemic New Zealand carabid beetles (Coleoptera: Carabidae): Mecodema oconnori Broun and Megadromus capito (White)

Abstract

Seasonal activity and life histories of two large endemic carabid beetles, Mecodema oconnori Broun and Megadromus capito (White), were investigated in two lowland forest remnants in the Horowhenua region, North Island, New Zealand. Seasonal activity was estimated from manual searches of surface refuges, plus pitfall trapping from October to March. Adults of both species were present throughout the year, although M. oconnori was much more abundant in spring. Megadromus capito larvae were caught from October to March, and the three instars overlapped temporally, implying a long period of larval emergence and development. Adult females were dissected to investigate their reproductive phenology. Both species carried eggs for at least 6 months of the year. M. oconnori females contained very few eggs, with a maximum of three eggs each and a mean of 1.6 eggs per female. Mg. capito females had up to 28 eggs each and a mean of 10.5. Both species appear to have low egg loads, a relatively long period of reproductive activity and long life spans, but longer‐term studies on the pre‐adult stages and reproductive activity of adults are required to complete understanding of their life cycles.     

Epibiont foraminifera of Sertella frigida (Waters) (Bryozoa, Cheilostomata) from Terranova Bay, Ross Sea, Antarctica

Several arenaceous and calcareous foraminifera epibionts of Sertella frigida (Waters) (Bryozoa, Cheilostomata), collected in Terranova Bay during the 1989–90 Italian Antarctic expedition, were studied. Arenaceous species, such as Psammosphaera fusca Schulze f. adhearescens Rhumbler, Haplophragmoides canariensis d'Orbigny, Portatrochammina antarctica (Parr), and Trochammina arctica Hedley, Hurdle and Burdett, are a conspicuous component of the epifaunal community. The calcareous foraminifers encrusting S. frigida are mostly represented by Cibicides refulgens Montfort and Rosalina globularis d'Orbigny. Foraminifera were found only in the basal part of the bryozoan colonies. The presence of many juveniles provided evidence that foraminifers were reproducing at the time of sampling. Psammosphaera fusca f. adhaerescens was found exclusively inside the zooids of S. frigida. The large number of individuals associated with the bryozoan suggest that P. fusca f. adhaerescens finds optimum conditions (shelter, food, and grains for shell building) for growth inside the zooids of S. frigida. The other species occurred most commonly adhered to the trabeculae or to the zooid orifices of S. frigida. An elevated position offers a better chance to catch food particles from the overlying water column. Foraminifers could benefit also from bryozoan feeding currents. The availability of resuspended organic material could provide a more consistent source of food in a highly seasonal oligotrophic environment. Received: 8 February 1995/Accepted: 30 June 1996     

Simulated Intraspecific Usurpations in Paper Wasps: Different Reproductive Tactics Affect Foreign Brood Destruction in Polistes fuscatus and Polistes dominulus

We simulated intraspecific usurpation in two species of paper wasps at a field site in Michigan to compare the species' treatment of foreign brood. Queens of Polistes fuscatus, a species that commonly uses intraspecific usurpation as an alternative reproductive tactic, destroyed significantly fewer large larvae and significantly more small larvae than queens of P. dominulus, a sympatric species that rarely usurps. The pattern of brood destruction exhibited by P. fuscatus was consistent with the previously published findings that P. fuscatus usurpers destroy reproductive‐destined brood (eggs and small larvae), but not worker‐destined brood (large larvae and pupae) that are subsequently used by a usurper to raise her own reproductives. The pattern of brood destruction displayed by P. dominulus differed from that of Polistes species that frequently engage in intraspecific usurpation. The brood destruction pattern in P. dominulus may have been shaped by nest adoption, a common alternative reproductive tactic in this species. If so, it is not clear why P. dominulus would destroy large, worker‐destined larvae.     

Host use of a specialist lichen-feeder: dealing with lichen secondary metabolites

Host use by herbivores is largely determined by host properties such as nutrient content and chemical defence against foragers. The impacts of these attributes on a herbivore may largely depend on its life cycle stage. Lichen species are known to differ in nutritional quality and level of chemical defence and, consequently, vary as fodder for herbivores. The aim of this study was to explore the impact of several lichen species and the presence of their secondary metabolites on their use as hosts by a specialist lichen-feeder, Cleorodes lichenaria. This study also addressed, for the first time, how a specialist lichen-feeder deals with different lichen secondary metabolites. In the beginning of their development, larvae grew better on Xanthoria parietina than on the other host lichens, whereas older larvae grew best on Ramalina fraxinea. Lichen secondary chemicals in R. fraxinea and Parmelia sulcata hindered larval growth in the beginning but after 75 days lichen secondary chemicals had no impact on the mass of larvae. Physodic acids in Hypogymnia physodes were lethal to larvae. In general, larvae metabolized 70–95% of ingested lichen secondary chemicals and the rest of these were excreted in frass. Lichen secondary metabolites in P. sulcata restrict and in H. physodes prevent their use as a host for C. lichenaria larvae. Our main finding, the ability of larvae to metabolize several lichen secondary metabolites, indicates digestive adaptation to these chemicals. No signs of sequestration of these chemicals were found.