Molecular data suggest a hybrid origin for the invasive Caulerpa racemosa (Caulerpales,Chlorophyta) in the Mediterranean Sea

Morphological data has provided a basis for the hypothesis that three taxa belonging to the Caulerpa racemosa complex occur in the Mediterranean Sea: var. turbinata–uvifera, var. lamourouxi, and the `invasive variety'. In order to test this hypothesis and to determine the origin of the `invasive variety', the transcribed spacer ITS1–ITS2 and an 18S ribosomal DNA (rDNA) intron were analysed from 16 isolates of Caulerpa racemosa. The `invasive variety' shows intraindividual polymorphism for both types of sequences. The ITS1–ITS2 data confirm that the three morphological varieties of C. racemosa from the Mediterranean Sea are distinct taxonomic units. The 18S intron data suggest that the new `invasive variety' could be a recent hybrid between var. turbinata–uvifera and an unknown tropical strain. Incongruence between the phylogenetic tree computed from ITS1–ITS2 regions and the 18S intron indicates that homogenization processes of concerted evolution have run at different rates.     

Blitzkrieg in a Marine Invasion: Caulerpa racemosa var. cylindracea (Bryopsidales, Chlorophyta) Reaches the Canary Islands (North-East Atlantic)

On the basis of morphological and genetic studies (rDNA ITS1, 5.8S, ITS2, and a 18S rDNA intron), we confirm here that Caulerpa racemosa var. cylindracea (Sonder) Verlaque, Huisman et Boudouresque, a southwestern Australian taxon recently introduced into the Mediterranean Sea also occurs in the Canary Islands. This is the first report of C. racemosa var. cylindracea in the Atlantic. It was observed for the first time in the Canary Archipelago in 1997–1998. The speed and regional scale of expansion (north Atlantic Ocean and the Mediterranean Sea) of this invasive species appear to be among the most dramatic ever recorded. The possible outcome of this introduction in the Atlantic is discussed.     

Removal of methylene blue by invasive marine seaweed: Caulerpa racemosa var. cylindracea

Caulerpa racemosa var. cylindracea is one of the well-known invasive species in the Mediterranean Sea. In the present study, dried biomass of C. racemosa var. cylindracea was shown to have adsorption capacity for methylene blue. The adsorption reached equilibrium at 90 min for all studied concentrations (5-100mg/L). The pseudo-second-order model is well in line with our experimental results. There was a sharp increase in the adsorbed dye amount per adsorbent amount from 3.3 to 16.7 g/L, then a slight increase up to 66.7 g/L was observed. Langmuir and Freundlich's models were applied to the data related to adsorption isotherm. According to Langmuir's model data, the observed maximum adsorption capacity (qm) was 5.23 mg/g at 18 degrees C. The enthalpy of adsorption was found to be 33 kJ/mol, which indicated a chemical adsorption between dye molecules and C. racemosa var. cylindracea functional groups.     

The indirect role of nutrients in enhancing the invasion of Caulerpa racemosa var cylindracea

The effects of nutrients dynamics on biological invasions in marine habitats have not been fully investigated. The present study aimed to evaluate the role of nutrient pollution in determining the successful invasion of the introduced seaweed Caulerpa racemosa var. cylindracea in a Mediterranean subtidal rocky habitat. To this end, a manipulative field experiment was carried out by supplying the water column with nutrients for a one-year period. Afterwards C. racemosa was transplanted in both nutrient-enriched and control macroalgal assemblages. Results showed that the penetration and biomass of C. racemosa in the assemblages previously treated with nutrients were higher than in control ones, indicating that communities stressed by nutrient pollution are more vulnerable to invasion. The nutrient enrichment changed the structure of natural macroalgal assemblages, mainly by encouraging the growth of turf species. No significant effects were detected for the erect and prostrate layers and for the number of species, suggesting that the susceptibility of the community to invasion may depend more on the morpho-functional identity of macroalgae rather than on the diversity of assemblages. Changes in the availability of seawater nutrients may foster the spread of C. racemosa, both by enhancing its growth and eroding the natural resistance of macroalgal assemblages. Thus, the maintenance of good water quality may play an important role in containing both C. racemosa and other introduced seaweeds.     

Comparative study of the growth of the two co-occurring introduced green algae Caulerpa taxifolia and Caulerpa racemosa along the Tuscan coast (Italy,western Mediterranean)

Caulerpa taxifolia and C. racemosa are tropical green algae now found in the Mediterranean, where the former was accidentally introduced and the latter presumably also introduced; both species have spread rapidly and invaded a variety of habitats. Since 1996, the two algae have co-inhabited a region along the Tuscan coast near Livorno. The present study has been carried out to evaluate the invasive capacity of the two species. After one year, the increase in surface area was 67.5 % for C. taxifolia and 284.8 % for C. racemosa, while the increase in patch number was 11.4 % and 121.4 % respectively. The stolon growth of the two algae followed a similar trend, as is showing the same temporal variations with the maximum growth rate between August and October. At the end of the study period, the increase in stolon length and number was significantly higher in C. racemosa than in C. taxifolia. The results of the present study show a higher invasive potential in C. racemosa than in C. taxifolia, although further studies are necessary to evaluate the competition between the two algae and the effect of their simultaneous presence on the benthic community.     

Habitat formed by the invasive macroalga Caulerpa filiformis (Suhr) Hering (Caulerpales,Chlorophyta) alters benthic macroinvertebrate assemblages in Peru

The green macroalga Caulerpa filiformis has been spreading on shallow soft sediment habitats along the Peruvian coast, colonizing previously unvegetated sediments to create monospecific meadows. We examined the nature of the impact of C. filiformis meadows on the density, taxonomic richness and assemblage structure of epifaunal and infaunal benthic macroinvertebrates. Specifically, we tested whether the spread of C. filiformis has resulted in different macroinvertebrate assemblages than those formed by the dominant native macroalgae (i.e., Rhodymenia spp.) and unvegetated sediments. Surveys were undertaken in two bays in each of two locations, in central and southern Peru, during winter 2017 and summer 2018. In general, our results show that macroinvertebrate assemblages were similar across all three habitats, although there were some differences, related to location and time, but with no clear patterns observed. Taxonomic richness and density was generally higher in the vegetated habitats than the unvegetated habitat, and where there were differences between the two vegetated habitats there was no consistent pattern of which habitat supported the highest richness or density. Given invading C. filiformis is primarily colonizing unvegetated habitats it would appear that this species is creating a new niche which supports similar assemblages, but higher taxonomic richness and density than unvegetated habitats. While our study suggests that C. filiformis is having a limited ecological impact we recommend that actions be put in place to limit the spread of this invasive species at the same time as increasing monitoring of the ecological impacts of this species as lags in the ecological impacts of invasive species are common.

     

Further confirmation of the presence of DNA in the pyrenoid core of the siphonous green algal genus Caulerpa (Caulerpales, Ulvophyceae, Chlorophyta)

We re-examined the distribution of chloroplast DNA (ct-DNA) in the pyrenoid core of Caulerpa okamurae Weber van Bosse and C. lentillifera J. Agardh by fluorescence microscopy after staining the squashes and Technovit sections with DNA fluorochromes such as 4′6-diamidino-2-phenylmdole (DAPI), ethidium bromide, Hoechst 33258 and chromomycin A3. All fluorochromes stained specifically the pyrenoid core on the squashes and Technovit sections. In addition, we present new data on the localization of ct-DNA in the pyrenoid core of two other species of the genus Caulerpa: C. cactoides (Turner) Agardh and C. geminata Harvey.     

Phylogenetic analyses of Caulerpa taxifolia (Chlorophyta) and of its associated bacterial microflora provide clues to the origin of the Mediterranean introduction

The accidental introduction of Caulerpa taxifolia into the Mediterranean is no longer under dispute. What has eluded researchers until now, is definitive evidence for the original, biogeographical source population. Here we present two independent lines of evidence that support an Australian origin for the Mediterranean populations of C. taxifolia. First, we reanalysed algal rDNA-internal transcribed spacer (rDNA-ITS) sequences, combining previously published sequences from different studies with 22 new sequences. The ITS sequence comparison showed that the Australian sample is the sister group of the Mediterranean-aquarium clade. Second, cloned bacterial 16S rDNA gene sequences were analysed from the associated microflora of C. taxifolia collected from Australia, Tahiti, the Philippines and the Mediterranean. Five bacterial lineages were identified, of which three were dominant. Alpha Proteobacteria were the most abundant and were found in all samples. In contrast, members of the beta Proteobacterial line and Cytophaga-Flexibacter-Bacteroides line (CFB) were mainly associated with Mediterranean and Australian samples. Frequency distributions of the five bacterial lineages were significantly different among biogeographical locations. Phylogenetic analyses of the 54 bacterial sequences derived from the four C. taxifolia individuals resulted in a well-resolved tree with high bootstrap support. The topologies of the beta Proteobacteria and CFB mirror the geographical sources of their algal hosts. Bacterial-algal associations provide an identification tool that may have wide application for the detection of marine invasions.     

Diterpenoid metabolites from pacific marine algae of the order Caulerpales (Chlorophyta)

Chemical investigations of two new species of Udotea, reinvestigation of Tydemania expeditionis and Chlorodesmis fastigiata from Guam and Saipan, Marianas Islands, as well as studies of Caulerpa brownii from Southern Australia, have led to the isolation of four new related diterpenoids and several previously described metabolites. These metabolites possess antimicrobial activities and inhibit cell division in the fertilized sea urchin egg bioassay.     

First record of Caulerpa lentillifera J. Agardh (Bryopsidales,Chlorophyta) from China

ABSTRACT

An edible green algal species Caulerpa lentillifera J. Agardh is reported from China for the first time. The species was collected from the southwest of Hainan Island and morphologically identified to be C. lentillifera based on the grape-like branches arising from cylindrical stolons. Phylogenetic analysis using tufA and rbcL DNA sequences also confirmed the monophyly of C. lentillifera-microphysa clade.     

An investigation on the antioxidant status of the invasive alga Caulerpa racemosa var. cylindracea (Sonder) Verlaque, Huisman, et Boudouresque (Caulerpales, Chlorophyta)

Caulerpa racemosa var. cylindracea is a potential invader of the Mediterranean Sea and 11 Mediterranean countries are under threat from this alga. In the present study, in order to investigate seasonal changes in the antioxidant status of C. racemosa var. cylindracea, antioxidant enzyme activities such as superoxide dismutase (SOD), catalase (CAT), glutathione peroxidase (GSH-Px), and lipid peroxidation (LPO) levels were determined in C. racemosa var. cylindracea samples, which were collected in May 2003, September 2003, December 2003, and March 2004 from Turkish coastlines. SOD and CAT activities showed similar trends. These enzymes increased from May to December, then sharp decreases were observed. On the other hand, GSH-Px enzyme activities showed a fluctuation between May 2003 and March 2004. In contrast to increases in SOD and CAT activities up to December 2003, LPO level decreased in this period. No significant correlation was observed between antioxidant status and solar radiation. In conclusion, the antioxidant status of C. racemosa var. cylindracea is strictly not affected by both solar radiation and seawater temperature; however, the growth of epiphytes on fronds may change antioxidant status. Further investigations are strongly warranted to understand the contributions of non-enzyme-based antioxidants such as glutathione, vitamin E, and vitamin C.     

Rhizome gravitropism precedes gravimorphogenesis after inversion of the green algal coenocyte Caulerpa prolifera (Caulerpales)

Previous investigations reported that the marine alga Caulerpa prolifera, a giant coenocyte, had a fast morphogenetic response to a change in position with regard to gravity: the next rhizoid formed approximately 1 day later on the new underside of the inverted rhizome (Jacobs and Olson, 1980, American Journal of Botany 67: 141–146). Preceding the change in site of rhizoid development was a striking accumulation of amyloplasts at the new prospective rhizoid-initiation site in the rhizome tip. Detailed monitoring with video equipment of inverted Caulerpa plants, growing under controlled conditions, now reveals that after the amyloplast accumulation, but before the gravimorphogenetic effect on rhizoid development, the inverted rhizome tip shows a negative gravitropism that restores it to its normally upturned position. Because the experiments were performed with two-sided illumination, this was clearly not a phototropic response.     

Biochemical composition of enigmatic green macroalgae,Caulerpa macrodisca Decaisne (Bryopsidales,Chlorophyta)

Journal of Applied Phycology - The green macroalga Caulerpa macrodisca Decaisne was recently reported in East Malaysia with limited information recorded on the nutritional properties of...     

Pollination partners of Mucuna macrocarpa (Fabaceae) at the northern limit of its range

Mucuna macrocarpa is a plant found in tropical and subtropical regions that requires an “explosive opening.” Explosive opening is the process that exposes the stamen and pistil from the opening of the carina. This process is needed for cross pollination; however, the plant cannot open itself and opening by an animal is needed. The most common opener of Mucuna flowers is several nectar‐eating bats (e.g., Syconycteris), but the flying fox, Pteropus dasymallus, is the only opener of M. macrocarpa on the subtropical island of Okinawajima. Here, we present the explosive openers and possible pollinators in the northernmost and temperate Kamae region, Kyushu, Japan, where nectar‐eating bats are absent. The Japanese macaque, Macaca fuscata, and the Japanese marten, Martes melampus, were the explosive openers observed during our survey in Kamae. Martens opened flowers using their snout in a manner similar to that of the flying fox, whereas macaques opened flowers using their hands. This is the first time that an animal has been observed opening these flowers with its hands rather than snout. In total, 97% (n = 283) of explosively opened flowers were opened by macaques, and the macaque largely contributed to the overall flower opening. Because many pollen grains become attached to the explosive openers, they are considered to be primary pollinators. Furthermore, two bee species, Apis cerana japonica and Bombus ardens ardens, also visited opened flowers and collected pollen, and they were possibly secondary pollinators.     

The oogonial aperture of Oedogonium decipiens var. decipiens (Oedogoniales, Chlorophyta) and its systematic significance

Pollen morphology of ten Brazilian species within the South American clade (ca. 20 species) of Symplocos section Barberina (Symplocaceae) was analyzed with scanning electron microscopy to assess their reported androdioecious breeding system. All species exhibited pollen dimorphism. Pollen from male individuals is well developed and 3-colporate, whereas that from morphologically hermaphroditic individuals is malformed, often completely fragmented, lacks cytoplasm, and has no germination pores. Our results suggest that the morphologically hermaphroditic species of S. section Barberina with malformed pollen are cryptically dioecious.     

Caste differentiation and seasonal changes in Vespa velutina (Hym.: Vespidae) colonies in its introduced range

Since its introduction in France 10 years ago, the yellow‐legged hornet, Vespa velutina, has rapidly spread to neighbouring countries (Spain, Portugal, Belgium, Italy and Germany). It showed efficient social traits facilitating its invasive success. Only scarce and incomplete natural history studies were known from its native distribution area. Studying the biology of this species in its invasive distribution range was thus a prerequisite to the implementation of efficient control methods in a near future. During a 3‐year field survey, we collected 77 nests to investigate several of the species' key colony characteristics. Our results enabled us to accurately quantify each of the castes and to better understand their synchronicity throughout the season. Our study showed that mature nests are able to produce up to 13 000 individuals and that the size of mature nests is correlated to the number of individuals produced. This correlation enables the inference of one characteristic from the other. Furthermore, each mature nest can produce up to several hundreds of potential founder queens, a crucial datum in the light of today's unregulated spring queen trapping control campaigns. In addition, nest dissections enabled to record the incidence of nest relocation for the first time in this species. Results are discussed with regards to what is known in other Vespidae species, with a focus on Vespula species that are known to be invasive in many other countries worldwide.     

Morphological and molecular clarification of the enigmatic Caulerpa floridana W.R. Taylor (Chlorophyta,Bryopsidales) from the Dry Tortugas,Florida

Morphological and molecular evidence is provided to further document the status of the enigmatic taxon known as Caulerpa floridana W.R. Taylor from White Shoal, Dry Tortugas, Florida. DNA sequencing of three historical herbarium specimens (WRT329, WRT345 and WRT349) housed at the University of Michigan Herbarium (MICH) demonstrated the molecular separation of this species based on the reconstruction of 931 nucleotides of the chloroplast gene tufA. Caulerpa floridana is sister to the western Atlantic endemic C. ashmeadii Harvey and an unknown Caulerpa taxon from the Florida Middle Grounds. Caulerpa floridana most reliably differs from C. ashmeadii by the presence of a sharp, unequivocal apiculus at the tip of each pinnule. A morphological review of southwestern Atlantic records of C. floridana from Brazil excludes these reports as representative of the species.     

Species and ecads of Caulerpa (Ulvophyceae,Chlorophyta) in Malesia (South-East Asia): Taxonomy,biogeography and biodiversity

In the present paper 29 species ofCaulerpa from Indonesia, the Philippines and Papua New Guinea are listed, with reference to their distribution and occurrence of ecological phenotypes (ecads). Other names ofCaulerpa taxa recorded for Malesia in literature are dealt with in the paragraph on excluded and changed names. The variability and nomenclature of mainly the species recorded from Indonesia are discussed, especially regarding the implications of the recognition of ecads in several species. The ecads in the sectionSedoideae are discussed as an example of use of the designation ecad for a number of growth forms. Biogeography of the genusCaulerpa is discussed with focus on areas of high biodiversity, both inside and outside Malesia, followed by a discussion on biodiversity assessment. It is suggested that the genusCaulerpa is a good indicator for the quantitative assessment of biodiversity. The need for a modern world-monograph of this genus is stressed. Such a monograph can be a stimulus for ecosystem studies ofCaulerpa stands.     

Research note: Induction of gamete discharge by hypertonic treatment in the green alga Bryopsis plumosa (Caulerpales,Chlorophyta)

Brief treatment with hypertonic solutions induced gamete discharge from gametangia in the coenocytic green alga Bryopsis plumosa (Hudson) C. Agardh. It is known that gamete discharge in this alga is triggered by a change from darkness to light. However, in this study mature gametangia, incubated for 30–120 s in artificial seawater (ASW) with an additional 0.4–0.6 M NaCl and then transferred into pure ASW, discharged gametes in darkness. The treatment did not affect the motility of the gametes. Addition of sucrose (1.0–1.2 M) to ASW also induced gamete discharge in darkness. Similar results were obtained by adding KCl (0.4 M) or mannitol (1.2 M) to ASW. Continuous incubation of gametangia in such hypertonic solutions also induced gamete discharge but led to a delay and a reduction in the rate of gamete discharge, and a loss of gamete motility. In gametangia treated with the hypertonic solutions, as well as in those exposed to light, shortening of the gametangial length was observed before gamete discharge.