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Course:EOSC270/2023/Killer Algae in the Mediterranean

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Introduction/ background.

What is C. taxifolia?

Caulerpa taxifolia

Caulerpa taxifolia (C. taxifolia) is an invasive strain of green macroalgae located in the Mediterranean. Sea[1]. The C. taxifolia strain was created in Australia; the goal for it was so it could be used in and survive aquarium environments.[2]. The strain was implemented into aquariums as decorations, as well as for their great nutrient export[1]. The C. taxifolia strain was later commercialized and expanded through trade, which is how it came to areas surrounding the Mediterranean Sea[1].

How did it enter the ecosystem?

Where Monaco is in respect to the Mediterranean Sea.

C. taxifolia accidentally entered the Mediterranean Sea through a water circuit from the Oceanographic Museum of Monaco, in Monaco[3]. The process of the water circuit is when a pump moves water in an aquarium tank out and filters the water, in order for the tank to be cleaned, and then back into the tank.[2][4] When the water is being filtered, it filters out debris, uneaten food, and toxic compounds, which is how some C. taxifolia were able to escape through the water circuit and into the Mediterranean Sea[3][4].

What is the location?

When C. taxifolia entered the Mediterranean Sea, it spread quickly along the northwestern Mediterranean coastline[5]. C. taxifolia was along the coast of Croatia, France, Italy, Monaco, Spain, Tunisia, and Turkey[5]. Although C. taxifolia is native to Southeast Asia, the Pacific Islands, and Australia, it thrives in tropical waters[5][6]. Part of the reason why C. taxifolia is native to those areas in the Indo-Pacific, is that C. taxifolia has more predators in those areas than in the Mediterranean Sea[6].

The problem: Invasive C. taxifolia outbreak.

What is the C. taxifolia outbreak?

Dense meadow of C. taxifolia

Once C. taxifolia entered the Mediterranean Sea, it spread rapidly through vegetative propagation[1][6]. Vegetative propagation is a form of asexual reproduction, where part of the thallus, which is a vegetative plant body, breaks apart into smaller fragments, and this allows C. taxifolia to colonize disturbed habitats quickly[6][7]. Since C. taxifolia exhibits invasive behavior when in non-native ecosystems, it formed dense meadows that impacted native seagrass[1][6]. Those dense meadows compete with the native seagrass Posidonia oceanica for light, space, and nutrients[1]. P. oceanica is an important species ecologically in the Mediterranean, and any changes in its density and quality would majorly impact food webs as they are a spawning ground and nursery[1]. The dense meadows C. taxifolia form an obstruction to fish that feed on benthic invertebrates[1].

In areas where C. taxifolia invaded P. oceanica seagrass beds, the total species richness, density, and biomass of fish are much lower in the invaded seagrass beds than in the native P. oceanica seagrass beds[1]. C. taxifolia was mainly found in depths of 0 to 30 m and on rocky and sandy substrates and on dead mattes of P. oceanica[8]. C. taxifolia is able to survive in varying habitats, which enhances its invasive behavior when in non-native ecosystems, like the Mediterranean Sea[6][8].

Sea urchin (Paracentrotus lividus)

It is hard for native species in the Mediterranean to get rid of C. taxifolia, because C. taxifolia has toxic secondary metabolites[9]. Toxic secondary metabolites are enzymes that are used as a defense mechanism when a potential predator or grazer, like a sea urchin, Paracentrotus lividus, tries to attack the algae[9]. From the presence of the toxic secondary metabolites, predators tend to stay away from C. taxifolia, as those that did prey upon the algae died a few months after[9]. A decline in sea urchin populations was detected in areas invaded by C. taxifolia[9]. The spread of C. taxifolia negatively impacted the Mediterranean Sea, decreasing its biodiversity and limiting native organisms by competing for light, food, and nutrients[1][6][7][9].

How does C. taxifolia impact marine ecosystems?

Rhizosphere structure

C. taxifolia are able to stimulate nitrogen fixation in low-nutrient environments [10]. This occurs when C. taxifolia releases photosynthetic products (Ex, organic carbon) into the rhizosphere [10]. The rhizosphere is the zone around the root where microorganisms and processes important for growth and plant health are located [11]. This process mimics the behaviour of saltwater vascular plants [10]. The excreted organic carbon then stimulates fermenting bacteria[10]. This fermenting bacterial community breaks down the organic carbon into substrates that support sulphate-reducing bacteria [10]. This creates low-oxygen (anaerobic) conditions [10]. C. taxifolia caused nitrogen fixation rates to increase in sediment below the algae by a factor of 28 [10]. This nitrogen fixation enhances the decomposition of native seagrass [10]. The nitrogen fixation also increases nutrient availability and improves sediment conditions for C. taxifolia [12]. This creates a positive feedback loop that allows for C. taxifolia to grow and spread quickly in poor sediment conditions, further enhancing the invasive species' success [10][12].

Measurable Ecosystem Changes to Primary Productivity:

The invasive Caulerpa taxifolia disrupts Mediterranean ecosystems through intense competition for light and space, notably reducing the summer productivity of sensitive species like Cystoseira barbata by 20-30%. By utilizing the allelopathic effects of Caulerpenyne, C. taxifolia suppresses competitors. This chemical interference is seasonally mediated, with toxicity peaking in summer, when caulerpenyne concentrations can reach ~9.6 µg/mL. Caulerpenyne also deters generalist herbivores such as sea urchins, reducing grazing pressure and allowing C. taxifolia to persist. Thus, primary productivity becomes increasingly concentrated within the invasive species.[13]

Furthermore, exposure to invasive algal toxins forces native species into defensive physiological responses. For instance, the seagrass Posidonia oceanica exhibits a stress response when exposed to waterborne Caulerpa taxifolia toxins. To mitigate toxic exposure, the seagrass reallocates metabolic energy away from primary growth toward the synthesis of protective phenolic compounds and tannins. While these antioxidants protect tissues, the diversion of resources leads to diminished health and stunted growth. Over time, the chronic stress weakens the seagrass beds, reducing habitat complexity and negatively impacting the diverse communities that rely on these meadows for survival. [13]

Measurable Ecosystem Changes to Trophic strcuture:

The invasion of Caulerpa taxifolia triggers measurable shifts in ecosystem connectivity, fundamentally altering the base of the benthic food web. This disruption is best reflected in the declining density of sediment-dwelling meiofauna, which play a vital role in nutrient cycling and energy transfer. In intact Posidonia oceanica meadows, meiofaunal densities are reported to be up to ten times higher than in adjacent areas dominated by C. taxifolia.  This shift is driven in part by the algae’s dense growth, which traps organic matter and lowers oxygen availability, creating inhospitable conditions for infaunal communities.[14]

In addition, the release of toxic compounds (such as caulerpenyne) selectively alters microbial assemblages by suppressing Gram-positive bacteria while promoting Gram-negative strains[13]. Thus, it effectively acts as a biological filter that reshapes microbial composition. These changes disrupt natural nutrient cycling processes and reinforce the persistence of C. taxifolia, further degrading ecosystem structure. As a result, energy pathways become simplified and less efficient, weakening connections between trophic levels. This is reflected at larger scales through habitat homogenization and a documented 25–50% reduction in native algal diversity in invaded areas[14].

Unique characteristics of this habitat that make it vulnerable to C.taxifolia invasion

Marine Litter and Semi Enclosure

Enclosure of the Mediterranean Sea

Marine litter can be defined as “any persistent, manufactured or processed solid material discarded, disposed of or abandoned in the marine and coastal environment” [15]. The Mediterranean Sea is a semi-enclosed area with anti-estuarine circulation; these characteristics cause the Mediterranean Sea to trap and retain materials that enter it. The Mediterranean Sea’s coastlines house over 150 million people, it handles 15-30% of global shipping, and the countries that surround it experience approximately one third of global tourism each year. These human activities introduce massive amounts of marine litter into the Mediterranean Sea. During peak summer, humans leave an estimated 40.6 million litter items per day across all Mediterranean island beaches [15]. The vast majority (94%) of this litter is plastic-based, meaning it persists in the ecosystem for a very long time. Marine litter pollution places significant stress on the Mediterranean Sea coastal ecosystem as organisms suffer entanglement, toxicity of water, and endocrine disruption. This stress reduces biodiversity and  makes the ecosystem more susceptible to establishment and spreading of invasive species such as C. taxifolia [16].

Marine traffic, boating, and anchoring

As mentioned in the previous section, the Mediterranean Sea handles 15-30% of global shipping traffic [15], making marine vessel traffic one of the most significant sources and drivers of species invasion. Vessels facilitate the spread of invasive species through multiple mechanisms including: hull biofouling, ballast water discharge, and fragmentation of algae beds by boat anchoring.

Commercial and recreational marine vessel hulls can act as a substrate for attachment and transport of marine organisms, such as C. taxifolia. This accumulation of marine organisms on boat hulls is known as “hull biofouling” [17]. Marchini et al. (2019) surveyed more than 600 recreational marine vessels across 25 marinas in the Mediterranean Sea. They found that 71% of these vessels harboured non-indigenous species, and over 50% of the vessels harboured non-indigenous species that did not yet exist in the Mediterranean Sea. The high volume of marine traffic through the Mediterranean Sea makes it vulnerable to invasive species introduction from hull biofouling.

Ballast water uptake and discharge is an one of the biggest sources of invasive species introduction and spreading. Large shipping vessels often take up large amounts of water from a port to stabilize the ship during travel, and discharge it in another location; this water can contain spores of algae, such as C. taxifolia[18] . Flagella & Abdulla (2005) estimate that 3000-4000 species have been transported across the globe by commercial vessels, and 30% of these species have been brought to the Mediterranean Sea.

On a local scale, anchoring of marine vessels into C. taxifolia beds is a major driver of C.taxifolia spread and colonization within the Mediterranean Sea ecosystem. West et al. (2007)  studied C.taxifolia fragmentation by different anchor and anchor attachment types (rope vs. chain). They found that 82% of anchors lowered into C.taxifolia beds removed fragments, and that chain anchor attachments produced larger fragments compared to rope attachments. The size of fragments is a relevant consideration because larger fragments have a higher fragment survivorship rate[14]. Additionally, they found that shade and moisture provided by anchored boats and damp ropes enhances fragment survivorship by preventing desiccation.

Anthropogenic Impacts on the C. taxifolia invasion

Ocean Acidification

Ocean acidification driven by increasing levels of atmospheric CO₂ dissolving into the ocean is a poses a major threat to many marine ecosystems and organisms; however, C.taxifolia may uniquely benefit from acidification. Roth-schulze et al. (2018) examined the effect of pH, pCO2, and temperature on samples populations of native and invasive of C.taxifolia in controlled laboratory tanks. They created different combinations of these variables, and observed the growth rate, mortality, and photosynthetic performance of C. taxifolia, as well as the diversity and composition of its ectosymbiotic bacteria. They found that low pH, high CO2 conditions conditions enhanced the growth rate of C. taxifolia, and had no impact on its photosynthetic performance or bacterial symbionts [14]. This evidence is significant because it demonstrates that C. taxifolia not only tolerates ocean acidification conditions, but actually thrives under them. This unique characteristic of C. taxifolia gives it a major competitive advantage over other macroalgae species in the Mediterranean Sea as oceans become increasingly more acidic.

Ocean Warming

The Mediterranean Sea is subject to significant warming driven by global climate change. Rivetti et al. (2014) analyzed temperature profiles for the Mediterranean Sea from 1945 to 2011, and discovered widespread temperature increases of approximately 0.07°C per year in a large portion of the basin, and more localized short-term temperature increases of up to 5.2°C above the average. Additionally, they demonstrate that these warming trends are associated with mass mortality events among benthic invertebrates (i.e. cnidarians, sponges, bivalves). Such mortality events result in biodiversity loss, which reduces the ecosystem’s resilience to invasive species establishment[14]. Roth-Schulze et al.’s (2018) study described in the previous section on ocean acidification found that increased temperature generally enhanced the growth rate of C. taxifolia. This suggests that C. taxifolia has a competitive advantage over other macroalgae species in the continuously-warming Mediterranean Sea.

What organisms does it impact?

P. oceanica in the Mediterranean Sea

Native Seagrass

C. taxifolia and P. oceanica, a native seagrass to the Mediterranean, directly compete for both space and light [19]. C. taxifolia produces a phytotoxin called caulerpenyne [20]. This toxin can severely damage P. oceanica by inhibiting the growth of adult leaves and preventing new leaf formations [20]. The toxin can also damage growth tissues called meristems, which threaten the seagrass’s survival [20]. It disrupts P. oceanica’s ability to photosynthesize and produce energy by reducing key proteins [20]. Caulerpeyne triggers P. oceanica’s stress responses [20]. These responses include oxidative stress, which occurs when the seagrass produces excess reactive oxygen species (ROS) [20]. When under stress, ROS accumulate, causing cellular damage [20]. C. taxifolia also produces caulerpin, a secondary metabolite [20]. This metabolite had little to no effect on leaf growth or formation but still caused metabolic and cellular changes, including impacts on transport and photosynthesis [20]. It also induces a defensive response in the seagrass, but to a lesser degree than caulerpeyne [20]. As competition between the two organisms increases, seagrass leaf length and age decrease and the leaves show more damage [21].

Sabella spallanzanii, a species of marine polychaete worms

Macrofauna

C.taxifolia provide a significant amount of detritus [22]. This detritus negatively affects invertebrates, as it lowers the abundance and reduces biodiversity [22]. Caulerpenyne, a phototoxin mentioned above, can be toxic or stressful for many invertebrates [20] [22]. Since invertebrates are unable to consume C. taxifolia, many species die or move to different areas with a more abundant food source [22]. As seen above, C. taxifolia stimulate nitrogen fixation [10]. This nitrogen fixation, causes anaerobic sediment condition [10]. These low oxygen conditions make it hard for organisms to survive [20][22]. C. taxifolia also change the habitat. Accumulated detritus can physically alter the sediment by making it more compact or unstable [22]. Macrofauna rely on seagrass like P. oceanica for shelter, so when P. oceanica is outcompeted by the algae, macrofauna lose their habitat and are more vulnerable to predation [23].

Native Fish

The spread of Caulerpa taxifolia directly alters fish communities by restructuring benthic habitats in ways that reduce feeding efficiency and increase energetic stress. Its dense mats of stolons and rhizomes form tightly interwoven networks that physically limit access to prey, particularly for benthic foragers. As a result, fish must adjust their behaviour in response to this constrained environment. [24]

Mullus surmuletus

Studies on the striped red mullet (Mullus surmuletus) provide clear evidence of this shift, showing that individuals in C. taxifolia habitats travel shorter distances, make more frequent feeding attempts, and spend significantly more time searching for food compared to those in Posidonia oceanica seagrass meadows. Despite this increased effort, feeding success is reduced, indicating a decline in overall foraging efficiency. This is further reflected in population structure, where large individuals comprise only about 1.19% of fish in C. taxifolia beds, compared to approximately 27.8% in seagrass habitats, suggesting that limited resource access constrains growth and survival. As prey becomes less accessible, intraspecific competition intensifies, forcing individuals to expend more energy for diminishing returns. These changes not only impact individual fitness, disrupt broader energy transfer, and contributing to a simplified and less productive food web.[24]

Scorpaena porcus

A second study conducted in France further highlights how Caulerpa taxifolia can impact fish at a physiological level. Research on the Mediterranean scorpionfish (Scorpaena porcus) found that exposure to the alga alters the fish’s Cytochrome P450 system, a group of liver enzymes responsible for detoxification and hormone regulation. This disruption was evident in the way the fish processed progesterone, where one key metabolite decreased by 25% while another increased significantly, indicating a substantial shift in internal biochemical pathways. Notably, these effects occurred even when fish did not ingest the alga, as simply inhabiting the same water was sufficient to trigger changes. [25]

This demonstrates that C. taxifolia exerts influence through waterborne chemical exposure, extending its impact beyond direct trophic interactions. As a result, impaired enzyme function may reduce the fish’s ability to properly break down toxins, increasing the likelihood of toxin accumulation within tissues. This not only compromises individual health but also raises broader ecological and human concerns, as these accumulated toxins may transfer through the food web and potentially affect higher-level consumers (such as H. sapiens). [25]

Are their unique characteristics of C. taxifolia that make it vulnerable?

Even though C. taxifolia has shown that it can outcompete and thrive in the Mediterranean, there are characteristics that make it vulnerable. An example of this is that C. taxifolia reproduce asexually [26]. The main mode of dispersion that C. taxifolia use is vegetative dispersion [26]. A key mechanism to this is fragmentation [26]. Fragmentation is the process where the algae splits into parts and reattaches to new substrates, growing into clone of the original algae [26][27]. This process can occur due to factors such as wave action, anchors, or other disturbances [26]. C. taxifolia release their gametes synchronously [26]. This release occurs in short coordinated windows, typically in the early morning [26]. However, these releases are highly dependent light conditions [26]. If there is low light intensity, the release time will be prolonged [26]. It was also found that during these releases, only male gametes were produced [26]. This eliminates any chance of sexual reproduction [26]. Since all gametes are identical to each other, there is no genetic diversity [26]. Even though C. taxifolia are able to thrive in the current conditions, if the conditions change unfavourably, they are unable to adapt [26].

Where are we today?

The spatial extent of C. taxifolia peaked around 1994, then steadily declined from the early 2000s onward, and now remains at relatively low densities. However, this decline does not indicate recovery. At its peak, C. taxifolia covered thousands of hectares, fundamentally reshaping habitats. These changes disrupted food availability and ecosystem structure, and their effects continue even as the algae itself has receded. As a result, the system has not fully recovered and may, in fact, be more vulnerable to further invasions. This aligns with the invasion meltdown hypothesis, which suggests that one invasive species can facilitate the success of others.

Ineffective and effective solutions

Fortunately, since 2006, the killer algae C. taxifolia has been successfully eradicated with minimal coverage at all spread locations[5]. After years of efficient eradication methods and natural degradation, the invasive algae is no longer an ecological issue [28]. The boom and bust phenomena in the Mediterranean and global solutions involving legal, chemical, and physical actions effectively combated the invasion[29]

Area covered by reappearance of C. taxifolia at 4 sites in Malinska. E represents the time of eradication.

Local Solutions

After recognition in Malinska, a coastal town in Croatia, suction pumps were tested at 4 different sites in the Mediterranean in 1996 and 1997. However, eradication was only successful if the patch was small. All 4 sites needed re-pumping shortly after due to the algae’s quick and aggressive reappearance. The reappearance at 4 different sites displayed different colonization patterns and area covered, as supported by the bar graphs [28].

After 2001, all four graphs show C. taxifolia covers were not found at any sites if not sparsely distributed. The spontaneous decline of the algae after repopulation supports the assumption that C. taxifolia experienced a natural solution of the boom and bust phenomena in the Mediterranean, especially when water temperatures were cooler during Winter months, which made them more susceptible to death.

It is important to note that these conflicting patterns of recolonization cannot confidently conclude the ecological cause of regression of C. taxifolia at these sites. Thus, although the local solution was not as effective as expected, the possibility of boom and bust naturally eradicated the algae along the coast of Croatia.

In other areas of across the Mediterranean, eradication methods like biocontrol and dredging were largely unsuccessful due to the algae's uncontrolled spread. Efforts have only mitigated algae growth but did not completely eradicate coverage. Today, there is still a reduced amount of coverage in areas of the Mediterranean including the coast of Italy, France, and Spain[30].

Global Solutions

Legal

At the first infestations of C. taxifolia in the waters surrounding Europe, Australia, and eventually the US, many countries took immediate legal action in managing and mitigating the spread of the algae. (Schaffelke). After the algae spread to the waters in Southern California, the U.S. government listed C. taxifolia on the U.S. Federal Noxious Weed List in 1999. Due to the 15 year history of the killer algae dispersal starting from the Mediterranean, the aquarium strain of C. taxifolia was banned in sales and possession[31]. The US Animal and Plant Health Inspection Service maintains a list of federal noxious weeds including aquatic, parasitic, and terrestrial plants that are prohibited because of their damage to native ecosystems, agriculture, and trade. Other countries like Spain, France, and Australia implemented the same policies of prohibited sale, possession, and transport of C. taxifolia[32].

Chemical

List of globally tested eradication methods. Red boxes are Southern California and Australia eradication methods that saw complete success.

Since the spread of C. taxifolia dominating Southern California, there have been numerous scientifically planned attempts to eradicate the algae. Chemical methods were worked on small infestations first with minimal ecological impacts[29]. Once success has been achieved, they can then carry out these procedures to eradicate C. taxifolia in a large scale way. In Southern California, covering and sealing of Polyvinylchloride tarps and injecting liquid chlorine at a 12% stock solution underneath the tarp has seen complete success[29]. To be considered “eradicated” no trace of C. taxifolia should be present at the initial site; however this is complicated to confirm due to water turbidity and epiphytism[29]. In California, after 2 years of monitoring closely with no observed C. taxifolia, it was successfully eradicated in July 2006.


The invasive algae’s spread in Southern Australia raised immediate concern because of its great ability to outcompete native species. A chemical method that worked for Australia was applying coarse sea salt of 99.5% NaCl specifically in the cooler months. The most successful dosage was 50kg/m^2[33]. The salt killed the algae within a few hours via osmotic shock and cell lysis while unharming native flora and fauna[33]. Taking advantage of C. taxifolia’s extreme sensitivity to salt conditions effectively decreased their population while native species like Z. capricorni in South Australia can adapt and recover to the salinity change[33].

Spreading Awareness

Since algae can rapidly transcend into national waters, the problem is considered global, requiring global action. For efficiency of reducing this spread, rapid response plans along with immediate funding must be adequate in coastal regions around the world. If marine invasions could be treated to the same extent as natural disasters like tornados, fires, floods, then rapid response should be achievable. To deal with future invasions, researchers suggested preparing biological experts, ownership of waterways, past eradication strategies, and funding[34].

References

  1. ↑ 1.00 1.01 1.02 1.03 1.04 1.05 1.06 1.07 1.08 1.09 Galil, B.S. (12 January 2007). "Loss or gain? Invasive aliens and biodiversity in the Mediterranean Sea". Marine Pollution Bulletin. 55: 314–322 – via Elsevier Science Direct.
  2. ↑ 2.0 2.1 Meinesz, Alexandre; Belsher, Thomas; Thibaut, Thierry; Antolic, Boris; Mustapha, Karim Ben; Boudouresque, Charles--Francçois; Chiaverini, Danièle; Cinelli, Francesco; et al. (June 2001). "The Introduced Green Alga Caulerpa Taxifolia Continues to Spread in the Mediterranean". Biological Invasions. 3: 201–210 – via Springer Nature Link. Explicit use of et al. in: |last9= (help)
  3. ↑ 3.0 3.1 Siguan, Maria A. R. (2002). "Review of Non-Native Marine Plants in the Mediterranean Sea". Invasive Aquatic Species of Europe. Distribution, Impacts and Management. 1: 291–310 – via Springer Nature Link.
  4. ↑ 4.0 4.1 de Blok, J.W. (September 1975). "The Texel aquarium". Netherlands Journal of Sea Research. 9: 231–232 – via Elsevier.
  5. ↑ 5.0 5.1 5.2 5.3 Montefalcone, Monica; Morri, Carla; Parravicini, Valeriano; Bianchi, Carlo Nike (26 May 2015). "A tale of two invaders: divergent spreading kinetics of the alien green algae Caulerpa taxifolia and Caulerpa cylindracea". Biological Invasions. 17: 2717–2728 – via Springer Nature Link.
  6. ↑ 6.0 6.1 6.2 6.3 6.4 6.5 6.6 Windarto, Seto; Wijaya, Yusuf Jati; Putri, Diwyacitta Antya; Umam, Ahmad Khoirul (September and October 2025). "Green Gold of the Ocean: Unlocking the Potential of Caulerpa in Global Seaweed Markets – A Review". Egyptian Journal of Aquatic Biology and Fisheries. 29(5): 2673–2700 – via Egyptian Journal of Aquatic Biology and Fisheries. Check date values in: |date= (help)
  7. ↑ 7.0 7.1 Thomaz, Sidinei Magela (30 July 2025). "Asexual reproduction of aquatic macrophytes via stem fragments: A review on determinants of plant fragmentation and propagule dispersal". Hydrobiologia – via Springer Nature Link.
  8. ↑ 8.0 8.1 Mannino, Anna Maria; Balistreri, Paolo (13 November 2017). "An updated overview of invasive Caulerpa taxa in Sicily and circumSicilian Islands, strategic zones within the NW Mediterranean Sea". Flora Mediterranea. 27: 221–240 – via Flora Mediterranea.
  9. ↑ 9.0 9.1 9.2 9.3 9.4 Bourdouresque, Charles F.; Meinesz, Alexandre; Ribera, María A.; Ballesteros, Enric (1995). "Spread of the green alga Caulerpa taxifolia (Caulerpales, Chlorophyta) in the Mediterranean: possible consequences of a major ecological event". Scientia Marina. 59: 21–29 – via Digital CSIC.
  10. ↑ 10.00 10.01 10.02 10.03 10.04 10.05 10.06 10.07 10.08 10.09 10.10 Chisholm, J. R. M; Moulin, P. (2003). "Stimulation of Nitrogen Fixation in Refractory Organic Sediments by Caulerpa taxifolia (Chlorophyta)". Limnology and Oceanography.
  11. ↑ Bakker, Peter A. H. M.; Berendsen, Roeland; Doornbos, Rogier F.; Wintermans, Paul C. A; Pieterse, Corné M (2013). "The rhizosphere revisited: root microbiomics". Frontiers in Plant Science.
  12. ↑ 12.0 12.1 Renn,, Chloe; Vadillo Gonzalez, Sebastian; Marzinelli, Ezequiel M.; Dafforn, Katherine A. (2025). "Propagule pressure and native macrophyte biomass mediate the success of an invasive alga: the role of below-ground microbial communities". Biological Invasions.CS1 maint: extra punctuation (link)
  13. ↑ 13.0 13.1 13.2 Boudouresque, C. F., & Verlaque, M. (1997). Ecology of Caulerpa taxifolia in the Mediterranean Sea. Marine Ecology Progress Series, 149, 279–289. https://www.int-res.com/articles/meps/149/m149p279.pdf
  14. ↑ 14.0 14.1 14.2 14.3 14.4 Cvitkovic, I., Despalatovic, M., Uljevic, A., Matijevic, S., Bogner, D., Lusic, J., & Travizi, A. (2017). Structure of epibiontic and sediment meiofauna in the area invaded by invasive alga caulerpa taxifolia. Marine Biology, 164(1), 1-15. https://doi.org/10.1007/s00227-016-3034-4
  15. ↑ 15.0 15.1 15.2 Grelaud, M; Ziveri, P (2020). "The generation of marine litter in Mediterranean Island beaches as an effect of tourism and its mitigation". Scientific Reports. 10(1): 20326–11.
  16. ↑ Deudero, S; Alomar, C (2015). "Mediterranean marine biodiversity under threat: Reviewing influence of marine litter on species". Marine Pollution Bulletin. 98(1-2): 58–68.
  17. ↑ Marchini, A; Seebens, H; Marina, B; Occhipinti-Ambrogi, A; Forcada, A; Ulman, A; Ferrario, J (2019). "Alien species spreading via biofouling on recreational vessels in the Mediterranean Sea". The Journal of Applied Ecology. 56(12): 2620–2629.
  18. ↑ Flagella, M. M.; Abdulla, A. A. (2005). "Ship ballast water as a main vector of marine introductions in the Mediterranean Sea". WMU Journal of Maritime Affairs. 4(1): 95–104.
  19. ↑ Ceccherelli, Giulia; Cinelli, Francesco (1999). "Effects of Posidonia oceanica canopy on Caulerpa taxifolia size in a north-western Mediterranean bay". Journal of Experimental Marine Biology and Ecology.
  20. ↑ 20.00 20.01 20.02 20.03 20.04 20.05 20.06 20.07 20.08 20.09 20.10 20.11 Oliva, Daniela; Piro, Amalia; Carbone, Marianna; Mollo, Ernesto (2024). "Physiological and proteomic responses of Posidonia oceanica to phytotoxins of invasive Caulerpa species". Environmental and Experimental Botany.
  21. ↑ Dumay, Olivier; Fernandez, Catherine; Pergent, Gérard (2002). "Primary production and vegetative cycle in Posidonia oceanica when in competition with the green algae Caulerpa taxifolia and Caulerpa racemosa". Marine Biological Association of the United Kingdom. Journal of the Marine Biological Association of the United Kingdom.
  22. ↑ 22.0 22.1 22.2 22.3 22.4 22.5 Taylor, Sl; Bishop, Mj; Kelaher, Bp; Glasb, Tm (2010). "Impacts of detritus from the invasive alga Caulerpa taxifolia on a soft sediment community". Marine Ecology Progress Series.
  23. ↑ Cvitkovic, Ivan; Despalatovic, Marija; Uljevic, Ante; Matijevic, Slavica; Bogner, Danijela (2017). "Structure of epibiontic and sediment meiofauna in the area invaded by invasive alga Caulerpa taxifolia". Marine Biology.
  24. ↑ 24.0 24.1 Longepierre, S., Robert, A., Levi, F., & Francour, P. (2005). How an invasive alga species (caulerpa taxifolia) induces changes in foraging strategies of the benthivorous fish mullus surmuletus in coastal mediterranean ecosystems. Biodiversity & Conservation, 14(2), 365-376. https://doi.org/10.1007/s10531-004-5356-7
  25. ↑ 25.0 25.1 Uchimura, M., Sandeaux, R., & Larroque, C. (1999). The enzymatic detoxifying system of a native mediterranean scorpio fish is affected by caulerpa taxifolia in its environment. Journal of Agricultural and Food Chemistry, 47(4), 1754-1759. https://pubs.acs.org/doi/10.1021/jf9808650
  26. ↑ 26.00 26.01 26.02 26.03 26.04 26.05 26.06 26.07 26.08 26.09 26.10 26.11 26.12 Žuljević, Ante; Antolić, Boris (2000). "Synchronous release of male gametes of Caulerpa taxifolia (Caulerpales, Chlorophyta) in the Mediterranean Sea". Phycologia.
  27. ↑ "Fragmentation - Definition and Examples - Biology Online Dictionary". Biology Articles, Tutorials & Dictionary Online. 2026.
  28. ↑ 28.0 28.1 Ivesa, Jeklin (2006). "Vegetation patterns and spontaneous regression of Caulerpa taxifolia". pp. 324–330.
  29. ↑ 29.0 29.1 29.2 29.3 Walters, L. (2009). Ecology and management of the invasive marine macroalga caulerpa taxifolia. Management of invasive weeds, 287-318. https://doi.org/10.1007/978-1-4020-9202-2_15
  30. ↑ R.G., Creese (June 2004). "Control Techniques for Caulerpa taxifolia" (PDF). NSW Fisheries. 64: 59–76.
  31. ↑ United States Department of Agriculture (2007). Federal Noxious Weed List. December 10, 2010. https://www.aphis.usda.gov/sites/default/files/weedlist.pdf
  32. ↑ Schaffelke, Britta. Capi Digital Library. (2008, March). Caulepra taxifolia (killer algae). [Datasheet]. CAPI Compendium. https://doi.org/10.1079/cabicompendium.2929
  33. ↑ 33.0 33.1 33.2 Glasby, T. M., Creese, R. G., & Gibson, P. T. (2005). Experimental use of salt to control the invasive marine alga Caulerpa taxifolia in New South Wales, Australia. Biological Conservation, 122(4), 573-580. https://doi.org/10.1016/j.biocon.2004.09.012
  34. ↑ Anderson, L. W. J. (2005). California's reaction to Caulerpa taxifolia: A model for invasive species rapid response. Biological Invasions, 7(6), 1003-1016. https://doi.org/10.1007/s10530-004-3123-z
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