Botanica Marina 2021; aop

Short Communication

João Faria*, Afonso C.L. Prestes, Ignacio Moreu, Gustavo M. Martins, Ana I. Neto
and Eva Cacabelos

Arrival and proliferation of the invasive seaweed

Rugulopteryx okamurae in NE Atlantic islands
https://doi.org/10.1515/bot-2021-0060 Keywords: Azores; macroalgae; marine bioinvasions; non-
Received June 15, 2021; accepted September 2, 2021; indigenous species; oceanic islands.
published online November 4, 2021

The brown seaweed Rugulopteryx okamurae (E.Y. Dawson)

Abstract: The present study reports the recent occurrence
I.K. Hwang, W.J. Lee et H.S. Kim, which has been introduced
and expansion of Rugulopteryx okamurae in the Azores ar-
in the Mediterranean (Verlaque et al. 2009) is a native and
chipelago (NE Atlantic). Morphological and molecular
widely distributed species in the subtropical to temperate
characters confirmed the species identification. Quick sur-
western Pacific Ocean (Lee and Kang 1986; Silva et al. 1987;
veys around the island of São Miguel showed that it has
Yoshida 1998). It grows in the shallow subtidal and can be
successfully colonized the island and is quickly expanding.
In some locations, R. okamurae is currently the dominant present year-round as dormant rhizoidal bases. At macro-
organism smothering all other benthic biota and posing a scopic level, individuals of R. okamurae are characterized by
serious threat to the benthic ecosystems across the region. a membranous and erect dichotomously branched thallus
The species first record dates from 2019 near the main with smooth margins 10–20 cm high, attached by rhizoids
harbour of the island, suggesting that its introduction was restricted to the basal parts of the thallus. The fronds grow
driven by human-assisted transport, via boat ballast waters into a compressed flabellum and in dense groups, and
or adhered to ship hulls and likely originating from the intraspecific variation is known to occur (Hwang et al. 2009;
Mediterranean populations that have been proliferating in Sun et al. 2006). Microscopically, the thallus consists of a
recent years across the Strait of Gibraltar. medulla, one cell thick centrally and two to three cells thick
near the margins (Womersley 1987; De Clerck et al. 2006;
Hwang et al. 2009). The reproductive structures are evenly
distributed over the entire thallus surface, rather than being
Ana I. Neto unfortunately deceased prior to the publication of this
confined to the concavities as in the other Rugulopteryx
article.
species. Just like other Rugulopteryx species, the sporangia
*Corresponding author: João Faria, Department of Biology, Faculty of of R. okamurae are borne on two stalk cells. The sporangia
Sciences and Technology, cE3c - Centre for Ecology, Evolution and can readily form propagules, proliferous branchlets arising
Environmental Changes/Azorean Biodiversity Group; University of on the thallus surface which grow into new plants of the
Azores, Ponta Delgada 9501-801, São Miguel, Azores, Portugal,
same ploidy level. Although it may be confused with some
E-mail: joao.fo.santos@uac.pt. https://orcid.org/0000-0002-5797-
0505 species of the genus Dictyota, R. okamurae is a single taxo-
Afonso C.L. Prestes, Ignacio Moreu and Ana I. Neto, Department of nomic entity and has been described in Hwang et al. (2009).
Biology, Faculty of Sciences and Technology, cE3c - Centre for Ecology, In European waters, the non-indigenous R. okamurae
Evolution and Environmental Changes/Azorean Biodiversity Group; was first detected in spring 2002 close to a small harbour in
University of Azores, Ponta Delgada 9501-801, São Miguel, Azores,
Thau Lagoon (France), a known hot spot of marine species
Portugal
Gustavo M. Martins, CIBIO - Research Centre in Biodiversity and introductions in Europe (Verlaque et al. 2009). It was more
Genetic Resources, InBIO Associate Laboratory, Pólo dos Açores – recently detected in Ceuta and in Andalusian waters,
Departamento de Biologia da Universidade dos Açores, Ponta southern Iberian Peninsula, in 2015 and 2016, respectively
Delgada 9501-801, Portugal (Altamirano et al. 2016, 2017; Ocaña et al. 2016). In the Strait
Eva Cacabelos, MARE - Marine and Environmental Sciences Centre,
of Gibraltar, R. okamurae has expanded massively in recent
Agência Regional para o Desenvolvimento da Investigação Tecnologia
e Inovação (ARDITI), Edifício Madeira Tecnopolo, Piso 0, Caminho da
years across subtidal marine hard-bottoms, causing evident
Penteada, Funchal 9020-105, Madeira, Portugal. ecological impacts on coastal communities (García-Gómez
https://orcid.org/0000-0002-8543-6510 et al. 2018, 2020). For instance, after its first record in the
2 J. Faria et al.: Rugulopteryx okamurae in Azores

area, R. okamurae became the most abundant species in liquid specimens at the AZB Herbarium Ruy Telles Palhinha
only one year (>90% coverage at 10–20 m depth), followed of the University of the Azores. Liquid specimens were
by a significant change in the structure of benthic commu- collected using 5% buffered formaldehyde seawater and
nities (García-Gómez et al. 2020; Sempere-Valverde et al. then replacing it by the fixing agent Kew (Bridsen and For-
2021). Additionally, hundreds of tons of R. okamurae man 2010). Pressed specimens were carried out as described
biomass have been reported to accumulate at coastal bed- by Gayral (1986).
rocks and/or beaches across the region with serious impli- Collected specimens were identified by their morpho-
cations for tourism and public health (Ocaña et al. 2016, logical and anatomical characteristics using both binocular
García-Gómez et al. 2018, 2020, 2021). Fishermen have also and light microscopes (see Hwang et al. 2009; Verlaque et al.
reported fishing nets becoming clogged with R. okamurae 2009). Macroscopically, individuals attained 8–20 cm in
and a significant reduction in their ability to catch fish height and were 95–130 mm thick (e.g. Figure 2A). Many
(Sempere-Valverde et al. 2019). displayed numerous proliferous branchlets on both surfaces.
The present study reports the recent occurrence of Transverse sections revealed the occurrence of mature
Rugulopteryx okamurae in the Azores archipelago (NE sporangia subtended by two stalk cells in most individuals, a
Atlantic). The species was first observed in early-2019 on the key distinctive morphological feature of the genus Rugu-
south coast of São Miguel Island and registered as an un- lopteryx (Figure 2B and C). Only asexual monosporangia
identified species of Dictyotales. A search for the species was were observed, 90–140 µm in diameter. Gametophytes were
conducted throughout the island in 2020 (June–July) and not encountered in the field. As in Sun et al. (2006), the upper
again in 2021 (May). Sampling was performed on rocky part of the thalli exhibited a unilayered medulla, which
bottoms at several locations by snorkelling and/or scuba sometimes become multilayered at the margins of thallus
diving at about 5 m depth. The most abundant organisms at due to transverse division of the internal cells (Figure 2D)
this depth are mainly macroalgae, which include articulated (see Hwang et al. 2009). A medulla with more than two cell
corallines (e.g. Ellisolandia elongata (J. Ellis et Solander) K.R. layers was often found at the lowest part of the blades.
Hind), encrusting corallines (e.g. Lithophyllum sp.), coarsely Moreover, DNA sequences of two chloroplast protein-
branched algae (e.g. Halopteris scoparia (Linnaeus) Sauva- encoded genes rbcL (ribulose bisphosphate carboxylase
geau, Asparagopsis taxiformis (Delile) Trevisan) and corti- large chain) and psbA (photosystem II reaction center
cated foliose algae (e.g. Zonaria tournefortii (J.V. Lamouroux) protein D1) from six collected samples were confirmed as
Montagne, Dictyota spp.) (Martins et al. 2013; Wallenstein R. okamurae using the GenBank BLASTn search. Resulting
et al. 2008). Invertebrates, although common, rarely occur at sequences matched with R. okamurae individuals from
high densities. Conspicuous species include sea-urchins the western Pacific Ocean (rbcL: GenBank accession no
(e.g. Arbacia lixula and Sphaerechinus granularis), star-fish MZ393479 to MZ393484; psbA: GenBank accession no
(e.g. Marthasterias glacialis, Ophidiaster ophidianus), poly- MZ393485 to MZ393490). DNA sequences for rbcL and
chaetes (e.g. Sabella spallanzani, Hermodice carunculata), psbA were amplified using primers listed in Draisma et al.
decapods (e.g. Calcinus tubularis, Percnon gibbesi) and gas- (2001) and Saunders and Moore (2013), respectively (see
tropods (e.g. Stramonita haemastoma, Phorcus sauciatus) Supplementary Tables S1, S2 for methodological details).
(Baptista et al. 2021; Martins et al. 2005). The arrival of R. okamurae in the Azores represents its
At each location, the DAFOR scale, an acronym for first record in the NE Atlantic islands. Although it may have
Dominant, Abundant, Frequent, Occasional and Rare, was been initially misidentified as a Dictyota sp., a posteriori an-
applied to measure the relative abundance of R. okamurae. alyses of video data taken in April 2019 indicate that R. oka-
The DAFOR scale is a method of defining relative abundance murae may have been already present in the island since
of species during rapid surveys (Hawkins and Jones 1992). early that year (see Supplementary Video). This finding was
The following criteria were used to rate the occurrence of the recorded in a location a few hundred meters from the main
species in this study: dominant (>75%), abundant (50–75%), shipping port of São Miguel Island, which may indicate the
frequent (25–50%), occasional (10–25%) and rare (<10%) likely vector of introduction of the species in the region (see
(Figure 1A). At each location, an identical slow and steady below). In subsequent years, the species underwent a rapid
swim perpendicular to the coastline was adopted. Under- proliferation throughout the South coast of São Miguel
water visual surveys took around 10 min. Observations were (Figure 1 and Table S3) dominating subtidal vegetation on
carried out by the same observer and performed on a virtual both well illuminated and shaded rock surfaces. In some
corridor of 4–5 m width along the pre-selected depth (∼5 m). locations occupancy rates were close to 100%. Substantial
In both field campaigns, live plants were collected, taken to amounts of the seaweed have also been cast on the beach as
the laboratory for examination, and housed as pressed and wracks as observed during the first semester of 2021
 J. Faria et al.: Rugulopteryx okamurae in Azores 3

Figure 1: Map of São Miguel Island (A) and locations (numbers) sampled for the presence of Rugulopteryx okamurae from 2019 to 2021. Black
circles indicate the occurrence and relative abundance of the species (see Supplementary Table S3). Images to the right show R. okamurae in
its natural environment (B) and as wrack cast onto the beach (C and D).

(Figure 1C and D). Similar beach accumulations of this brown hulls). These are known to be two of the main introduction
alga were reported at another island in Azores (Faial Island) vectors of exotic species in the marine environment (Hewitt
in May 2021, but species identification requires further et al. 2009). Both routes could explain the introduction of
confirmation. R. okamurae in the region from Mediterranean populations
As mentioned before, and given the location where the that are proliferating across the Strait of Gibraltar, while
species was first detected around the area of the main port taking advantage of the archipelago location as a known
of São Miguel, the introduction of R. okamurae into the stop for transatlantic crossing. For instance, Rosas-Guer-
Azores is likely to have been driven by human-assisted rero et al. (2018) observed that adult specimens of R. oka-
transport, via ballast waters and/or fouling (inlays in boat murae had survival rates between 80 and 100% after being
4 J. Faria et al.: Rugulopteryx okamurae in Azores

Figure 2: Rugulopteryx okamurae from the

Azores: (A) habit of the thallus showing the
anchoring rhizoids and dichotomous
branching; (B and C) thallus surface with
asexual monosporangia that are borne on 2
stalk cells (1 and 2); (D) transverse section
of thallus showing the margin with two
layers of medullary cells.

grown in dark conditions (a key condition during transport has the potential to affect the ecological balance of the
in ballast waters) for three weeks, maintaining the same community, but also fundamental economic activities such
survival rates when they were restored to the light. Also, it as fishing and tourism. It can contribute to the loss of marine
has been shown that the species is capable of adhering to biodiversity and alteration of the structure of the commu-
surfaces of very diverse nature and composition (García- nities, causing the physical displacement of native species
Gómez et al. 2018) increasing its capacity to travel attached due to substrate occupation and preventing the fixation of
to boat hulls. larvae or propagules of other species. This, together with the
In São Miguel Island, R. okamurae is showing explosive amounts of biomass observed at invaded locations, poses a
growth and a surprising adaptation capacity. Its prolifera- significant ecological pressure on native ecosystems and
tion and high seabed occupancy rate in São Miguel may local economies. This new invader should be kept moni-
be explained by its great ability to propagate by vegetative tored so that its status and potential impact on Azorean
(i.e. propagules) and asexual mechanisms (i.e. proliferous native biota can be evaluated. Given the high level of pro-
branchlets arising on the thallus surface which grow into liferation of the species there is a need for a rapid scientific
new plants; Verlaque et al. 2009), as highlighted in the Strait response in providing valuable information so that appro-
of Gibraltar (Altamirano et al. 2016, 2017; García-Gómez et al. priate preventive measures and mitigation actions can be
2020). Also, anthropogenic disturbance (e.g. fisheries, applied by regional authorities to safeguard coastal marine
ocean warming) and previous biological invasions (e.g. habitats in Azores.
Asparagopsis armata; Martins et al. 2019) may have reduced
niche competition in the region and favoured the arrival and Acknowledgments: We thank the editor and reviewers for
spread of R. okamurae in such remote oceanic islands. their helpful comments on the manuscript. In loving
The species is known to cause serious impacts over memory of Ana Neto.
previously established benthic communities in the Strait Author contribution: The author has accepted respon-
of Gibraltar (García-Gómez et al. 2020, 2021; Sempere- sibility for the entire content of this submitted manuscript
Valverde et al. 2021). Rugulopteryx okamurae not only and approved submission.
 J. Faria et al.: Rugulopteryx okamurae in Azores 5

Research funding: Funding was provided from National Gayral, P. (1986). Connaitre et reconnaitre les algues marines.
Funds through FCT – Fundação para a Ciência e Tecnologia, Éditions Ouest-France, Rennes.
Hawkins, S.J., and Jones, H.D. (1992). Marine field course guide. 1.
within the projects UID/BIA/00329/2015–2019 and UID/BIA/
Rocky shores. marine conservation society. Immel Publishing
00329/2020–2023. ACLP was supported by PhD grant Ltd, London.
awarded by FRCT- Fundo Regional da Ciência e Tecnologia Hewitt, C.L., Gollasch, S., and Minchin, D. (2009). The vessel as a
(M3.1.a/F/083/2015). vector – bio-fouling, ballast water and sediments. In: Rilov, G.
Conflict of interest statement: The author declare no and Crooks, J.A. (Eds.). Biological invasions in marine
conflicts of interest regarding this article. ecosystems: ecological, management, and geographic
perspectives. Springer-Verlag Berlin, Germany, pp. 117–131.
https://doi.org/10.1007/978-3-540-79236-9_6.
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where he coordinates the experimental laboratory for cultivation of
Yoshida, T. (1998). Marine Algae of Japan. Uchida Rokakuho
marine macroalgae at off-shore structures. He was a post-doctoral
Publication, Tokyo.
researcher between 2010 and 2018. He has broad research interests
that include the community ecology of intertidal rocky shores, the
Supplementary Material: The online version of this article offers population and community level effects of harvesting, the importance
supplementary material (https://doi.org/10.1515/bot-2021-0060). of biodiversity for ecosystem processes, the impact of man-made
structures on coastal ecosystems and the effects of climate change on
species interactions.

Bionotes Ana I. Neto

Department of Biology, Faculty of Sciences and Technology, cE3c -
João Faria Centre for Ecology, Evolution and Environmental Changes/Azorean
Department of Biology, Faculty of Sciences and Technology, cE3c - Biodiversity Group; University of Azores, Ponta Delgada 9501-801, São
Centre for Ecology, Evolution and Environmental Changes/Azorean Miguel, Azores, Portugal
Biodiversity Group; University of Azores, Ponta Delgada 9501-801, São
Ana I. Neto passed away in May 2021. She was a professor at the
Miguel, Azores, Portugal
University of Azores and since 1988 investigated marine botany and
joao.fo.santos@uac.pt
coastal ecology, using macroalgae as model organisms and the
https://orcid.org/0000-0002-5797-0505
Azorean littoral communities as model systems. She was the lead
João Faria holds a PhD in biology from the University of Lisbon and has scientist of the Island Aquatic Ecology SubGroup of the Azorean
been accumulating considerable knowledge on coastal habitats. He Biodiversity Group, and she coordinated the Herbarium AZB Ruy Telles
has proven to be skillful in applying experimental ecology designs and Palhinha of DB/UAc.
molecular and genetic tools to better understand how biological
diversity is generated. His main research interests lie in the field of Eva Cacabelos
marine conservation and resilience of marine organisms to natural MARE - Marine and Environmental Sciences Centre, Agência Regional
and/or anthropogenic-derived disturbances (i.e. biological invasions, para o Desenvolvimento da Investigação Tecnologia e Inovação
fisheries, ocean warming and ocean acidification). (ARDITI), Edifício Madeira Tecnopolo, Piso 0, Caminho da Penteada,
Funchal 9020-105, Madeira, Portugal
Afonso C.L. Prestes https://orcid.org/0000-0002-8543-6510
Department of Biology, Faculty of Sciences and Technology, cE3c -
Eva Cacabelos is a post-doctoral researcher and has extensive
Centre for Ecology, Evolution and Environmental Changes/Azorean
experience in the analysis of factors affecting littoral communities.
Biodiversity Group; University of Azores, Ponta Delgada 9501-801, São
She has participated in projects investigating the effects of invasions,
Miguel, Azores, Portugal
climate change or anthropogenic perturbations in benthic
Afonso C.L. Prestes is a PhD student of biology of the 3CBIO in the communities, as well as related to valorization of marine resources.
University of the Azores, studying the impact of invasive algae in the She is well acquainted with the methods of investigating effects of
Azorean algal community. He has a research grant in the non-indigenous species in littoral systems.