Figure 9
Morphological variation of Caulerpa racemosa-peltata complex taxa discussed in the present study. (A) C. nummularia Harvey ex. J. Agardh (AD-A91369) from Heron Island, Great Barrier Reef, Australia, showing ramuli disk arising from margin of parent disk; (B) Syntype specimen of C. nummularia (TCD 00111032). (C) C. macrodisca Decaisne (L 0509359/FM965053) from Indonesia with characteristic upright assimilator morphology; (D) Copy of the lectotype illustration of C. macrodisca by Decaisne (1846 (1846-1864), pl. 1, ); (E) C. megadisca Belton & Gurgel sp. nov. (AD-A90107) from Lizard Island, Great Barrier Reef, Australia showing typical single peltate disk morphology. Scale bars: A = 0.5 mm; B = 25 mm; C, D, E = 20 mm.
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Although recent molecular studies have indicated the presence of a number of distinct species within the Caulerpa racemosa–peltata complex, due to the difficulties presented by high levels of phenotypic plasticity and the large number of synonyms, infra-specific taxa, and names of uncertain affinity, taxonomic proposals are yet to be made. In this...
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... 7 included small, delicate, strictly peltate specimens with both crenate and entire disk margins, from which secondary peltate ramuli arose ( of syntype specimens in TCD and NSW (see Fig. 9B), and previous accounts of Caulerpa species from the Pacific region (i.e., Cribb and Cribb 1985, South and Skelton 2003, Abbott and Huisman 2004, Skelton and South 2007. The morphology of specimens in this lineage was ...
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... 8 and 9 included specimens that were identified as C. racemosa var. macrodisca that could easily be morphologically distinguished from other lineages, although specimens in lineage 9 could at times resemble peltate specimens from lineage 6. Specimens in lineage 9 (Fig. 9C) closely resembled the lectotype illustration (Decaisne 1846-1864; pl. 1, fig. 1 as C. macrodisca; reproduced as our Fig. 9D), whereas specimens from lineage 8 had ramuli that were not arranged around an upright assimilator, but rather the assimilators consisted of a single peltate branch (Fig. 9E). No intermediates between the two ...
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... as C. racemosa var. macrodisca that could easily be morphologically distinguished from other lineages, although specimens in lineage 9 could at times resemble peltate specimens from lineage 6. Specimens in lineage 9 (Fig. 9C) closely resembled the lectotype illustration (Decaisne 1846-1864; pl. 1, fig. 1 as C. macrodisca; reproduced as our Fig. 9D), whereas specimens from lineage 8 had ramuli that were not arranged around an upright assimilator, but rather the assimilators consisted of a single peltate branch (Fig. 9E). No intermediates between the two forms were observed during the present ...
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... specimens from lineage 6. Specimens in lineage 9 (Fig. 9C) closely resembled the lectotype illustration (Decaisne 1846-1864; pl. 1, fig. 1 as C. macrodisca; reproduced as our Fig. 9D), whereas specimens from lineage 8 had ramuli that were not arranged around an upright assimilator, but rather the assimilators consisted of a single peltate branch (Fig. 9E). No intermediates between the two forms were observed during the present ...
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... Herb. Agardh 16809, 16811 (in part), in LD; BM; 287054, in NSW; DSC01032, in TCD. Note: as type material was never designated by J. Agardh, the specimen marked 77 Friendly Isl. C. nummularia in TCD (TCD 0011105) is designated here as the lectotype (reproduced in Fig. ...
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Knowledge of spatial patterns of genetic differentiation between populations is key to understanding processes in evolutionary history of biological species. Caulerpa is a genus of marine green algae, which has attracted much public attention, mainly because of the impacts of invasive species in the Mediterranean. However, very little is known abou...
Taxonomy and species richness estimates for the genus Caulerpahave proven to be challenging due to the difficulty of assessing morphological species limits. In the present study we evaluate the taxonomy of the genus, and assess species distributions in the Eastern Tropical Pacific (ETP), based on morphology and tufA sequence data. Molecular analyse...
Citations
... This complicates species identification by researchers who are not familiar with Caulerpa. For example, Zuldin et al. (2019) concluded that they found C. macrodisca based on tufA, but they were not aware that they also sampled C. megadisca sequences from GenBank (JN645149, JN645154, Belton et al. 2014). Their queried specimen had morphological features of both C. macrodisca (ramuli attached to assimilators) and C. megadisca (very large ramuli), but this clade remained internally unresolved in the phylogeny they presented, so it cannot be concluded with certainty that they found C. macrodisca. ...
... Moreover, the authors did not submit their newly generated tufA sequence to GenBank. Belton et al. (2014) proposed tufA reference sequences for nine Caulerpa species of the so-called C. racemosa-peltata complex (Sauvage et al. 2013) (Suppl . Table S1). ...
... These studies also recognized 'dark taxa', which are genetically distinct specimens that could not be linked to existing species or have not yet been formally described as new species (Page 2016). Belton et al. (2014) reported Caulerpa sp. 3 (JQ894932) and Caulerpa sp. 10 (JN645159), both having globose ramuli like in C. racemosa. Both unidentified species were only represented by a single specimen and, therefore, the authors refrained from attributing a name to them as they did for nine other clades in the C. racemosa-peltata complex. ...
The circumtropical, multinucleate, siphonous, green algal genus Caulerpa is the single genus in the family Caulerpaceae and exhibits a remarkably high morphological variability. Morphological plasticity, overlapping morphological species boundaries, and the occurrence of cryptic species, has led several scientists to advocate the use of DNA barcodes for reliable species identification. The chloroplast-encoded tufA gene has been the DNA barcode marker of choice for Caulerpa, in particular the 820 nucleotides central region. Molecular identification through DNA barcodes depends on the availability of a curated reference library with pre-defined species. However, currently there are many incorrect Caulerpa species names in the GenBank sequence data base, potentially causing error propagation. This complicates species identification by researchers who are not familiar with Caulerpa. The present study reviewed the published CaulerpatufA sequences, generated new tufA sequences for four species, and selected reference sequences for ¾ of the currently accepted Caulerpa species. Reference sequences were primarily selected based on geographic proximity to the type location. This resulted in a tufA sequence alignment of 89 Caulerpa taxa of which four remained unnamed. Caulerpa ambigua, C. verticillata, and C. lanuginosa represent cryptic species complexes of, respectively, nine, three, and two species. The tufA sequences of C. macra, C. manorensis, and C. veravalensis were compared for the first time and the small sequence difference suggests that these could represent a single variable species.
... This plasticity resulted in the placement of several species as synonyms and the creation of forms, varieties, or ecads to accommodate the different or intermediate forms of species (e.g., C. cupressoides comprises eight forms and nine varieties). This proliferation of taxonomic ranks has highlighted the need for DNA barcodes for reliable identification [2,9,10]. ...
... To complete our dataset at a broader phylogenetic scale, we also selected one reference sequence per accepted species clade from Genbank. Firstly, we have added holotype sequences (i.e., four sequences) and tuf A references sequences proposed by Belton et al. (2014Belton et al. ( , 2019; i.e., 35 sequences) (See Table S1). Then, the selection for unrepresented species among the sequences available from GenBank was as follows: selection of sequence from a sample collected from the type locality of the species, and if not, near the type locality of the species. ...
... We have tried as far as possible to integrate reliable sequences that can be considered as good references for the species in question. In particular, we relied on the reference sequences proposed by previous studies [9,10,30], but for the vast majority of Caulerpa species, these have not yet been proposed. It would therefore be necessary to start creating a database of reference sequences for all existing Caulerpa species, in order to eliminate doubtful sequences and select the most reliable and accurate ones for each species. ...
Caulerpa J.V. Lamouroux is a genus of green macroalgae belonging to the family Caulerpaceae in the order Bryopsidales. The genus comprises 104 currently accepted species, of which 51 have been recorded from Pacific Islands. Among these islands, French Polynesia is found in the middle of the South Pacific Ocean and includes five archipelagos (i.e., the Austral, Gambier, Marquesas, Society, and Tuamotu Islands) where seaweed inventories have reported a total of 16 Caulerpa species so far based on morphology. Here, based on a sampling covering the five archipelagos of French Polynesia, we attempt to (i) verify the taxonomy of Caulerpa species present in these regions based on phylogeny, (ii) describe in more detail the specific diversity between the five archipelagos, and (iii) provide a morphological identification tool for these species. We successfully obtained 134 new tufA sequences for phylogenetic analyses, which corresponded to 13 species. We propose to resurrect C. pickeringii Harvey & Bailey for representatives of C. webbiana var. pickeringii and classify it in the Caulerpa subgenus Araucarioideae. We also transfer C. seuratii to C. pickeringii based on genetic results. A new morphological identification key is provided as well as an updated distribution of Caulerpa species across French Polynesia.
... In Japan, several other species of Caulerpa, such as Caulerpa chemnitzia var. laetevirens (Montagne) Fern andez-García et Riosmena-Rodriguez (Belton et al. 2014;Fern andez-García et al. 2016;Norris et al. 2017; = Caulerpa racemosa var. laetevirens [Montagne] Weber Bosse) are also found, and these species are also distributed in the warm temperate regions of Japan (Yoshida 1998). ...
The effects of four stressors, temperature, irradiance, desiccation and salinity, on the photochemical efficiency (ΔF/F 0 m) of an edible green alga, Caulerpa chemnitzia var. laetevirens from Kagoshima, Japan, were determined for optimizing cultivation and shipping environments using a pulse amplitude modulation-chlorophyll fluorometer. The ΔF/F 0 m remained stable at 24-34 C during the 3-day temperature exposures ranging from 8 to 36 C; however, it dropped at higher and lower temperatures. During continuous exposure for 6 h to irradiance levels of 400 (low) and 1000 (high) μmol photons m À2 s À1 at 16, 24 and 32 C, the decrease in ΔF/ F 0 m was pronounced at high irradiance. Moreover, ΔF/F 0 m also dropped at 16 C even under low irradiance, suggesting the occurrence of low temperature-light stress. Desiccation experiments under 50% humidity and up to 5 h of aerial exposure at 24 C and dim light (20 μmol photons m À2 s À1) indicated that ΔF/F 0 m was well tolerated within 1 h of desiccation; however, it dropped quickly as the desiccation period prolonged, suggesting that, if more than 20% of the interior water was lost, ΔF/F 0 m dropped progressively. However, during a similar experiment up to 14 days of aerial exposure under saturated humidity (99%) and dim light (12:12 h light/dark photocycle), as the interior water of the frond was maintained because of the saturated humidity, ΔF/F 0 m appeared to be well maintained for up to 5 days. Similar to desiccation , this alga exhibited stenohaline characteristics (30-40 psu) under the salinity gradient of 0-80 psu for up to 7 days of culture at 24 C and under dim light (12:12 h light/dark). In conclusion, this alga can be cultivated by modifying the techniques used for cultivating Japanese Caulerpa lentillifera and adjusting its environment accordingly. To ensure appropriate shipping conditions, it is essential to maintain saturated humidity in a plastic container at room temperature.
... cylindracea (Sonder) Verlaque and Boudouresque [22]. Finally, molecular investigations stated the genetic independence of the taxa and established the reinstatement of the original binomial as Caulerpa cylindracea Sonder [23]. This Indo-Pacific and thermophilic macroalga is able to colonize every kind of substrate, from sandy to muddy and rocky bottoms [24] and even areas affected by sedimentation [25]. ...
Caulerpa cylindracea, Indo-Pacific and thermophilic macroalgal species, have spread in Italian coastal waters, especially in the western Mediterranean Sea. Also, along the Calabria coastline, this Invasive Alien Species (IAS) was able to colonize most of the western and eastern seasides of the region. This research was conducted between 1999 and 2021 through a detailed cost-effective citizen science program. The novelty of this research was to determine the overall distribution of Caulerpa cylindracea along Calabria seawaters from the Tyrrhenian to the Ionian coasts. The results of the monitoring survey highlighted the presence of Caulerpa cylindracea in 45 collecting stations throughout the Calabrian coastline. This regional assessment showed the extensive adaptability of the species to different environmental conditions: from exposed to sheltered sites, in shaded and sunny coastal areas and also throughout pristine and polluted waters. The rapid and increasing spread of the species could affect the structure of Mediterranean biota or, otherwise, could lead in time to new ecological niches. As highlighted in the most recent literature, it is necessary to update monitoring and mapping plans for the protection of marine biodiversity through close coordination between scientists, citizens and policy makers, all engaged to ensure effective management of invasive processes.
... Our analyses also showed that the M. haageana complex is a recently diversified lineage (2.1 mya; Cervantes et al., 2021) whose evolution and speciation are not well understood. Although traditional morphological and conventional molecular markers have traditionally been used for species delimitation in plants (Belton et al., 2014), animals (Schwarzfeld and Sperling, 2014) and fungi (Stielow et al., 2011), their usefulness decreases considerably when dealing with taxonomically challenging species complexes such as the M. haageana complex. In this context, the objective is to assess species boundaries using an integrative framework incorporating genomic data (GBS data analysis, phylogenetic network, and coalescentbased analyses), complex phenotypes (morphometric analysis) and environmental variables (ecological niche models). ...
Species complexes consist of very close phylogenetic relatives, where morphological similarities make it difficult to distinguish between them using traditional taxonomic methods. Here, we focused on the long-standing challenge of species delimitation in the Mammillaria haageana complex, a group that presents great morphological diversity that makes its taxonomy a puzzle. Our work integrates genomic, morphological, and ecological data to establish the taxonomic limits in the M. haageana complex, and we also studied the evolutionary relationships with the remainder of the M. ser. Supertextae species. Our genetic analyses, as well as morphological and ecological evidence, led us to propose that the M. haageana complex is made up of six distinct entities (M. acultzingensis, M. conspicua, M. haageana, M. lanigera, M. meissneri, and M. san-angelensis), mainly as a result of ecological speciation. A recent taxonomic proposal considered these taxa as a single species; therefore, we propose their recognition at the species level. Our results also show a high level of incomplete lineage sorting rather than reticulation, which is especially likely in recently diverged species such as those comprising M. ser. Supertextae. The species hypotheses proposed here may be useful in future extinction risk assessments and conservation strategies.
... In Puerto Rico three distinct morphological forms of C. chemnitzia occur (all previously considered to be varieties of C. racemosa). In their circumscription of Caulerpa chemnitzia based on genetic analyses that included taxa that showed a continuous morphological gradient between them, Belton et al. (2014) concluded that the species is broadly defined and did not recognize varieties. Norris et al. (2017) in their treatment of Caulerpa chemnitzia in the Gulf of California chose to acknowledge the different morphologies in the genus as varieties. ...
... chemnitzia have not been recognized for Puerto Rico. See Belton et al. (2014: fig. 8E) for an illustration of the holotype. ...
... Caulerpa chemnitzia var. occidentalis J. Agardh Syntype Localities: "ß, ad oras calidiores atlanticas Americae a sinu superiore Mexicano saltem ad Pernambucco usque descendens" (Agardh 1873: 37); upper Gulf of Mexico to Pernambuco, Brazil [Veracruz, Mexico, Gulf of Mexico, fide Belton et al. (2014: fig. S4B)]. ...
p>This treatment is a taxonomic study of the benthic species of Chlorophyta known from Puerto Rico, Caribbean Sea. In all, 2 classes, 8 orders, 25 families, 50 genera, and 150 species occur in the benthic marine communities in Puerto Rico. Along with date, place, and author(s) of valid publication for all genera, species, and infraspecific taxa, type locality information and descriptive accounts of vegetative morphological and reproductive anatomy are provided. Distribution of each species is given, and where relevant, comments on their habitat and taxonomic and nomenclature status are discussed. A key to the genera and keys to species within genera are included. Either an in situ image or an illustration accompanies most species. Two varieties each of Bryopsis pennata and Caulerpa cupressoides ; one variety each of Caulerpa chemnitzia, C. racemosa, and Udotea occidentalis ; and a single forma each of Halimeda opuntia and Udotea cyathiformis are reported for Puerto Rico for the first time. Avrainvillea mazei, Anadyomene howei, A. rhizoidifera, and Caulerpa parvifolia are also reported as new geographic records for Puerto Rico. A new section of Ulva , sect. Chaetomorphoides J. N. Norris et D. L. Ballant., is established. Descriptions of one new combination, Halimeda acerifolia (D. L. Ballant.) D. L. Ballant. et J. N. Norris, and two new forma, Udotea cyathiformis f. mesophotica D. L. Ballant, H. Ruiz, et J. N. Norris and U. occidentalis f. radiata D. L. Ballant. et J. N. Norris, are included. Smithsonian Contributions to Botany, number 117, x + 164 pages, 116 figures, 2023.</p
... Uprights axes are called fronds (or assimilators) and bear branchlets (or pinnules), whose overall morphotypes are used to distinguish species. Unfortunately, many Caulerpa species are phenotypically very plastic as a response to environmental factors (Ohba et al., 1992;Littler and Littler, 2000;Belton et al., 2014), which complicates their taxonomic identification. ...
... This is the case of Caulerpa taxifolia (M. Vahl) C. Agardh 1817 whose strain is thought to have originated from southeastern Australian waters (Jousson et al., 2000;Meusnier et al., 2004) and Caulerpa cylindracea Sonder 1845 (Belton et al., 2014, previously described as C. racemosa var. cylindracea, Verlaque et al., 2003, initially thought as endemic to southwestern Australia, although later published records of this species in New Caledonia and Indonesia challenge this hypothesis (Sauvage et al., 2013;Darmawan et al., 2021). ...
... Identification of the genus Caulerpa is often challenging due to the difficulty of assessing morphological boundaries for some species (Belton et al., 2014;Pattarach et al., 2019). In this context, DNA-assisted identification (e.g. ...
The green macroalga Caulerpa ashmeadii was recently found in the North-eastern Atlantic Ocean, on the southern coast of Porto Santo Island (Madeira archipelago). Caulerpa ashmeadii was detected in a subtidal soft-bottom forming scattered patches, mostly among the green calcareous macroalgae Halimeda incrassata and Penicillus
capitatus, within a depth range of 18–26 m. Some isolated patches were also observed in bare sand. The species is currently distributed over an estimated area of approximately 0.22 km2. Caulerpa ashmeadii may represent a recent settler (Nov. 2020), although the hypothesis of the species being present elsewhere in Porto Santo prior to our discovery cannot be discarded. Alternatively, the presence of C. ashmeadii in Porto Santo may indicate the influence of climate change on the composition and structure of benthic communities in the Webbnesia region, which further stresses the importance of conducting more intensive biological and environmental monitoring programmes. Overall, this new record constitutes an exceptional eastern expansion of this marine species by
more than 4200 km, comprising its first register in the Eastern Atlantic Ocean.
... Anyway, in both cases, the absence of Cystoseira sp. and the ubiquitous presence of Ellisolandia elongata (J. Ellis et Solander) K.R. Kind et G.W. Saunders indicated a poor coastal water quality (Arevalo et al., 2007), whereas C. cylindracea is highly adapted also in polluted areas (Ballesteros et al., 1999;Ruitton et al., 2005). ...
A survey to state the spread of Caulerpa cylindracea in the Calabrian Tyrrhenian coasts has been un-dertaken. The research aims to value the role of shipping activities in the ten-year's study from 1999 to 2009, as a vector in the spreading of the species. The outcome of this study has shown that, during the last ten years, the species has colonized most of the regional coastline, on all kinds of substrata, in areas closed to harbours and subjected to high rate of sedimentation. The main effects of Caulerpa cylindracea colonization have resulted in a gradual decrease of crustose species while the turf ones have increased their abundance, altering the native structure of the macroalgal assemblages. These results confirm the extremely invasive behaviour of this strain in the Calabrian Tyrrhenian coasts and, more generally, in the Mediterranean Sea.
... According to Farnsworth et al., [11] species identification is essential to documenting, managing, and sustaining organism diversity. The Caulerpa algae are well-known for their great phenotypic plasticity, which refers to the ability of the same species to exhibit a variety of morphological structures in response to changing environmental conditions [12,13]. Accurate identification and characterization of seaweed species are critical for resolving their taxonomic uncertainty, especially for those with commercial interests. ...
... Caulerpa cylindracea has been raised from Caulerpa racemosa var. cylindracea because of its genetic independence [12]. C. cylindracea is one of the most invasive algae species that has been reported by several researchers [55][56][57][58]. ...
The studies of the Bulung Boni and Bulung Anggur (Caulerpa spp.) species and secondary metabolites are still very limited. Proper identification will support various aspects, such as cultivation, utilization, and economic interests. Moreover, understanding the secondary metabolites will assist in developing algae-based products. This study aimed to identify these indigenous Caulerpa algae and analyze their bioactive components. The tufA sequence was employed as a molecular marker in DNA barcoding, and its bioactive components were identified using the GC-MS method. The phylogenetic tree was generated in MEGA 11 using the maximum likelihood method, and the robustness of the tree was evaluated using bootstrapping with 1000 replicates. This study revealed that Bulung Boni is strongly connected to Caulerpa cylindracea. However, Bulung Anggur shows no close relationship to other Caulerpa species. GC-MS analysis of ethanolic extracts of Bulung Boni and Bulung Anggur showed the presence of 11 and 13 compounds, respectively. The majority of the compounds found in these algae have been shown to possess biological properties, such as antioxidant, antibacterial, anticancer, anti-inflammation, and antidiabetic. Further study is necessary to compare the data obtained using different molecular markers in DNA barcoding, and to elucidate other undisclosed compounds in these Caulerpa algae.
... turbinata (J. Agardh) Eubank [Belleza and Liao, 2007] Remarks: In an elegant study of the morphologically complex Caulerpa racemosa group, Belton et al. (2014) provided detailed molecular and morphological evidence recognizing eleven lineages within this large assemblage. One of them (Lineage 6) is composed of plants variously recognized as three varieties of Caulerpa racemosa namely, C. racemosa var. ...
... Caulerpa lamourouxii (Turner) C. Agardh =Caulerpa racemosa var. lamourouxii (Turner) Weber Bosse [Cordero, 1980] Remarks: In the same study of the morphologically complex Caulerpa racemosa group, Belton et al. (2014) provided detailed molecular and morphological information defining Lineage 2 which included this particular variety of C. racemosa. Another small group of plants identified under the same name has been recognized at the species level in the same study under the name Caulerpa oligophylla Montagne after comparison with its type specimen. ...
... [Trono, 1997[Trono, , 2018Belleza and Liao, 2007;Magdugo et al., 2020] Remarks: This species is particularly notorious for its phenotypic plasticity. Belton et al. (2014) have used molecular and morphological characters to clarify the phylogenetic positions of its various varieties and forms. The reports cited above have not indicated the particular subspecific type from this region. ...
The marine benthic flora of the Philippines is among the richest in the Indo-West Pacific region. However, some regions like the eastern Caraga in the northeastern section of Mindanao Island are poorly known in terms of their seaweed diversity. A survey of the published literature was conducted to compile baseline information. This report on the diversity of the marine green algae recorded a total of 62 species and one form in 25 genera, 13 families, and four orders. Many species have ecological and economic importance such as contributors to reef building, sources of valuable chemicals and food for humans. The enumeration of the marine green algae as well as those of other seaweed groups is useful for better resource management and conservation, guided extraction and sound policy formulation towards greater protection of coastal habitats and aquatic resources therein.
