identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
DA5BB272BF35FFF02EAD6D81FD48F89A.text	DA5BB272BF35FFF02EAD6D81FD48F89A.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Karenia papilionacea A. I. Haywood et K. A. Steidinger 2004	<div><p>3.1.1 Karenia papilionacea A.Į.Haywood et K.A.Steidinger 2004</p><p>The cells were solitary, transversely elongated, wider than long, dorsoventrally compressed, dorsally convex and ventrally concave, 34 – 41 μm long, and 38 – 44 μm wide, with a length to width ratio of about 0.9 (Figure 2). The epicone possessed a pointed apical protrusion (carina). The hypocone was bilobed with a deeply excavated antapex (Figure 2A, B, D – F, H – N). The cingulum was slightly pre-median to median, descending, and displaced by about one cingular width (Figure 2B, I, L). The sulcus continued as a short open extension onto the epicone (Figure 2B, D, L). A short linear apical groove bisected the apex and extended to about one-third of the dorsal epicone (Figure 2A, D, I, L, M). The nucleus was spherical and located in the left lobe of the hypocone, as seen in the ventral view (Figure 2A, B, E, H, K, N). The cytoplasm contained numerous (16 – 26) kidney-shaped to elongated, golden-brown chloroplasts located peripherally (Figure 2C, G). Swimming cells were able to bend along the longitudinal axis, resulting in slow movements resembling a flying butterfly (Figure 2O – R).</p></div>	https://treatment.plazi.org/id/DA5BB272BF35FFF02EAD6D81FD48F89A	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Al-Kandari, Research Article Manal;Saburova, Maria;Polikarpov, Igor;Larsen, Jacob;Lundholm, Nina;Hussain, Sumaiah	Al-Kandari, Research Article Manal, Saburova, Maria, Polikarpov, Igor, Larsen, Jacob, Lundholm, Nina, Hussain, Sumaiah (2025): Morphological and molecular characterization of Kareniaceae (Dinophyceae, Gymnodiniales) in Kuwait’s waters. Botanica Marina (Warsaw, Poland) 68 (2): 155-173, DOI: 10.1515/bot-2024-0083, URL: https://doi.org/10.1515/bot-2024-0083
DA5BB272BF35FFF02EAD688EFA94FB35.text	DA5BB272BF35FFF02EAD688EFA94FB35.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Karenia selliformis A. I. Haywood, K. A. Steidinger et L. MacKenzie 2004	<div><p>3.1.2 Karenia selliformis A.Į.Haywood, K.A.Steidinger et L.MacKenzie 2004</p><p>The cells were solitary, oval, longer than wide, strongly dorsoventrally compressed, 30 – 33 μm long, and 24 – 27 μm wide, with a length to width ratio of about 1.2 (Figure 3). The epicone was broadly dome-shaped with slightly convex sides. The hypocone was hemispherical with a bilobed, centrally excavated antapex and a slightly protruding right lobe (Figure 3A, B, E – I). The cingulum was slightly premedian, descending, and displaced by 1.5 – 2 cingular widths. The sulcus was narrow; its left margin opened onto the epicone as a narrow extension (Figure 3A, B, E, F). The narrow and linear apical groove originated just above and slightly to the right of the proximal end of the cingulum and extended across the apex to about one-third down on the dorsal side of the epicone. The apex was slightly indented by the apical groove (Figure 3A, B, E, F, H, I). The nucleus was large, oblong, transversely elongated, and located in the center of the hypocone (Figure 3C, I, J, L). The cytoplasm contained numerous (48 – 56) plate-like, elongated, golden-brown chloroplasts with internal pyrenoids (Figure 3C, D, K).</p></div>	https://treatment.plazi.org/id/DA5BB272BF35FFF02EAD688EFA94FB35	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Al-Kandari, Research Article Manal;Saburova, Maria;Polikarpov, Igor;Larsen, Jacob;Lundholm, Nina;Hussain, Sumaiah	Al-Kandari, Research Article Manal, Saburova, Maria, Polikarpov, Igor, Larsen, Jacob, Lundholm, Nina, Hussain, Sumaiah (2025): Morphological and molecular characterization of Kareniaceae (Dinophyceae, Gymnodiniales) in Kuwait’s waters. Botanica Marina (Warsaw, Poland) 68 (2): 155-173, DOI: 10.1515/bot-2024-0083, URL: https://doi.org/10.1515/bot-2024-0083
DA5BB272BF35FFF12D3F6BE0FEEFF862.text	DA5BB272BF35FFF12D3F6BE0FEEFF862.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Karlodinium ballantinum Salas 2008	<div><p>3.1.3 Karlodinium ballantinum Salas 2008 (Figure 4)</p><p>The cells were solitary, more or less oval in ventral view, 14 – 18 μm long, and 10 – 13 μm wide, with a length to width ratio of about 1.4. The cingulum was equatorial, and the epicone and hypocone were both hemispherical and nearly equal in size (Figure 4A – D, L). The cingulum originated at the longitudinal axis of the cell, descended, and displaced about two cingular widths or 25 – 30 % of the cell length. The sulcus was sigmoid; it formed a short finger-like projection onto the epicone and continued a narrow, oblique connection between the two ends of the cingulum before fading and becoming wider posteriorly (Figure 4A – D). A short apical groove was linear and extended across the apex and briefly down the dorsal epicone (Figure 4B – E). In squashed cells, a pattern of knob-like structures lining the lower margin of the cingulum was discerned (Figure 4G). No ventral pore was observed. The nucleus was located in the central part of the cell, close to the dorsal side (Figure 4H – J). Numerous (8 – 19) irregularly shaped chloroplasts with internal pyrenoids were present (Figure 4K). Cell division occurred at the motile stage (Figure 4F).</p></div>	https://treatment.plazi.org/id/DA5BB272BF35FFF12D3F6BE0FEEFF862	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Al-Kandari, Research Article Manal;Saburova, Maria;Polikarpov, Igor;Larsen, Jacob;Lundholm, Nina;Hussain, Sumaiah	Al-Kandari, Research Article Manal, Saburova, Maria, Polikarpov, Igor, Larsen, Jacob, Lundholm, Nina, Hussain, Sumaiah (2025): Morphological and molecular characterization of Kareniaceae (Dinophyceae, Gymnodiniales) in Kuwait’s waters. Botanica Marina (Warsaw, Poland) 68 (2): 155-173, DOI: 10.1515/bot-2024-0083, URL: https://doi.org/10.1515/bot-2024-0083
DA5BB272BF38FFFD2E836EBBFB6EF8FA.text	DA5BB272BF38FFFD2E836EBBFB6EF8FA.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Karenia selliformis A. I. Haywood, K. A. Steidinger et L. MacKenzie 2004	<div><p>4.1 Karenia selliformis</p><p>K. selliformis was the first species of the family Kareniaceae identified in Kuwait ’ s waters. Its appearance was linked to red tide and large fish kills in Kuwait Bay in September- October 1999 (Heil et al. 2001; Husain and Faraj 2000). It was subsequently recorded again in 2001 but did not form a bloom and was reported at a concentration of 400 cells l −1 (Glibert et al. 2002). This species was initially reported as Gymnodinium cf. mikimotoi (Husain and Faraj 2000) and then as Gymnodinium sp. or G. selliforme (Heil et al. 2001), and Karenia sp. (Glibert et al. 2002) (Table 4). In the course of a taxonomic examination of bloom material by A. Haywood and K. Steidinger, the Kuwaiti taxon was believed to be conspecific with yet undescribed Gymnodinium sp. from New Zealand coastal waters, which possessed the laterally elongated nucleus in the hypocone (Chang 1995; MacKenzie et al. 1995, 1996). Upon the description of the genus Karenia (Daugbjerg et al. 2000), this taxon was recognized as fitting the newly erected genus and redescribed as K. selliformis (Haywood et al. 2004) . The authors noted the conspecificity of material from Kuwait due to similar morphology (Haywood et al. 2004; Steidinger et al. 2008). K. selliformis was recurrently found in Kuwait ’ s waters during the phytoplankton taxonomic survey in 2003 – 2007, when its morphology was illustrated by LM and SEM (Al-Kandari et al. 2009, pl. 7) (Table 4).</p><p>The observed morphological characters of K. selliformis cells from field samples and monoclonal culture (strain KW- D10-60) isolated from Kuwait ’ s waters are consistent with the original description of this species (Haywood et al. 2004) and previous records from Kuwait (Al-Yamani and Saburova 2019; Al-Kandari et al. 2009; Heil et al. 2001) in terms of their shape, apical groove length and path, and position of the nucleus (Figure 3; Table 3). The cells observed in this study were slightly larger than those described initially. The cells were 30 – 33 µm long, as compared to 26 – 32 µm (MacKenzie et al. 1996) and 20 – 27 μm (Haywood et al. 2004), but our measurements fall within the range previously reported from Kuwait (17 – 37 μm, Heil et al. 2001; 30 – 34 μm, Al-Yamani and Saburova 2019). The most notable characteristics distinguishing this species from the other Karenia are the large, transversely elongated nucleus located posteriorly and the higher number of chloroplasts (Table 3).</p><p>Molecular phylogenetic analysis using partial LSU rDNA showed that the strain from Kuwait clearly belonged to the K. selliformis clade and was genetically distinct from other Karenia species (Figure 5). Kuwait ’ s strain was most closely related to isolates from New Zealand and was 99.86 % identical (1 bp difference) to the holotype sequence of K. selliformis (U92250). This study provides the first molecular data on K. selliformis isolated from Kuwait ’ s waters and confirms the taxonomic identity of this dinoflagellate that has caused harmful blooms.</p><p>Recently, the genetic variability among the LSU rDNA and ITS sequences of K. selliformis strains originating from distant geographical areas worldwide was found to be high enough to form distinct nested subclades, suggesting the existence of two or three distinct phylotypes, depending on the molecular marker used (Iwataki et al. 2022; Mardones et al. 2020; Orlova et al. 2022). Moreover, the delineated phylotypes were distinguished phenotypically and by their temperature tolerance (Iwataki et al. 2022; Mardones et al. 2020; Orlova et al. 2022), pointing to the potential K. selliformis “ species complex ” (Mardones et al. 2020).</p><p>Our morphological and molecular data support the hypothesis of phylotype separation within K. selliformis, differentiating phenotypically and by temperature preference. In the phylogeny inferred from partial LSU rDNA, the topology of the K. selliformis clade is consistent with that previously described (Mardones et al. 2020). The K. selliformis strains were grouped into distinct nested subclades representing the different phylotypes (Figure 5). The basal subclade (phylotype I) comprised strains isolated from K. selliforms bloom along the coast of the Kamchatka Peninsula in 2020. The strain from subtropical Kuwait clustered with other warm-water strains originating from New Zealand, which were closely related to the Asian strains, belonging to phylotype II, and separated from another subclade that included the Chilean strains (phylotype III). The warm-water strain from Kuwait was represented by strongly dorso-ventrally flattened cells with a distinct antapical notch consistent with the described initially K. selliformis from New Zealand waters. In contrast, strains blooming in Chile (1999), Kamchatka, Russia (2020), and Hokkaido, Japan (2021), which belong to phylotype I, exhibited cells with abundant granular to strap-shaped chloroplasts and bloomed at low temperatures &lt;20 °C (Iwataki et al. 2022; Orlova et al. 2022; Uribe and RuÍz 2001). The strains isolated during the K. selliformis bloom in Chile in 2018 (MN203220 and MN203221) were genetically distinct enough to form phylotype III. Morphologically, cells of this phylotype differed from the holotype strain by weak dorsal-ventral compression, shallower antapical excavation, and a smaller number of lateral pores in the hyposome (Mardones et al. 2020).</p></div>	https://treatment.plazi.org/id/DA5BB272BF38FFFD2E836EBBFB6EF8FA	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Al-Kandari, Research Article Manal;Saburova, Maria;Polikarpov, Igor;Larsen, Jacob;Lundholm, Nina;Hussain, Sumaiah	Al-Kandari, Research Article Manal, Saburova, Maria, Polikarpov, Igor, Larsen, Jacob, Lundholm, Nina, Hussain, Sumaiah (2025): Morphological and molecular characterization of Kareniaceae (Dinophyceae, Gymnodiniales) in Kuwait’s waters. Botanica Marina (Warsaw, Poland) 68 (2): 155-173, DOI: 10.1515/bot-2024-0083, URL: https://doi.org/10.1515/bot-2024-0083
DA5BB272BF38FFFA2D1D688BFC01FBEB.text	DA5BB272BF38FFFA2D1D688BFC01FBEB.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Karenia papilionacea A. I. Haywood et K. A. Steidinger 2004	<div><p>4.2 Karenia papilionacea</p><p>The taxonomic survey by Al-Kandari et al. (2009) revealed the second morphotype of Karenia in Kuwait ’ s waters. It was distinguished from K. selliformis by its cell shape, being wider rather than oblong. The small globular nucleus of this morphotype was located in the left lobe of the hypocone versus the larger, transversely elongated, and posteriorly located nucleus in K. selliformis (Table 3). While the morphology of this taxon resembled K. brevis, its unambiguous identification remained uncertain for a long time. In taxonomic and ecological phytoplankton studies in Kuwait ’ s waters, this taxon has been variously referred to as either Gymnodinium breve (Al-Yamani et al. 2004), Karenia cf. brevis (Al-Kandari et al. 2009, plate 8A, LM), K. brevis (Devlin et al. 2019), or K. papilionacea (Al-Yamani and Saburova 2019; Al-Yamani et al. 2012, plate 60, LM; Polikarpov et al. 2020) (Table 4).</p><p>Morphologically, K. papilionacea is similar to K. brevis, and these two species can be distinguished only by minor details. The cells of K. papilionacea are wider (24 – 36 µm vs 18 – 48 µm), possess pointed rather than bulbous apical protrusions (carina), and have slightly deeper antapical excavations (Haywood et al. 2004). Given the high morphological variability observed in both species, even within the same strain (Persson et al. 2013; Stuart 2011), it is doubtful whether species discrimination based on morphological criteria alone can be considered reliable. Nevertheless, these two species are genetically distinct (Haywood et al. 2004).</p><p>K. papilionacea may have previously been misidentified as K. brevis worldwide. The geographic range of K. brevis appears to be limited to the Gulf of Mexico (Brand et al. 2012; Magaña et al. 2003; Steidinger 2009). Previous reports of fish mortality caused by K. brevis in Asia and the Mediterranean Sea require molecular analysis to verify if they are actually misidentifications of K. papilionacea or other Karenia species (e.g., Tsikoti and Genitsaris 2021; Yamaguchi et al. 2016; Yeung et al. 2005).</p><p>The seven strains of K. papilionacea examined in this study displayed similarity to each other, and the morphological characters observed under LM were consistent with previous reports of this species (e.g., Al-Yamani and Saburova 2019; Kim et al. 2023; Yamaguchi et al. 2016), including its original description (Haywood et al. 2004). Despite some morphological variability within and among strains in cell size and shape, the typical cells were wide, dorsoventrally flattened, with short apical grooves, pointed apices, and round nuclei in the left lobe of the hypocone (Figure 2; Table 3).</p><p>In the phylogenetic tree based on partial LSU rDNA sequences, the strains from Kuwait were grouped together within the monophyletic clade of K. papilionacae among strains isolated from New Zealand (the type locality) and Japan with strong statistical support (Figure 5). The sequences obtained from Kuwait ’ s strains were identical, sharing 99.7 % similarity (2 bp differences) to that of the type strain of K. papilionacea (U92252), and were clearly genetically distinct from the K. brevis clade. Therefore, our results provide the first unambiguous molecular identification of K. papilionacea from Kuwait ’ s coastal waters and resolve the previous regional taxonomic uncertainty for this species. This implies that earlier records of K. brevis -like species in Kuwait ’ s waters need to be reconsidered and attributed to K. papilionacea . Furthermore, the genetic similarity among the strains isolated seven years apart (first in 2014 and then in 2021) and from different sampling sites (Kuwait Bay and semi-enclosed marina, Figure 1C; Table 1) suggests the persistent presence of the same population of this species throughout the study area.</p><p>Similar to K. selliforms, recent phylogenetic analyses based on both LSU and ITS have revealed genetic divergence among the K. papilionacea strains, resulting in the distinction of at least two phylotypes within this species (Kim et al. 2023; Yamaguchi et al. 2016). No morphological differences were observed between phylotypes, while different physiological growth traits were assumed to distinguish them (Yamaguchi et al. 2016). In our LSU-based phylogeny, the topology of the K. papilionacea clade was almost the same as previously reported by Yamaguchi et al. (2016) and Kim et al. (2023) and consisted of three distinct, well-supported subclades (Figure 5). All of Kuwait ’ s strains were placed in a large, diverse, and strongly supported clade (89/0.99) among strains from Australia, New Zealand, Japan, China, and Korea, corresponding to the original phylotype (as per Yamaguchi et al. 2016). The second distinct subclade, with high nodal support (85/0.99), solely comprised K. papilionacea strains restricted to the western Japanese coast (Yamaguchi et al. 2016), representing phylotype I. Two strains originating from the French Atlantic and north-western Mediterranean Sea, Spain, were genetically divergent from those of the original phylotype and phylotype I, clustering separately into a well-supported subclade (86/0.99).</p></div>	https://treatment.plazi.org/id/DA5BB272BF38FFFA2D1D688BFC01FBEB	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Al-Kandari, Research Article Manal;Saburova, Maria;Polikarpov, Igor;Larsen, Jacob;Lundholm, Nina;Hussain, Sumaiah	Al-Kandari, Research Article Manal, Saburova, Maria, Polikarpov, Igor, Larsen, Jacob, Lundholm, Nina, Hussain, Sumaiah (2025): Morphological and molecular characterization of Kareniaceae (Dinophyceae, Gymnodiniales) in Kuwait’s waters. Botanica Marina (Warsaw, Poland) 68 (2): 155-173, DOI: 10.1515/bot-2024-0083, URL: https://doi.org/10.1515/bot-2024-0083
DA5BB272BF3FFFFB2D3F6B85FB24FC0B.text	DA5BB272BF3FFFFB2D3F6B85FB24FC0B.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Karlodinium ballantinum Salas 2008	<div><p>4.3 Karlodinium ballantinum</p><p>Like Karenia, Karlodinium species possess the characteristic straight apical groove but differ in the presence of a ventral pore on the epicone in most species and a unique type of amphiesma with plugs. Typical Karlodinium cells are small, globular rather than flattened like most Karenia, and their straight versus sigmoid apical groove distinguishes them from the morphologically similar Takayama species (Daugbjerg et al. 2000; de Salas et al. 2003). Most Karlodinium species share similar morphology and often overlap in size, shape, and diagnostic morphological characters. Species differentiation within this genus is based on cell shape, the presence/ absence of a ventral pore, the position of the nucleus, and the arrangement of chloroplasts. These characters are often difficult to distinguish in preserved samples but may be discerned by observing live cells (e.g., Bergholtz et al. 2006; de Salas et al. 2008).</p><p>Until recently, dinoflagellates belonging to the genus Karlodinium have not been recognized from Kuwait ’ s waters due to research being restricted to Lugol ’ s preserved samples and possible misidentification with other small gymnodinioid taxa. Five Karlodinium morphotypes have recently been reported from Kuwait based on the observations of live cells in freshly collected water samples (Table 4), including Karl. australe, Karl. decipiens, Karl . digitatum (as Karenia digitata), Karl. gentienii, and Karl. veneficum, as reported by Al-Yamani and Saburova (2019); however, no molecular data were provided for these species. In this study, two Kuwait strains, KW-JL-07 and KW-E9-046, exhibited the typical Karlodinium morphology, with small, nearly spherical to ellipsoidal cells, possessing a short straight apical groove, a large centrally located nucleus, and several elongated peripheral chloroplasts. The lack of a ventral pore in this species (as discerned by LM) sets it apart from most Karlodinium taxa previously reported in Kuwait, suggesting the finding of an unrecorded taxon. Among the currently described Karlodinium species, an inconspicuous or lacking ventral pore has been reported in Karl. antarcticum, Karl. ballantinum, Karl . digitatum, Karl. jejuense, and Karl. zhouanum (de Salas et al. 2008; Li and Shin 2018; Luo et al. 2018; Siano et al. 2009; Yang et al. 2000). The cell morphology of Kuwait ’ s strains differed from that described in four out of five pore-lacking Karlodinium species in terms of shape (ellipsoidal instead of elongated in Karl. antarcticum), nucleus position (centrally located nucleus versus those in the epicone in Karl. zhouanum or in the hypocone in Karl. antarcticum and Karl . digitatum), the apical groove length that extends for a short distance onto the dorsal epicone compared to its longer path (1/4 – 1/2 down the dorsal epicone) in Karl. antarcticum, Karl . digitatum, Karl. jejuense, and Karl. zhouanum, and the knob-like structures (microprocesses) lining the lower margin of the cingulum (Figure 4G) have not been observed in other Karlodinium species. The morphology observed in Kuwait ’ s strains corresponds well with that described for Karl. ballantinum (Table 3). Although de Salas et al. (2008) originally reported this species as lacking a ventral pore, further SEM analysis by Benico et al. (2020) of strains from the Philippines and Japan revealed a pore or pore-like shallow depression in this species. As our morphological observations of this species were limited to light microscopy, further SEM examination is required to determine whether this character varies in Kuwait ’ s material.</p><p>The phylogenetic tree inferred from the LSU rDNA sequences in the present study also showed a high genetic similarity between the sequences obtained from Kuwait and the type material of Karl. ballantinum from Tasmania (Figure 5). Kuwait ’ s strains shared 98.19 – 99.27 % similarity (5 bp differences) with the holotype strain EF469232. Through combined evidence from molecular phylogeny and morphology, this study reports the identification of K. ballantinum in Kuwait ’ s waters, marking the first record of this species in the entire Gulf.</p></div>	https://treatment.plazi.org/id/DA5BB272BF3FFFFB2D3F6B85FB24FC0B	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	Al-Kandari, Research Article Manal;Saburova, Maria;Polikarpov, Igor;Larsen, Jacob;Lundholm, Nina;Hussain, Sumaiah	Al-Kandari, Research Article Manal, Saburova, Maria, Polikarpov, Igor, Larsen, Jacob, Lundholm, Nina, Hussain, Sumaiah (2025): Morphological and molecular characterization of Kareniaceae (Dinophyceae, Gymnodiniales) in Kuwait’s waters. Botanica Marina (Warsaw, Poland) 68 (2): 155-173, DOI: 10.1515/bot-2024-0083, URL: https://doi.org/10.1515/bot-2024-0083
