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Underwater images of target black coral species: (A) Antipathella subpinnata (Ellis & Solander, 1786), (B) Antipathella wollastoni (Gray, 1857), (C, D), Antipathes dichotoma Pallas, 1766 (branched and whip morphotypes, respectively), (E) Leiopathes glaberrima (Esper, 1792), (F) Parantipathes larix (Esper, 1790), (G) Phanopathes cf. rigida (Pourtalès, 1880). (B courtesy of Federico Betti, Pico Island, Azores.)
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The Mediterranean black coral fauna includes type species of four antipatharian genera belonging to four different families, therefore phylogenetic studies hold great potential for enhancing systematics within the order. The analysis of six Mediterranean antipatharian species by means of nuclear sequence data of internal transcribed spacer (ITS1 an...
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... B4 includes two specimens within the genus Stichopathes (Stichopathes cf. occidentalis and Stichopathes sp.) and seven specimens within the genus Antipathes (Antipathes elegans (Brook, 1889), Antipathes cf. fruticosa, Antipathes grandis Verrill, 1928, Antipathes caribbeana Opresko, 1996, A. griggi, Antipathes sp. 1 and Antipathes sp. ...
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... These habitats are biodiversity hotspots because the colonies can modify the local physiography and tridimensionality due to their branched morphology (Wagner et al. 2012;Bruckner 2016;Gress and Andradi-Brown 2018). They also play an important ecological role as essential habitat for feeding, recruitment, and reproduction of many organisms, including important commercial fish species (Bo et al. 2009(Bo et al. , 2018Wagner et al. 2012;Bruckner 2016). Antipatharians have been used for religious and medical purposes since ancient times (Bruckner et al. 2008;Bo et al. 2012;Lauretta and Penchaszadeh 2017). ...
Antipathes galapagensis, Deichmann (Smithson Misc Collect 9:1–18, 1941), has been the object of intensive fishing in the Eastern Tropical Pacific due to its large and arborescent colonies and dense forests. Despite its importance as a habitat-forming species, little information exists about its basic biology. Thus, the objective of this study is to describe its reproductive cycle. Samplings were performed in Espiritu Santo Archipelago (La Paz Bay, Gulf of California, Mexico) over 22 months. Histological analyses were conducted on 197 coral samples collected to assess their reproductive strategy from 2018 to 2019. For the first time, male and female gametogenic development stages are described for the species, determining the mean diameter and size range of oocytes and spermatocysts for each gametogenesis sub-stage. The black coral A. galapagensis is an external spawner, adopting a partial spawning strategy, showing evidence of sequential hermaphroditism, and this latter representing the first documentation for the order Antipatharia. The estimated colony sexual maturity height is 102 and 93 cm for females and males, respectively. Gametogenesis begins in June and reaches the reproductive peak in September–October, where the highest frequency is observed of mature females and males and partial spawning. The reproductive cycle shows a correlation with seawater surface temperature increase in the study area, which reaches its maximum from September–October. The results provide the first knowledge contribution to the species biology, essential for its protection and conservation management.
... Several studies in the past 15 years have relied on an integrated morpho-molecular approach to clarify the phylogenetic relationships of the order Antipatharia using a combination of mitochondrial and nuclear genes, including 5. 8S,18S,28S,ITS1,ITS2, and trnW-IGR-nad2 (IgrW), (Lapian et al. 2007;Wagner et al. 2010;Bo et al. 2012;Brugler et al. 2013;Macisaac et al. 2013;Bo et al. 2018;Opresko et al. 2020;Chimienti et al. 2022). Most of the studies focused on inferring relationships among and within the families Antipathidae, Aphanipathidae and Myriopathidae (Lapian et al. 2007;Lapian 2009;Wagner et al. 2010), whereas Macisaac et al. (2013), Lü et al. (2021) and Chimienti et al. (2022) investigated the relationships within the family Schizopathidae. ...
... Furthermore, an additional representative of the genus Aphanipathes clustered in clade VII, within the lineage Aphanipathidae 2. To clarify the phylogenetic position of the family Antipathidae, we need to refer to sequences of the type species of Antipathes, Antipathes dichotoma Pallas, 1766. Even if excluded from our study, this species has been shown to be closely related to representatives of the family Aphanipathidae Bo et al. 2018;Horowitz et al. 2020Horowitz et al. , 2022Opresko et al. 2020). This evidence suggests that the clade in which sequences of A. dichotoma are nested represents the real phylogenetic position of the family Antipathidae. ...
... sarothamnoides with the taxa Aphanipathes pedata and Phanopathes cf. rigida (Bo et al. 2009(Bo et al. , 2018Terrana et al. 2021). In our reconstructions these species were sister to clade VII, leading to the hypothesis that this is the real phylogenetic position of the Aphanipathidae and the other lineages are ripe for revision. ...
Black corals occur as part of benthic assemblages from shallow to deep waters in all oceans. Despite the importance in many benthic ecosystems, where these act as biodiversity aggregators, antipatharians remain poorly studied, with 75% of the known species occurring below recreational SCUBA diving depth limits. Currently, information regarding the diversity and evolutionary history is limited, with most studies focusing on Hawaii and the South Pacific Ocean. Other regions of the world have received less attention, such as the Red Sea, where only two black coral families and four genera have been recorded. We provide the first analysis of the molecular diversity of black corals in the eastern Gulf of Aqaba and the northern and central Saudi Arabian Red Sea, based on a dataset of 161 antipatharian colonies collected down to 627 m deep. Based on specimen morphology, we ascribed our material to 11 genera belonging to 4 of the 7 known Antipatharia families, i.e. Antipathidae, Aphanipathidae, Myriopathidae and Schizopathidae. The genus level phylogeny of three intergenic mitochondrial regions, the trnW-IGR-nad2 (IgrW), nad5-IGR-nad1 (IgrN) and cox3-IGR-cox1 was reconstructed including previously published material. Overall, we recovered six molecular clades that included exclusively Red Sea sequences, with the highest diversity occurring at mesophotic depths. This study highlights that diversity of black corals in the Red Sea is much higher than previously known, with seven new generic records, suggesting that this basin may be a hotspot for antipatharian diversity as is known for other taxa. Our results recovered unresolved relationships within the order at the familial and generic levels. This emphasises the urgent need for an integration of genomic-wide data with a re-examination of informative morphological features necessary to revise the systematics of the order at all taxonomic levels.
... There are currently 36 nominal species of the genus Antipathes that have been described in the Indo-Pacific ocean, of which 13 lack type material. The purpose of this paper is to designate a neotype for Antipathes flabellum Pallas, especially in light of the fact that recent DNA studies indicate that Antipathes dichotoma (which is currently the type species of the genus Antipathes, the latter being the type genus of the Antipathidae) is more closely related to species in the family Aphanipathidae than to species that have been universally recognized as belonging in the family Antipathidae (Brugler et al., 2013;Bo et al., 2018;Horowitz et al., 2022). ...
A neotype is designated for the antipatharian coral Antipathes flabellum Pallas, 1766. The neotype was collected off Madagascar (the original type locality is given as the “Oceanus Indicus”). Morphologically, the neotype corresponds closely in corallum shape and skeletal spination to specimens that have traditionally been identified as Antipathes flabellum. Another specimen of A. flabellum from Madagascar, morphologically almost identical to the neotype and described here, has been sequenced using ultra conserved elements and exon nuclear loci, which showed that it falls within the family currently recognized as Antipathidae.
... In the Mediterranean Sea, the branched antipatharian Antipathella subpinnata (Ellis and Solander, 1786) was reported to be one of the most frequently observed species in the mesophotic zone (Bo et al., 2008(Bo et al., , 2018Ingrassia and Di Bella, 2021). Forests of A. subpinnata have been described in the Tyrrhenian Sea (Bo et al., 2009Gravier, 1918), the Ligurian Sea (Bo et al., 2008), the Adriatic Sea (Chimienti et al., 2020), the Sardinian coast (Cau et al., 2015) and the Sicily Channel (Deidun et al., 2014). ...
... Mediterranean Sea Remarks. Antipathes dichotoma is the type species of the Antipathidae; however, molecular studies (Bo et al. 2018;Brugler et al. 2013), including this study (Fig. 2), have found that the species is more closely related to species in the Aphanipathidae than the Antipathidae. A formal review with integrated morphological and molecular data of all species in each family is required to resolve this taxonomic issue. ...
... Although Rhipidipathes is currently in the Aphanipathidae, previous Bo et al. 2018;Terrana et al. 2021) and the present study indicate that the genus is more closely related to species in the Antipathidae. The present study suggests that Rhipidipathes shares a lineage with the genus Blastopathes Horowitz, 2020 (Fig 2). ...
We describe five new species of black corals from the Great Barrier Reef and Coral Sea, collected at depths ranging from 14 to 789 m: two in the family Antipathidae (Antipathes falkorae sp. nov. and Antipathes morrisi sp. nov.), two in the family Aphanipathidae (Aphanipathes flailum sp. nov. and Rhipidipathes helae sp. nov.), and one in the family Cladopathidae (Hexapathes bikofskii sp. nov.). We also present a phylogeny of 80 black corals reconstructed from a target capture dataset of ultraconserved elements and exons, to show the systematic relationships among new and nominal species. This phylogeny also represents a backbone for future species descriptions and research into the evolutionary history of the Antipatharia.
... In the Mediterranean Sea, the branched antipatharian Antipathella subpinnata (Ellis and Solander, 1786) was reported to be one of the most frequently observed species in the mesophotic zone (Bo et al., 2008(Bo et al., , 2018Ingrassia and Di Bella, 2021). Forests of A. subpinnata have been described in the Tyrrhenian Sea (Bo et al., 2009Gravier, 1918), the Ligurian Sea (Bo et al., 2008), the Adriatic Sea (Chimienti et al., 2020), the Sardinian coast (Cau et al., 2015) and the Sicily Channel (Deidun et al., 2014). ...
Antipatharians (black corals) are major components of mesophotic ecosystems in the Mediterranean Sea. The arborescent species Antipathella subpinnata has received particular attention as it is the most abundant and forms dense forests harbouring high levels of biodiversity. This species is currently categorized as "Near Threatened" in the IUCN Red List, due to increasing fishing pressure and bottom-trawling activities. Yet, the effects of ocean warming have never been investigated for this species, nor for any other antipatharians from temperate regions. Our study aimed at evaluating the effects of increasing seawater temperatures on A. subpinnata, by combining predictive distribution modelling with a physiological tolerance experiment. During the latter, we exposed A. subpinnata for 15 days to different temperature conditions spanning the current seasonal range to forecasted temperatures for 2100, while measuring biological endpoints such as oxygen consumption rates and different signs of stress (tissue necrosis, total antioxidant capacity). Unexpectedly, no stress was found at organism nor cellular level (wide thermal breadth) suggesting low susceptibility of this species to mid-term temperature increase. If the response to the 15-days heat stress is representative of the response to longer-term warming, ocean warming is unlikely to affect A. subpinnata. The species distribution model predicted the presence of A. subpinnata at depths that correspond to temperatures colder than its maximum thermal tolerance (as determined by the physiology experiment). This suggests that the presence of A. subpinnata at shallower depths is not limited by physiological constraints but by other ecological factors including interspecific competition.
... With the advent of molecular analyses aiming at the study of phylogenetic relationships and taxa boundaries, several inconsistencies between the phylogenies and the current taxonomy of black corals were revealed. This has led to several Antipatharia families and genera currently recognised as polyphyletic Bo et al. 2018; Barret et al. 2020;Lü et al. 2021). For example, the intergenic spacer between nad5-IGR-nad1 (igrN), which is the most variable region of the mitochondrial genome of black corals, in combination with trnW- , and the spacer between cox3 and cox1 (cox3-cox1), have been used to clarify the phylogenetic position and describe new genera and species within the order Antipatharia (Thoma et al. 2009;Brugler et al. 2013;MacIsaac et al. 2013;Horowitz et al. 2020;Opresko and Molodtsova 2021). ...
... For example, the intergenic spacer between nad5-IGR-nad1 (igrN), which is the most variable region of the mitochondrial genome of black corals, in combination with trnW- , and the spacer between cox3 and cox1 (cox3-cox1), have been used to clarify the phylogenetic position and describe new genera and species within the order Antipatharia (Thoma et al. 2009;Brugler et al. 2013;MacIsaac et al. 2013;Horowitz et al. 2020;Opresko and Molodtsova 2021). Moreover, the rRNA internal transcribed spacer region (comprising the 18S, ITS1, ITS2, and 5.8S) has been used in a growing number of publications to elucidate phylogenetic relationships within the families Antipathidae Ehrenberg, 1834, Aphanipathidae Opresko, 2004, and Schizopathidae Brook, 1889(Lapian et al. 2007Bo et al. 2018; Barret et al. 2020), as well as in population connectivity studies (Terrana et al. 2021). Finally, the use of next generation sequencing techniques, such as ultra-conserved elements, has recently proven useful in inferring relationships at species level in black corals (Horowitz et al. 2020). ...
... This is relatively unsurprising as previous works already highlighted that the genus Bathypathes, as currently defined based on morphological characters, is polyphyletic. For example, rDNA phylogenetic reconstructions showed that Bathypathes, comprising representatives of the type species, B. patula, clusters with Stauropathes (Bo et al. 2018;Lü et al. 2021), and one nuclear sequence of Bathypathes sp. (MG023167-YPM IZ 028566) is close to the genus Telopathes (Bo et al. 2018;Lü et al. 2021). ...
A black coral, Bathypathes thermophila Chimienti, sp. nov. is described from the Saudi Arabian coasts of the Gulf of Aqaba and north Red Sea (Neom area) using an integrated taxonomic approach. The morphological distinctiveness of the new species is confirmed by molecular analyses. The species thrives in warm and high salinity waters typical of the Red Sea at bathyal depths. It can form colony aggregations on muddy bottoms with scattered, small hard substrates. Colonies are monopodial, feather-like, and attached to a hard substrate through a thorny basal plate. Pinnules are simple, arranged biserially and alternately, and all the same length (up to approximately 20 cm) except for few, proximal ones. Spines are triangular, laterally compressed, subequal, smooth, and simple or rarely bifurcated. Polyps are elongated transversely, 1.5–2.0 mm in transverse diameter. Large colonies can have one or few branches, whose origin is discussed. The phylogenetic position of B. thermophila sp. nov. within the order Antipatharia, recovered using three mitochondrial markers, shows that it is nested within the family Schizopathidae. It is close to species in the genera Parantipathes , Lillipathes , Alternatipathes , and Umbellapathes rather than to the other available representatives of the genus Bathypathes , as currently defined based on morphology. In agreement with previous findings, our results question the evolutionary significance of morphological characters traditionally used to discriminate Antipatharia at higher taxonomic level.
... The anthozoan order Antipatharia (black coral) currently comprises 7 families, 7 subfamilies, 45 genera, and 279 species, with ~ 75% of species occurring deeper than 50 m (Cairns, 2007). With advances in SCUBA diving allowing access to greater depths and an increased number of remotely operated vehicles exploring the deep sea, black coral evolution, diversity, and life history are now the focus of research programs around the world (Baliński et al., 2012;Barrett et al., 2020;Bo et al., 2018;Gress et al., 2020;Hitt et al., 2020;Horowitz et al., 2020;Ruiz-Ramos et al., 2015;MacIsaac et al., 2013;McFadden et al., 2021;Molodtsova & Opresko, 2017;Rakka et al., 2020;Tapia-Guerra et al., 2021;Tazioli et al., 2007;Terrana et al., 2021;Wagner et al., 2012). With few exceptions (select scleractinians (Kitahara et al., 2016) and cerianthids (Stampar et al., 2014)), anthozoan mitochondrial DNA is known to evolve slowly (Hellberg, 2006;Shearer et al., 2002). ...
... Extremely low variability combined with mitogenomes of the same length suggests that one of these taxa may have been misidentified and thus does not represent a separate genus. Additionally, Bo et al. (2018) showed that specimens of A. dichotoma have affinity to species within the family Aphanipathidae, not Antipathidae. Sibopathes cf. ...
A 2013 study revealed that three morphologically distinct antipatharian genera (Dendrobathypathes, Lillipathes, Parantipathes) from the eastern North Pacific (ENP) are genetically indistinguishable using three mitochondrial and four nuclear markers (7,203 bp). To investigate whether this lack of molecular variability extends beyond three mitochondrial genes, we sequenced the complete mitogenome of a single representative within each genus. Dendrobathypathes was the only specimen from the 2013 study containing high molecular weight (HMW) DNA. In terms of geographic proximity to the ENP, the closest Lillipathes and Parantipathes yielding HMW DNA were from the central North Pacific near Hawai'i. Based on cox3-IGR-cox1, Lillipathes and Parantipathes each contained two variable sites and thus were not equivalent substitutes for specimens from the ENP. Nonetheless, variation was extremely low when comparing the mitogenomes, with 32 variable positions across 17,687 bp. Pairwise comparisons revealed 18 (Dendrobathypathes and Parantipathes) and 23 (Lillipathes and Parantipathes; Lillipathes and Dendrobathypathes) variable sites. An ML-based phylogenetic reconstruction using 13 protein-coding genes and two rRNAs revealed that the three North Pacific genera grouped in a clade with Atlantic Dendrobathypathes, while Atlantic Parantipathes spp. formed a sister clade. Previous research hypothesized that hybridization with subsequent introgression was responsible for the lack of variability among genera. Due to uniparental inheritance and lack of recombination, mtDNA cannot identify hybrids; however, finding Pacific Parantipathes grouping with Dendrobathypathes and Lillipathes rather than Atlantic Parantipathes suggests that the trigeneric complex has a unique evolutionary history. If high-resolution nuclear markers support hybridization, it will be important to elucidate the molecular mechanism that maintains three distinct morphological forms occurring in sympatry.
... The nuclear rDNA internal transcribed spacer (ITS) region spanning partial 18S rDNA, ITS1, 5.8S, ITS2, and partial 28S rDNA has been proved to successfully evaluate the phylogenetic relationships and establish new taxa for black coral species (Lapian et al., 2007;Bo et al., 2009Bo et al., , 2018Wagner et al., 2010). Meanwhile, two frequently used markers, mitochondrial cox3 -IGR-cox1 (COI) and trnW -IGR-nad2 (NAD2) were also selected for the phylogenetic analyses. ...
Three deep-sea black corals belonging to the family Schizopathidae, are reported from two seamounts on the Caroline Ridge in the tropical Western Pacific: Umbellapathes parva sp. nov. with depth of 1 488–1 766 m, Telopathes cf. magna MacIsaac & Best, 2013 with depth of 937–1 016 m, and Stauropathes cf. punctata (Roule, 1905) with depth of 942–1 353 m. The latter two species are the first records in the Western Pacific Ocean. U. parva sp. nov. is characterized by monopodial corallum, relatively long unpinnulated stalk with complex pinnulated branches developing from the lowermost primary pinnules and small spines. It differs from two known congeners by much smaller spines and the present of secondary pinnules. T. cf. magna MacIsaac & Best, 2013 has sparsely branched corallum, relatively long and simple pinnules arranged in subopposite or alternate pairs. S. cf. punctata (Roule, 1905) is characterized by the corallum with almost planar pinnulated branches, and pinnules arranged in subopposite pairs. Our results indicate that the corallum size and shape are variable in conspecific specimens particularly those at different growth stages. By contrast, the size of the polypar spines as well as the abpolypar spines have little variation and can be served as a main differing feature for schizopathid species. The phylogenetic analyses using nuclear internal transcribed spacer region (spanning partial 18S rDNA, ITS1, 5.8S, ITS2, and partial 28S rDNA) and two mitochondrial fragments cox3-IGR-cox1 (COI) and trnW-IGR-nad2 (NAD2) showed Telopathes, Stauropathes, and Bathypathes had close relationships, and Umbellapathes formed a sister clade with Alternatipathes.
... European Community 1999), in Annex III of the Berna Convention, in Annex II of the Barcelona Convention for the Mediterranean species, and are also categorized as "threatened" by the International Union for Conservation of Nature (IUCN) Red List of Mediterranean Anthozoa [8] (with the millennial life span L. glaberrima being the only one listed as "endangered"). The main anthropogenic impact affecting these corals is represented by fishing activity, particularly artisanal and recreational activities and bottom trawling, also causing the resuspension of fine sediments [4,9,10]. Rarer activity related to commercial employment for jewelry has also been reported [11]. ...
Antipatharian corals are important structural and complex organisms of the benthic communities inhabiting the Italian seafloor. In this study, the distribution of Italian black corals is reported and mapped for the first time. This review has permitted the identification of occurrences of such vulnerable marine ecosystems along the Italian coasts in a bathymetric range of 42 m to 790 m. Black corals appear to be most conspicuous and widely distributed in the mesophotic zone (from −60 to about −300 m), with a major occurrence on the rocky bottom and shoals. This review also highlights that these communities suffer direct damage from anthropogenic impacts (fishing activity and lost garbage). Finally, this study provides evidence that the reported Italian submarine sites associated with the occurrence of black corals probably represent only a small portion of their real distribution. This statement urges the need to increase monitoring efforts to ensure the protection and the preservation of these pristine assemblages by adopting conservation and management measures.
