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Raw target enrichment of conserved element sequence data for 24 black coral species
<p><span>Deep-sea lineages are generally thought to arise from shallow-water ancestors, but this hypothesis is based on a relatively small number of taxonomic groups. Anthozoans, which include corals and sea anemones, are significant contributors to the faunal diversity of the deep sea, but the timing and mechanisms of their invasion into this biome remain elusive. Here, we reconstruct a fully resolved, time-calibrated phylogeny of 83 species in the order Antipatharia (black coral) to investigate their bathymetric evolutionary history. Our reconstruction indicates that extant black coral lineages first diversified in continental slope depths (~250–3,000 m) during the early Silurian (~437 Ma) and subsequently radiated into, and diversified within, both continental shelf (<250 m) and abyssal (>3,000 m) habitats. Ancestral state reconstruction analysis suggests that the appearance of morphological features that enhanced the ability of black corals to acquire nutrients coincided with their invasion of novel depths. Our findings have important conservation implications for anthozoan lineages, as the loss of "source" slope lineages could threaten millions of years of evolutionary history and confound future invasion events, thereby warranting protection. </span></p>
Raw target enrichment of conserved element sequence data for 24 black coral species
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FIGURE 3 in A new distributional record of the black coral Antipathes cf. griggi in Mactan Island, Philippines
FIGURE 3. Skeletal branch diameter of Antipathes cf. griggi on the terminal branch when viewed under SEM. Scale bar: 0.2 mm.
FIGURE 1 in A new distributional record of the black coral Antipathes cf. griggi in Mactan Island, Philippines
FIGURE 1. Antipathes cf. griggi colony in-situ in the locality area of Barangay Agus, Mactan Island, Philippines. Photo courtesy of Antonio Ayop.
FIGURE 32 in Shallow-water black corals (Cnidaria: Anthozoa: Hexacorallia: Antipatharia) from SW Madagascar
FIGURE 32. Cupressopathes sp. 2, INV.131340. (a) In situ picture of the entire colony showing the asymmetry in the cylindrical shape. (b) Close-up view of the distal part of the colony where a branch is growing. (c) Cross section showing the irregular arrangement of the primary pinnules. (d) Lateral view of the pinnulation on a primary pinnule. (e) Top view of the pinnulation of a primary pinnule. (f) Section of subpinnule 0.15 mm in diameter, polypar side is on the right. (g) Section of secondary pinnule 0.34 mm in diameter, polypar side is on the right. (h) Section of primary pinnule 0.85 mm in diameter. (i) Section of branch 2.2 mm in diameter. (j) Polypar spine of subpinnule. (k) Abpolypar spine of subpinnule. (l) Polypar spine of primary pinnule. (m) Needle-like and bifid spines of a branch.
FIGURE 31 in Shallow-water black corals (Cnidaria: Anthozoa: Hexacorallia: Antipatharia) from SW Madagascar
FIGURE 31. Cupressopathes sp. 1, INV.131365. (a) In situ picture of the whole colony. (b) Detailed view of the pinnulation on a distal branch. (c) Cross section of a pinnulated branch showing the arrangement in irregular rows. (d) Lateral view of the scarce subpinnulation of a primary pinnule. (e) Section of subpinnule 0.1 mm in diameter, polypar side is on the right. (f) Section of primary pinnule 0.22 mm in diameter, polypar side is on the right. (g) Section of distal branch 0.27 mm in diameter. (h) Section of branch 1.10 mm in diameter. (i) Abpolypar spine of a subpinnule. (j) Polypar spine of subpinnule. (k) Tall, acicular needle-like spines occurring on thick branches. (l) Bifid spines occurring on a branch.
FIGURE 29 in Shallow-water black corals (Cnidaria: Anthozoa: Hexacorallia: Antipatharia) from SW Madagascar
FIGURE 29. Cupressopathes cf. pumila INV.131366. (a) In situ pictures of the whole colony (b) In situ close-up view of a primary pinnule. (c) Detailed view of the branching pattern and the pinnulation of a distal branch. (d) Cross section of a distal branch showing the arrangement of the primary pinnules in four irregular rows. (e–f) Detailed top views of the pinnulation of two primary pinnules showing the biserial arrangement of the subpinnules and the variation in their density. (g) Section of subpinnule 0.13 mm in diameter, polypar side is on the right. (h) Section of primary pinnule 0.25 mm in diameter, polypar side is on the right. (i–j) Sections of two branches 0.30 mm and 1.30 mm in diameter, respectively. (k) Abpolypar spine of subpinnule. (l) Polypar spine of subpinnule. (m) Polypar spine on primary pinnule. (n) Tall, acicular and needle-like spines occurring on thick branch.
FIGURE 27 in Shallow-water black corals (Cnidaria: Anthozoa: Hexacorallia: Antipatharia) from SW Madagascar
FIGURE 27. Myriopathes sp. INV.131336. (a) In situ picture of the entire colony. (b) Close-up view of a branch and the pinnules. (c) Close-up view showing the pinnulation in details. (d) Section of subpinnule 0.10 mm in diameter, polypar side is on the right. (e) Section of primary pinnule 0.18 mm in diameter, polypar side is on the right. (f) Section of primary pinnule 0.41 mm in diameter, polypar side is on the right. (g) Section of branch 1.8 mm in diameter. (h) Polypar spine of a subpinnule. (i) Abpolypar spine of a subpinnule. (j) Tall, cylindrical spine on a primary pinnule. (k) Needle-like spines of a branch.
FIGURE 26 in Shallow-water black corals (Cnidaria: Anthozoa: Hexacorallia: Antipatharia) from SW Madagascar
FIGURE 26. Myriopathes cf. ulex (Ellis & Solander, 1786) INV.131356. (a) In situ picture of the entire colony. (b) In situ close-up view of a branch and the primary pinnules. (c) Close-up view showing the pinnulation in details. (d) Section of secondary pinnule 0.12 mm in diameter. The polypar side is on the right. (e) Section of primary pinnule 0.14 mm in diameter. The polypar side is on the right. (f) Section of distal fragment of the main branch bearing primary pinnules and measuring 0.25 mm in diameter. (g) Section of thicker branch 0.34 mm in diameter and bearing primary pinnules. (h) Polypar spine from secondary pinnule. (i) Polypar spine from primary pinnule. (j) Tall, needle-like spine of a branch. (k) Branched, bifid or antler-like spines occurring on a branch measuring 1.8 mm in diameter.
FIGURE 30 in Shallow-water black corals (Cnidaria: Anthozoa: Hexacorallia: Antipatharia) from SW Madagascar
FIGURE 30. Cupressopathes pumila (Brook, 1889) type specimen. (a) Entire colony, holotype Brit. Mus. 83.9.13.15. (b) Schizoholotype USNM 100355 showing different sections of 0.08, 0.11, 0.12 and 0.22 mm in diameter, respectively.
FIGURE 28 in Shallow-water black corals (Cnidaria: Anthozoa: Hexacorallia: Antipatharia) from SW Madagascar
FIGURE 28. Cupressopathes abies (Linnaeus, 1758). (a–b) In situ pictures of a single-stem (INV.131367) and a branched colony (INV.131357). (c) Lateral view of the arrangement of the primary pinnules on a distal part of a colony. (d) Lateral detailed view of the subpinnules on a primary pinnule. (e) Section of subpinnule 0.09 mm in diameter, polypar side is on the right. (f) Section of subpinnule 0.18 mm in diameter, polypar side is on the right. (g) Section of primary pinnule 0.54 mm in diameter, polypar side on the right. (h) Section of main distal branch 0.70 mm in diameter. (i) Polypar spine of a subpinnule. (j) Abpolypar spine of subpinnule. (k) Tall, acicular spine occurring on thick primary pinnule. (l) Bifid spine occurring on branches.
FIGURE 25 in Shallow-water black corals (Cnidaria: Anthozoa: Hexacorallia: Antipatharia) from SW Madagascar
FIGURE 25. Myriopathes cf. stechowi (Pax, 1932) INV.131335. (a) Entire colony in situ. (b) Close-up view of the branching pattern in situ. (c) Detailed view of the pinnulation. (d) Section of subpinnule 0.1 mm in diameter, polypar side is on the right. (e) Section of primary pinnule 0.14 mm in diameter, polypar side is on the right. (f) Section of a branch 1.1 mm in diameter, showing numerous antler-shaped spines. (g) Polypar spine on primary pinnule. (h) Abpolypar spine on primary pinnule. (i) Antler-shaped spine on a branch. (j) Detailed view of the numerous bifid and antler-spines of a branch.
FIGURE 24 in Shallow-water black corals (Cnidaria: Anthozoa: Hexacorallia: Antipatharia) from SW Madagascar
FIGURE 24. Myriopathes cf. myriophylla (Pallas, 1766) INV.131334. (a) In situ picture of the entire colony. (b) Detailed view of a branch and its pinnulation. (c) Close-up view of incipient branches and primary pinnules (ib—incipient branches; pp—primary pinnules; sp—secondary pinnules; tp—tertiary pinnules). (d) An incipient branch with bilateral primary pinnules with up to four uniserial secondary pinnules. (e) Section of a subpinnule 0.09 mm in diameter, the polypar side is on the right. (f) Section of a primary pinnule 0.14 mm in diameter, the polypar side is on the right. (g–h) Branches of different diameter showing the spine arrangement and morphology. (i) Polypar spine of a primary pinnule. (j) Abpolypar spine of a primary pinnule. (k) Tall, needle-like spines of a branch.
FIGURE 23 in Shallow-water black corals (Cnidaria: Anthozoa: Hexacorallia: Antipatharia) from SW Madagascar
FIGURE 23. Stichopathes cf. maldivensis Forster Cooper, 1903. (a–c) In situ pictures showing the different morphologies that this species displays. A branched appearance is sometimes seen as a consequence of a breaking event, as shown by the white arrow. (d) White phenotype (INV.131372) with contracted polyps. (e) Brown-orange phenotype (INV.151020) with expanded polyps. The polyp row is twisting around the stem. (f) Section of a distal fragment measuring 4.5 mm in diameter (INV.131370). Polypar side is on the right. (g) Tall cylindrical polypar spines. (h) Conical abpolypar spines. (i) Close-up view of an abpolypar spine showing the tuberculation occurring on only one side. (j) Secondary spines.
FIGURE 22 in Shallow-water black corals (Cnidaria: Anthozoa: Hexacorallia: Antipatharia) from SW Madagascar
FIGURE 22. Stichopathes cf. diversa (Brook, 1889). (a–b) In situ pictures of the white (INV.131375) and brown (INV.131377) phenotypes. (c) Close-up view of the sagittally compressed polyps, which are found on the outer side of the coils. (d) SEM view of a 2.1 mm in diameter skeleton of an apical coil (INV.131375). The size difference between polypar spines (the outer ones) and abpolypar spines (the inner ones) is clear. (e–f) Blunt, conical polypar spines showing the variability between papillose and tuberculated surfaces and small tubercles at the tip. Small triangular, secondary spines can be seen all around. (g–h) Variability in abpolypar spines that can be sub-triangular or cylindrical, always showing a papillose surface. Small secondary, triangular spines are seen all around as well.
FIGURE 18 in Shallow-water black corals (Cnidaria: Anthozoa: Hexacorallia: Antipatharia) from SW Madagascar
FIGURE 18. Cirrhipathes cf. indica Summers, 1910 (a–b) Morphological variations of the species (a—INV.131348, b—INV.131343). (c) Close-up view of the coil in the apical part of the colony INV.131343, with expanded polyps. (d) Detailed view of a growing apical part in INV.131348 where the verticillated spines are found in the bright, narrow part. (e) Detailed view of an apical part lacking the growing skeleton in INV.131343. (f) Section of a basal part in INV.131343 just above the anchorage and measuring 4.2 mm in diameter. (g) Spines of the apical part shown in e). (h) Spine found just above the basal anchorage and showing a few tubercles at the top. (i) Succession of spines near the basal anchorage, where a fully tuberculated one is seen at the top, only 1 mm above the base of the stem.
FIGURE 15. Cirrhipathes densiflora Silberfeld, 1909 in Shallow-water black corals (Cnidaria: Anthozoa: Hexacorallia: Antipatharia) from SW Madagascar
FIGURE 15. Cirrhipathes densiflora Silberfeld, 1909. (a–e) In situ pictures showing the different phenotypes of the species (a—INV.131368; b—INV.131376; d/e—INV.131374). (f) Sections of the skeleton 1.8 mm in diameter (INV.131352), 2 mm in diameter (INV. 131376), 3.2 mm in diameter (INV.131368) and 2.5 mm in diameter (INV.131374), respectively. (g–i) Spines found on the contorted morphotype (INV.131374, conical polypar spines in (d), cylindrical abpolypar spines in (e, f). (j–k) Spines found on the whip morphotypes (INV.131368, INV.131376). (l) Detailed view of the tubercles at the tip of the spine.
FIGURE 8. Antipathes lentipinna Brook, 1889 in Shallow-water black corals (Cnidaria: Anthozoa: Hexacorallia: Antipatharia) from SW Madagascar
FIGURE 8. Antipathes lentipinna Brook, 1889 type specimen. (a) Entire colony, holotype BMNM 1973.9.20. (b) Schizoholotype USNM 100360 showing a section of a branch 0.3 mm in diameter. Photo of Brook's specimen of A. lentipinna courtesy of Paul Richens of the NHMUK.
FIGURE 19. Cirrhipathes rumphii van Pesch, 1910 in Shallow-water black corals (Cnidaria: Anthozoa: Hexacorallia: Antipatharia) from SW Madagascar
FIGURE 19. Cirrhipathes rumphii van Pesch, 1910. (a, b) In situ pictures of the specimen INV.131361 with large distal spirals. (c) In situ picture of the specimen INV.1312345. with a single distal spiral. (d) Close-up view of the polyp arrangement, leaving one side free of any polyps. (e) Section of the apical growing portion (INV.131345) measuring 1.26 mm in diameter, before tapering. (f) Distal section 1.22 mm in diameter (INV.131361). (g) Distal section 2.76 mm in diameter (INV.131345). (h) Spine found on the growing section shown in (e). (i–k) Spines found on the distal sections shown in (f).
FIGURE 1 in Shallow-water black corals (Cnidaria: Anthozoa: Hexacorallia: Antipatharia) from SW Madagascar
FIGURE 1. (a) Map showing the western position of Madagascar in the Indian Ocean, with the studied sites located off the southwestern coast in the Mozambic Channel. (b) Details of the southwestern coast and the coastal villages near where surveys were performed.
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