Corresponding author: James Davis Reimer (
Academic editor: Bert W. Hoeksema
The zoanthid genus
Zoanthids are a hexacorallian order (
While zoanthids can be found in a wide variety of marine environments from shallow waters to the deep sea, much of their diversity is found in subtropical and tropical coral reef ecosystems, particularly within the suborder
There is one additional family of brachycnemic zoanthids, the monotypic
In this study, utilizing both morphological and molecular techniques, we formally describe two new
Summary of morphological characters of major brachycnemic zoanthid genera compared with specimens examined in this study (adapted from
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Yes | Simple | Mesogleal | No | Yes | No |
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No | Double | Mesogleal | Yes | Yes | No |
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No | Simple | Mesogleal | No | No | Yes |
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Partial | Simple | Endodermal | No | No | No |
Specimens in this study | Partial | Simple | Endodermal | No | No | No |
Internal structure of
Specimens were collected as detailed in
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NSMT Co1554 |
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North West Reef, GBR, Australia |
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10 | Nov. 17, 2009 | JD Reimer | NA | NA |
HI141 |
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Sykes Reef, GBR, Australia |
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21 | Nov. 18, 2009 | JD Reimer |
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HI142 |
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Sykes Reef, GBR, Australia |
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21 | Nov. 18, 2009 | JD Reimer |
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HI143 |
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Sykes Reef, GBR, Australia |
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21 | Nov. 18, 2009 | JD Reimer | NA |
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HI144 |
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Sykes Reef, GBR, Australia |
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20 | Nov. 18, 2009 | JD Reimer | NA |
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HI145 |
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Sykes Reef, GBR, Australia |
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18 | Nov. 18, 2009 | JD Reimer |
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HI199 |
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Heron Channel, GBR, Australia |
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22 | Nov. 22, 2009 | JD Reimer | NA | NA |
HI200 |
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Heron Channel, GBR, Australia |
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23 | Nov. 22, 2009 | JD Reimer |
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HI209 |
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Sykes Reef, GBR, Australia |
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28 | Nov. 23, 2009 | JD Reimer |
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HI214 |
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Sykes Reef, GBR, Australia |
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9 | Nov. 23, 2009 | JD Reimer |
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MTQ G65793 |
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Sykes Reef, GBR, Australia |
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4 | Nov. 23, 2009 | JD Reimer |
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HI224 |
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Heron Channel, GBR, Australia |
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26 | Nov. 24, 2009 | JD Reimer |
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HI225 |
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Heron Channel, GBR, Australia |
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25 | Nov. 24, 2009 | JD Reimer |
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HI227 |
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Heron Channel, GBR, Australia |
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25 | Nov. 24, 2009 | JD Reimer |
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HI231 |
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Heron Channel, GBR, Australia |
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23 | Nov. 24, 2009 | JD Reimer |
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HI101114-13 |
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Sykes Reef, GBR, Australia |
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29 | Nov. 14, 2010 | JD Reimer | NA | NA |
RMNH Coel 40098 |
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Manza, Okinawa, Japan |
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25 | Sept. 1, 2008 | JD Reimer et al. | NA | NA |
MISE 545 |
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Teniya, Okinawa, Japan |
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1 to 2 | Sept. 5, 2008 | JD Reimer et al. | NA | NA |
MISE 546 |
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Teniya, Okinawa, Japan |
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1 to 2 | Sept. 5, 2008 | JD Reimer et al. | NA | NA |
USNM 1194728 |
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Teniya, Okinawa, Japan |
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1 to 2 | Sept. 5, 2008 | JD Reimer et al. |
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MISE 549 |
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Teniya, Okinawa, Japan |
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1 to 2 | Sept. 5, 2008 | JD Reimer et al. | NA | NA |
NSMT Co1553 |
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Teniya, Okinawa, Japan |
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1 to 2 | Sept. 5, 2008 | JD Reimer et al. |
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NA |
MISE 560 |
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Yona, Okinawa, Japan |
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13 | Sept. 24, 2008 | JD Reimer, T Fujii | NA | NA |
MISE 1092 |
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Teniya, Okinawa, Japan |
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1 | July 2008 | JD Reimer | NA | NA |
MISE 1093 |
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Teniya, Okinawa, Japan |
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Inter-tidal | July 2008 | JD Reimer | NA | NA |
MISE 1115 |
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Tinyuhama, Korijima, Okinawa, Japan |
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24 | Dec. 28, 2008 | JD Reimer |
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MISE 1116 |
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Tinyuhama, Korijima, Okinawa, Japan |
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24 | Dec. 28, 2008 | JD Reimer |
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MISE 1400 |
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Omonawa, Tokunoshima, Kagoshima, Japan |
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9 | March 9, 2010 | JD Reimer | NA | NA |
MISE 1401 |
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San, Tokunoshima, Kagoshima, Japan |
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10 | March 10, 2010 | JD Reimer | NA | NA |
MISE 1402 |
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San, Tokunoshima, Kagoshima, Japan |
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12 | March 10, 2010 | JD Reimer | NA | NA |
MISE 1403 |
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Zampa, Okinawa, Japan |
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NA | August 29, 2008 | JD Reimer | NA | NA |
MISE MO-100 |
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Tebiro, Amami-Oshima, Kagoshima, Japan |
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10 | March 16, 2011 | M Obuchi | NA | NA |
Abbreviations: GBR=Great Barrier Reef; NA=not acquired. Sample number abbreviations as in Methods.
Specimens were examined, decalcified, and sectioned as detailed in
External morphology of specimens was examined using both preserved specimens and in situ images. Polyp dimensions (oral disk diameter, polyp height) for both in situ and preserved specimens were obtained, as were the following data: tentacle number, color of polyp, color(s) of oral disk, relative amount of sand encrustation, associated/substrate species. Additionally, the relative development of the coenenchyme was examined.
For internal examinations, the following data were obtained: mesentery form (brachycnemic or macrocnemic arrangement), mesentery numbers, presence/absence of encrustations, location of encrustations, location and development of the sphincter muscle, presence/absence of gonads. Decalcification, histology and electron microscopy were performed as described in
Undischarged nematocysts were measured from tentacles, column, actinopharynx, and mesenterial filaments of polyps (specimens examined n=2-4 colonies/species) for both new species. 400x images of the nematocysts were obtained by optical microscope, and measured using the software ImageJ (National Institutes of Health, USA). Nematocyst nomenclature generally followed
Phylogenetic analyses are detailed and were performed on both species in
Museum of Tropical Queensland, Townsville, Australia.
Smithsonian National Museum of History, Washington D.C., USA
National Museum of Nature and Science, Tokyo, Japan
Naturalis Biodiversity Center, Leiden, the Netherlands
Molecular Invertebrate Systematics and Ecology Laboratory, University of the Ryukyus, Nishihara, Okinawa, Japan
Additional data related to both species, including tables, phylogenetic trees, and histological images, are reported in
Longitudinal section of
Differs from
Named for Dr. Julian Caley, the leader of the Australian Census of Coral Reef Ecosystems (CReefs) project. Dr. Caley’s acceptance of the first author’s participation in CReefs led to the discovery of this species. Noun in genitive.
Specimens from the Great Barrier Reef were found at depths from 4 to 29 m. Despite repeated surveys, no
This species can close its polyps much more rapidly than those of other zooxanthellate zoanthid genera (Reimer pers. obs).
Comparison of various features of
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Distribution | NE Madagascar | Middle Ryukyu Islands, Okinawa, Japan | Heron Island, Great Barrier Reef, Australia |
Depth | No data | Intertidal to 25 m | 4 to 29 m |
Oral disk color | Greenish-beige to yellow | Light gray-blue, white, rust or deep wine red | Light gray-blue, white, or deep wine red |
Polyp diameter (mm) | 1.5 to 5.0 | 2.2 to 5.1 | 2.3 to 5.0 |
Polyp height (mm) | 2.0 to 12.0 | 2.0 to 8.5 | 2.5 to 3.0 |
Number of tentacles (avg. ± SE) | 38–44 | 32–42 (38±3.0) | 28–40 (33±3.9) |
Cnidae | |||
Column | Microbasic mastigophores | Holotrichs | Holotrichs |
Pharynx | Microbasic mastigophores | Holotrichs, basitrichs, spirocysts | Holotrichs, basitrichs |
Tentacles | Holotrichs, spirocysts | Holotrichs, basitrichs, spirocysts | Holotrichs, basitrichs, spirocysts |
Filaments | Holotrichs, microbasic mastigophores | Holotrichs, |
Holotrichs, basitrichs, |
Originally listed in Table S1 in
Cytochrome oxidase subunit I: HM991247-HM991257
Mitochondrial 16S ribosomal DNA: HM991230-HM991242
Holotype, specimen number NSMT-Co1553. Colony of 17 polyps connected by stoloniferous coenenchyme on a rock approximately 4.5 × 3.0 cm. Polyps approximately 2.0–4.4 mm in diameter, and approximately 2.0–5.4 mm in height from coenenchyme. Polyps and coenenchyme encrusted with irregularly sized and colored sand grains. There was no noticeable variation between holotype and other specimens. Preserved in 99.5% ethanol. Original label.
Differs from
Named for the Okinawan dialect word for Okinawa, “uchina”, the prefecture where this species was first found. Noun in apposition.
Specimens from the Ryukyu Archipelago were found at depths from the intertidal zone to 25 m. Despite repeated surveys focused on zoanthids, no
This species can close its polyps much more rapidly than species of other zooxanthellate zoanthid genera (Reimer pers. obs).
Originally listed in Table S1 in
Cytochrome oxidase subunit I: HM991243-HM991246
Mitochondrial 16S ribosomal DNA: HM991227-HM991229
Cnidae of
As stated previously (
Both new species in this study were found in areas notable for their strong currents. Both species, although zooxanthellate, were found in areas somewhat sheltered from direct sunlight, unlike many
As mentioned in the species’ descriptions, for now it appears that both
Two new species of
The discovery of these two species (detailed in
We recommend the utilization of the combination of both molecular results (
The author thanks the following people at the University of the Ryukyus (UR): Dr. Mamiko Hirose (now Ochanomizu U.) for histology help, and Dr. Frederic Sinniger (now JAMSTEC), Masami Obuchi (now Biological Institute on Kuroshio) and Masaru Mizuyama for specimens, distribution information, and in situ images. On the Great Barrier Reef, the Census of Coral Reef Ecosystems (CReefs) Australia Project, and in particular Dr. Julian Caley and Shawn Smith (both AIMS) are thanked for sampling help, and Gary Cranitch (AIMS) for images. CReefs was a field project of the Census of Marine Life. Dr. Bert Hoeksema (Naturalis Biodiversity Center) and Julian Sprung are thanked for information on
Types, relative abundance and sizes of cnidae in