Cairns & Kitahara 2012 Key Scleractinia (Anthozoa)

A peer-reviewed open-access journal
ZooKeys 227: 1–47 (2012)

An illustrated key to the genera and subgenera of the Recent azooxanthellate Scleractinia... 1
doi: 10.3897/zookeys.227.3612 Research article
www.zookeys.org Launched to accelerate biodiversity research
An illustrated key to the genera and subgenera of

the Recent azooxanthellate Scleractinia (Cnidaria,
Anthozoa), with an attached glossary
Stephen D. Cairns1, Marcelo V. Kitahara2
1 Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution,

Washington, DC 20560, USA 2 Centro de Biologia Marinha, Universidade de São Paulo, São Sebastião,
SP 11600-000, Brazil
Corresponding author: Stephen D. Cairns ([email protected])
Academic editor: Bert W. Hoeksema | Received 30 July 2012 | Accepted 13 September 2012 | Published 5 October 2012
Citation: Cairns SD, Kitahara MV (2012) An illustrated key to the genera and subgenera of the Recent azooxanthellate
Scleractinia (Cnidaria, Anthozoa), with an attached glossary. ZooKeys 227: 1–47. doi: 10.3897/zookeys.227.3612
Abstract
The 120 presently recognized genera and seven subgenera of the azooxanthellate Scleractinia are keyed us-
ing gross morphological characters of the corallum. All genera are illustrated with calicular and side views
of coralla. All termes used in the key are defined in an illustrated glossary. A table of all species-level keys,
both comprehensive and faunistic, is provided covering the last 40 years.
Keywords
Azooxanthellate, Illustrated Key, Genera, Glossary, Scleractinia
Introduction
The ready identification of azooxanthellate Scleractinia (determined herein by depth of

occurrence and previously published observations) to the genus and species levels has
been hampered by a lack of a comprehensive key to the genera as well as a lack of species
level keys. For instance, the last comprehensive set of keys to the genera was published
by Vaughan and Wells (1943) almost 70 years ago, and relied in part on microstructural
characters that were both hard to observe (requiring thin sectioning) and interpret.
Copyright S.D. Cairns, M.V. Kitahara. This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0
(CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
2 Stephen D. Cairns & Marcelo V. Kitahara / ZooKeys 227: 1–47 (2012)
Since then the number of Recent azooxanthellate genera and species has almost dou-
bled, and new observations on apozooxanthellate species (species that have facultative
symbiosis with zooxanthellae) are also available. Furthermore, what keys exist to the
species level of various taxa or geographic regions are scattered throughout the literature
and of variable quality (Table 1). In this Table, tabular keys are included, as they provide
as much if not more information than a conventional dichotomous key. As result of the
application of molecular data (e.g. Fukami et al. 2008, Kitahara et al. 2010a, Huang
et al. 2011, Stolarski et al. 2011, Arrigoni et al. 2012), the higher taxonomic ranks of
the order Scleractinia were shown to be polyphyletic. As such, a key to this taxonomic
rank seems premature. Thus, it is the purpose of this paper to provide a single, com-
prehensive, illustrated key to the presently recognized 120 azooxanthellate scleractinian
genera and 7 additional subgenera. We constructed the key using gross morphological
characteristics of the corallum, which, when used in conjunction with the glossary and
illustrations, we hope will provide a guide to the proper genus identification. But one
must keep in mind that this key, as most, will not necessarily supply a definitive identi-
fication of the genus, as its use depends on the interpretation of the characters as well as
the variation of that character state. We have used many of the dichotomies published
by Vaughan and Wells (1943), but avoided the microstructural characters, and updated
the taxa. Whereas microstructure is undoubtedly a valuable set of characters to define
genera, in most cases it is not necessary to identify the genera. Among the 120 extant
azooxanthellate scleractinian genera, 74 are illustrated with its type species (~61%).
Within the remaining 46 genera, 20 (~16,6%) have an extinct species as type, repre-
sented by a fossil coral. For them and the remaining 26 genera, the illustrated species
present very well the most important morphological characters of their respective genus.
Table 1. Previously published keys to azooxanthellate taxa, divided as comprehensive keys to all taxa with
in a monophyletic taxon, and partial (faunistic) keys of species. Taxa listed alphabetically by taxon name.
Tabular keys (T) are included.
Comprehensive keys
Anthemiphyllia, species (T) Cairns (1999: 290)
Asterosmilia, species (T) Cairns and Wells (1987: 38)
Aulocyathus, species Cairns (1999: 104)
Caryophyllia, species (T) Cairns (1991: 12)
Caryophyllia, species Kitahara et al. (2010b: 112)
Conocyathus, species Cairns (2004a: 290)
Crispatotrochus, species Kitahara and Cairns (2008: 62)
Deltocyathus, species Kitahara and Cairns (2009: 236)
Dendrophylliidae, genera (T) Cairns (2001: 5)
Flabellidae, genera (T) Zibrowius (1974: 26); Cairns (1989: 45)
Guyniidae, genera (T) Cairns (1989: 41); Stolarski (2000: 23)
Javania, species Cairns (2004b: 10)
Micrabaciidae, genera Cairns (1989: 13)
Placotrochides, species Cairns (2004a: 307)
Scleractinia, families and genera Vaughan and Wells (1943)
Comprehensive keys
Stephanophyllia, species Cairns (1989: 21)
Trochocyathus (Aplocyathus), species (T) Cairns (1999: 85)
Cairns (1988a: 711; 1989: 25; 1997: 5 [T]);
Turbinoliidae, genera
Filkorn (1994: 44)
Faunistic keys
Astrangia, E. Pacific Durham and Barnard (1952: 60)
Azooxanthellate Scleractinia, Antarctica Cairns (1990: 18 [book])
Azooxanthellate Scleractinia, E. Gulf of Mexico Cairns (1977a: 5)
Azooxanthellate Scleractinia, New Zealand Squires and Keyes (1967: 13); Tracey et al. (2012)
Azooxanthellate Scleractinia, NE Pacific Cairns (1994: 13)
Azooxanthellate Scleractinia, NW Pacific Cairns (1994: 75)
Azooxanthellate Scleractinia, S. Australia Cairns and Parker (1992: 4)
Azooxanthellate Scleractinia, Cold Temp. NE Atl. Cairns (1981:3)
Azooxanthellate Scleractinia, Brazil Kitahara (2007: 510)
Balanophyllia, W. Atlantic Cairns (1977b: 133)
Balanophyllia, Japan Ogawa et al. (1998: 145 [in Japanese])
Balanophyllia, W. Atlantic (T) Cairns (2000: 163)
Caryophyllia, New Zealand Cairns (1995: 43)
Caryophyllia, W. Atlantic Cairns (1979: 46)
Caryophyllia, W. Pacific Cairns and Zibrowius (1997: 87, 96)
Caryophyllia and Premocyathus, Japan Ogawa et al. (1999: 115 [in Japanese])
Conotrochus and Trochocyathus, Japan Ogawa et al. (2003: 57 [in Japanese])
Culicia, Australia Cairns (2004a: 274)
Deltocyathus, W. Atlantic Cairns (1979: 91)
Deltocyathus, W. Pacific Cairns and Zibrowius (1997: 121)
Dendrophyllia, Japan Ogawa and Takahashi (1995: 25 [in Japanese])
Flabellum, New Zealand Cairns (1995: 96)
Flabellum, Japan Ogawa and Takahashi (2005: 56 [in Japanese])
Fungiacyathus, W. Pacific (T) Cairns (1989: 6, 7; 1999: 55)
Fungiacyathus, Japan Ogawa and Takahashi (2004: 11 [in Japanese])
Heterocyathus, W. Pacific Hoeksema and Best (1991: 222)
Heterocyathus, Japan Ogawa and Takahashi (2008: 248 [in Japanese])
Heteropsammia, W. Pacific Hoeksema and Best (1991: 222)
Heteropsammia, Japan Ogawa and Takahashi (2008: 248 [in Japanese])
Madracis, W. Atlantic Wells (1973: 19)
Paracyathus and Polycyathus, Japan Ogawa et al. (2000: 55 [in Japanese])
Trochocyathus, W. Pacific Cairns and Zibrowius (1997: 105)
Truncatoflabellum, W, Pacific Cairns (1989a: 62)
Truncatoflabellum, SW Indian Ocean Cairns and Keller (1993: 264)
Truncatoflabellum, Australia (T) Cairns (1998: 397)
Truncatoflabellum, Japan Ogawa (2006: 13 [in Japanese])
Tubastraea, Red Sea Scheer and Pillai (1983: 173)
Tubastraea, Galapagos Cairns (1991: 27)
Tubastraea, Japan Ogawa and Takahashi (1993: 97 [in Japanese])
Turbinoliidae, Japan Ogawa et al. (2002: 27 [in Japanese])
Methods
Some genera are keyed two or even three times because of the variation within those
genera regarding the characters used in the key. In theory, all variations of that genus
will be correctly keyed. Although most couplets are dichotomous, some are polychoto-
mous, such as the columella or colony shape, which allows the reader to clearly see the
multiple states of a particular character.
Although it would be desirable to follow the generic key with keys to all of the ap-
proximately 720 azooxanthellate species, it is a simple fact that not many species level
keys have been published. Those that have been published in the last 35 years are listed
in Table 1, separated as to whether they are keys to all of the taxa within a monophyletic
taxon (comprehensive) or to a more limited fauna of a region (faunistic). Keys made
before 1970 were found to be, in general, not up to date and are thus not included. It
should be noted that fully one-third of the genera (40) are monotypic, and thus do not
require a key following a correct genus identification, and another 22 genera have but
two species. Finally, although they do not include keys, the treatises of Wells (1956) and
Chevalier and Beauvais (1987) include diagnoses of all genera, including those repre-
sented only by extinct species, and thus provide a rich source of taxonomic information.
Other sources of useful taxonomic information include a list of all extant Recent
scleractinian species as of 1999 (Cairns et al. 1999), which also includes a rough in-
dication of their geographic range. The azooxanthellate component of this list is kept
up to date as an on-line resource (www.lophelia.org/online-appendices), which now
includes junior synonyms and depth ranges of the species, and authors of the genera.
A list of the 120 azooxanthellate genera, their authorship, and bathymetric ranges was
also published in Roberts et al. (2009: Table 2.7)
Geographic ranges within brackets in the key are not meant to be considered as
distinguishing characters, but simply informational, which may nonetheless hint at an
incorrect identification. Abbreviations: Ant. = Antarctic or Subantarctic, Atl. = Atlan-
tic, IP = Indo Pacific, IWP = Indo-West Pacific, Pac. = Pacific, SubAnt = Subantarctic;
Cosmopolitan implies occurrence in all three oceans as well as Subantarctic and/or
Antarctic. Museums and Institutions acronyms: AM = Australian Museum (Sydney);
AU = Auckland University Museum (Auckland); CSIRO = Commonwealth Scien-
tific and Industrial Research Organisation (Hobart); JCU = James Cook University
(Townsville); MNHN = Muséum national d’Histoire naturelle (Paris); SBMNH =
Santa Barbara Natural History Museum (Santa Barbara); SIO = Scripps Institute of
Oceanography (San Diego); NZOI = New Zealand Oceanographic Institution (now
the National Institute of Water and Atmospheric Research) (Wellington); USNM
= United States National Museum (now the National Museum of Natural History,
Smithsonian) (Washington, D.C.); YPM = Yale Peabody Museum (New Heaven).
Useful sources for more information about definitions of terms used in the glossary
include: Wells (1956), and Cairns (1981, 1989, 1994).
Key to the Genera and Subgenera of the Recent Azooxanthellate Scleractinia

(An asterisk indicates genera that have azooxanthellate and zooxanthellate representatives)
1a Corallum colonial........................................................................................2
1b Corallum solitary.......................................................................................43
2a Corallum free of attachment (recumbent, usually curved with a broken or
open base, or globular).................................................................................3
2b Corallum firmly attached (arborescent, bushy, encrusting, or reptoid).........5
3a Corallum recumbent (composed of a large primary corallite from which
smaller buds originate); no sipunculid commensalism..................................4
3b Corallum globular; pores in lateral base of colony associated with commensal
sipunculid.................[IWP] Heteropsammia* (in part) Plate 1, Figures A–B
4a Corallum not porous (solid); septa arranged normally...................................
................................................. [Atl. + IP] Anomocora Plate 1, Figures C–D
4b Corallum, especially septa porous; septa arranged in a Pourtalès Plan............
....................................... [Atl. + IWP] Eguchipsammia Plate 1, Figures E–F
5a Corallum arborescent or bushy....................................................................6
5b Corallum encrusting or reptoid..................................................................27
6a Branching intratentacular............................................................................7
6b Branching extratentacular............................................................................9
7a Equal distomadeal budding.........................................................................8
7b Unequal monostomaeous budding................................................................
............................................ [Cosmopolitan] Lophelia Plate 1, Figures G–H
8a Texture of corallum rough (like sandpaper), resulting from a porous theca;
septa arranged in a weak Pourtalès Plan.........................................................
...............................................[W. Pac.] Dichopsammia Plate 1, Figures I–J
8b Texture of corallum smooth or costate, solid; septa arranged normally..........
........................................ Cosmopolitan] Solenosmilia Plate 1, Figures K–L
9a Septal symmetry decameral or octameral, septa in only one cycle; columella
styliform........................[Atl. + IP] Madracis* (in part) Plate 2, Figures A–B
9b Septal symmetry hexameral, septa arranged in multiple cycles; columella pap-
illose, fascicular or absent...........................................................................10
10a Texture of theca and septa rough (like sandpaper), resulting from a porous
theca..........................................................................................................11
10b Texture of theca smooth, granular, or ridged (solid)..................................14
11a Septa arranged in a Pourtalès plan.............................................................12
11b Septa arranged normally............................................................................13
12a Corallum small (bushy), most corallites budding from a common basal coe-
nosteum or from the edge zone of corallites that originate from the basal
coenosteum...................... [Atl. + Pac.] Cladopsammia Plate 2, Figures C–D
12b Corallum large (bushy to arborescent), with multiple successive generations

of budding forming an erect colony...............................................................
............................................. [Atl. + IP] Dendrophyllia Plate 2, Figures E–F
13a Corallum porosity only apparent near calicular edge; found in deep-water:
110-2165 m...................[Atl. + IWP] Enallopsammia Plate 2, Figures G–H
13b Corallum porosity uniform: shallow-water: 0-110 m.....................................
...................................... [Atl. + IP] Tubastraea (in part) Plate 2, Figures I–J
14a Columella absent.......................................................................................15
14b Columella present (papillose, trabecular or fascicular)................................16
15a Corallum large (arborescent), with numerous budding cycles, adjacent coral-
lites often linked with hollow, tubular coenosteal bridges; tabular endothecal
dissepiments common.... [I–P + Subant.] Goniocorella Plate 2, Figures K–L
15b Corallum a small bush, corallites originating from a common basal coenos-
teum or from the sides of other corallites and from relatively few budding
cycles; endothecal dissepiments not prominent..............................................
...............[E. Atl. + New Zealand] Hoplangia (in part) Plate 5, Figures L–M
16a Columella fascicular...................................................................................17
16b Columella papillose or trabecular...............................................................18
17a Pali before septa of third cycle (P3)...............................................................
.............................................. [N. Atl.] Pourtalosmilia Plate 3, Figures A–B
17b Pali absent............................. [W. Pac.] Confluphyllia Plate 3, Figures C–D
18a Columella trabecular, composed of slender ( flattened laths); corallum never
with more than 4 generations of budding......................................................
........................................ [Atl. + W. Pac.] Coenosmilia Plate 3, Figures E–F
18b Columella papillose (composed of rods); corallum composed of many gen-
erations of budding....................................................................................19
19a Axial septal edges dentate.......[W. Pac.] Sympodangia Plate 3, Figures G–H
19b Axial septal edges smooth..........................................................................20
20a Pali absent............ [Cosmopolitan] Madrepora (in part) Plate 3, Figures I–J
20b Pali present................................................................................................21
21a Pali arranged in multiple crowns before septa of all but last cycle; axial edge
of septa minutely dentate...........................................................................22
21b Pali arranged in two crowns before S2 and S3 or S1-3; axial edges of septa
smooth......................................................................................................24
22a Coenosteum costate..................[Atl. + Pac.] Cladocora Plate 3, Figures K–L
22b Coenosteum not costate.............................................................................23
23a Axial corallite associated with each branch.....................................................
................................................ [SW Pac.] Petrophyllia Plate 4, Figures A–B
23b Axial corallites absent.................[Atl. + Pac.] Oculina* Plate 4, Figures C–D
24a P1-3 arranged in two palar crowns...... [IWP] Cyathelia Plate 4, Figures E–F
24b One palar crown of P2 or P3.....................................................................25
25a Only P2 present.........................................................................................26
25b Only P3 present ............. [SW Atl. + E. Pac.] Bathelia Plate 4, Figures G–H
26a Columella massive ...........................[SE Atl.] Sclerhelia Plate 4, Figures I–J

26b Columella rudimentary..................................................................................
........................... [Cosmopolitan] Madrepora (in part) Plate 4, Figures K–L
27a Septal symmetry decameral or octameral, septa in only one cycle; columella
styliform........................[Atl. + IP] Madracis* (in part) Plate 5, Figures A–B
27b Septal symmetry hexameral, septa arranged in multiple cycles; columella pap-
illose, fascicular, spongy, lamellar or absent................................................28
28a Texture of corallum rough (like sandpaper), resulting from a porous theca....29
28b Texture of corallum smooth or costate, solid.............................................31
29a Corallum increases by stoloniferous budding (reptoid), the connection
among corallites often obscured, thus sometimes appearing to be solitary;
Pourtalès Plan present... [W. Atl. + IP] Rhizopsammia Plate 5, Figures C–D
29b Corallum increases by budding from a common basal coenosteum, the con-
nection among polyps quite evident; septa normally inserted.....................30
30a Columella massive; epitheca surrounds each corallite.....................................
......................................................... [E. Atl.] Astroides Plate 5, Figures E–F
30b Columella of moderate to small size; epitheca lacking....................................
...................................[Atl. + IP] Tubastraea (in part) Plate 5, Figures G–H
31a Columella absent.......................................................................................32
31b Columella present......................................................................................33
32a Corallites united by thin basal stolons (reptoid).............................................
........................................[Atl. + IWP] Thalamophyllia Plate 5, Figures I–K
32b Corallites bud from a common basal coenosteum..........................................
...............[E. Atl. + New Zealand] Hoplangia (in part) Plate 5, Figures L–M
33a Axial edges of some or all cycles of septa finely dentate or beaded..................
.......................................................................................... (Rhizangiidae) 34
33b Axial edges of all septa smooth...................................................................38
34a Thin epitheca encircles corallites; axial edges of S1-2 smooth, sometimes lo-
bate (but inner edges of S3-4 dentate)........................................................35
34b Epitheca absent; axial edges of all septa dentate..........................................36
35a Corallite base polycyclic; one crown of large P3.............................................
....................................................[Atl.+ Pac.] Colangia Plate 6, Figures A–B
35b Corallite base monocyclic; pali, if present, of uniform size.............................
................................................................. [IP] Culicia Plate 6, Figures C–D
36a Corallite base polycyclic; pali absent....... [IP] Oulangia Plate 6, Figures E–F
36b Corallite base monocyclic; pali before septa of all but last cycle..................37
37a Corallum stoloniferous (reptoid) or cerioid; peritheca absent.........................
..................................................[Atl. + IP] Astrangia* Plate 6, Figures G–H
37b Corallum massive (subramose); peritheca unite corallites...............................
.........................................................[Indian] Cladangia Plate 6, Figures I–J
38a Pali or paliform lobes on axial edges of septal of all but last cycle...............39
38b Pali or paliform lobes present only on septa of penultimate cycle (usually
P3).............................................................................................................41
39a Corallum stoloniferous (reptoid)....[IWP] Rhizosmilia Plate 6, Figures K–L

39b Corallites bud from a common basal coenosteum......................................40
40a Corallites monocyclic; pali before septa of all but last cycle, and all of ap-
proximately the same size................ [IWP] Polycyathus Plate 7, Figures A–B
40b Corallites polycyclic; pali before septa of all but last cycle, those of P3 crown
much larger than others....... [W. Atl.] Phacelocyathus Plate 7, Figures C–D
41a Columella fascicular...................................................................................42
41b Columella trabecular................... [Atl. + IP] Phyllangia Plate 7, Figures E–F
42a Occurrence of pali variable: usually P4, occasionally also P3, occasionally
absent...................................... [E. Pac.] Bathycyathus Plate 7, Figures G–H
42b Pali in one crown before septa of third cycle (P3)..........................................
............................................. [Atl. + Pac.] Coenocyathus Plate 7, Figures I–J
43a Corallum firmly attached (fixed)................................................................44
43b Corallum unattached (free)........................................................................67
44a Theca granular, the granules usually occurring on longitudinally oriented
costae..................................................................................................... 45
44b Theca smooth (epithecate or stereome-reinforced), sometimes with fine
transverse ridges encircling the theca..........................................................53
44c Theca and septa porous, although in some genera a smooth epitheca may
cover the basal portion of the corallum......................................................61
44d Theca absent (corallum discoidal)......[E. Pac.] Nomlandia Plate 7, Figure K
45a Columella papillose...................................................................................46
45b Columella fascicular...................................................................................51
45c Columella absent.......................................................................................52
45d Columella labyrinthiform.... [Atl. + IP] Labyrinthocyathus Plate 8, Figures A–B
46a Pali or paliform lobes absent; base polycyclic.................................................
.................................... [W. Atl. + W. Pac.] Oxysmilia Plate 8, Figures C–D
46b Pali or paliform lobes present; base monocyclic.........................................47
47a Coralla usually arranged in pseudocolonial assemblages.................................
........................................[W. Pac.] Lochmaeotrochus Plate 8, Figures G–H
47b Coralla discrete..........................................................................................48
48a Pali before S1-2 (P1, P2), indistinguishable from columellar elements...........
...............................[W. Atl. + IWP] Monohedotrochus Plate 8, Figures E–F
48b Pali before septa of all but last cycle; palar crowns discrete.........................49
49a Multiple slender paliform lobes on axial edge of every lower cycle septum, not
arranged in crowns..................... [Atl. + IP] Paracyathus Plate 8, Figures I–J
49b Two crowns of discrete pali or paliform lobes (P1+P2 and P3), only one palus
or paliform lobe per septum.......................................................................50
50a True pali present, the P1-2 smaller than P3 but not significantly...................
....[Atl. + IP] Trochocyathus (Trochocyathus) (in part) Plate 8, Figures K–L
50b Paliform lobes present, the P1-2 much smaller than the broad P3.................
.................................. [W. Atl. + W. Pac.] Vaughanella Plate 9, Figures A–B
51a Pali before septa of penultimate cycle.............................................................

.[Cosmopolitan] Caryophyllia (Caryophyllia) (in part) Plate 9, Figures C–D
51b Pali absent................. [Cosmopolitan] Crispatotrochus Plate 9, Figures E–F
52a Corallum base monocentric; epitheca lacking; calice elliptical in outline; me-
nianes lacking............ [Cosmopolitan] Desmophyllum Plate 9, Figures G–H
52b Corallum polycentric; transverse epithecal bands near corallum base; calicu-
lar outline modified by calicular extensions; menianes on septal faces............
................................................[W. Pac.] Dactylotrochus Plate 9, Figures I–J
53a Columella absent or simply a rudimentary fusion of lower axial edges of ma-
jor septa deep in fossa................................................................................54
53b Columella present (papillose, fascicular or labyrinthiform)........................57
54a Pedicel reinforced (thickened) with stereome deposits...................................
...............................................[Cosmopolitan] Javania Plate 9, Figures K–L
54b Pedicel reinforced with hollow rootlets, most easily seen in cross section of
base or pedicel, or in a damaged corallum..................................................55
55a Rootlets non-contiguous with pedicel, 2-20 adventitious rootlets anchoring
the corallum............................... [IWP] Rhizotrochus Plate 10, Figures A–B
55b Rootlets (symmetrical or asymmetrical in placement) contiguous with pedi-
cel, forming an integral part of the lower corallum.....................................56
56a Calicular edge jagged...........[W. Atl. + IP] Polymyces Plate 10, Figures C–D
56b Calicular edge smooth....[E. Atl. + W. Pac.] Monomyces Plate 10, Figures E–F
57b Columella fascicular...................................................................................60
57c Columella labyrinthiform.... [W. Pac.] Stolarskicyathus Plate 10, Figures G–I
58a Corallum base polycyclic; no notch between upper outer edges of septa and
theca..........................................................................................................59
58b Base monocyclic, but may have an accessory basal rootlet; septal notch pre-
sent................................. [W. Atl. + IWP] Gardineria Plate 10, Figures J–K
59a Pali before septa of penultimate cycle.............................................................
................................. [Atl. + E. Pac.] Concentrotheca Plate 10, Figures L–M
59b Paliform lobes present before septa of S1-2 (P1-2).........................................
................................ [E. Atl. + E. Pac.] Ceratotrochus Plate 11, Figures A–B
59c Pali before septa of all but last cycle in two crowns........................................
......................................... [Atl. + Pac.] Tethocyathus Plate 11, Figures C–D
60a Corallum cylindrical and very small (calicular diameter less than 2 mm); a
row of thecal spots or pores present in every interseptal region; octameral
septal symmetry; only 1 columellar element...................................................
..................................................[Atl. + IWP] Guynia Plate 11, Figures E–G
60b Corallum trochoid and larger (adult calicular diameter over 10 mm); the-
cal spots and pores lacking; hexameral symmetry; numerous columellar ele-
ments............................. [IWP] Conotrochus (in part) Plate 11, Figures H–I
61a Septa arranged in a Pourtalès Plan.............................................................62
61b Septa arranged normally............................................................................63

62a Corallum base polycyclic; theca costate..........................................................
....... [Cosmopolitan] Balanophyllia (Balanophyllia)* Plate 11, Figures J–K
62b Corallum base monocyclic; theca hispid (not costate)....................................
.......................... [W. Atl. + SW Pac.] Thecopsammia Plate 11, Figures L–M
63a Columella absent or rudimentary...............................................................64
63b Columella spongy......................................................................................65
64a Corallum trochoid; theca costate...................................................................
.......................................... [W. Atl.] Trochopsammia Plate 12, Figures A–B
64b Corallum subcylindrical (sometimes scolecoid); theca uniformly hispid (not
costate)........................ [S. Africa] Pourtalopsammia Plate 12, Figures C–D
65a Costae absent; axial edges of all septa smooth; no endothecal dissepiments....
.............................................[W. Atl.] Bathypsammia Plate 12, Figures E–F
65b Costae granular or hispid; axial edges of higher cycle septa dentate to lacini-
ate; endothecal dissepiments present in an elongate corallum.....................66
66a Columella not discrete (merging with lower axial edges of septa); costae
weakly granular …...................... [IP] Endopsammia Plate 12, Figures G–H
66b Columella discrete; costae hispid...................................................................
.................................... [E. Atl. + IWP] Leptopsammia Plate 12, Figures I–J
67a Corallum unattached (free) in every growth stage (lacking transverse divi-
sion)..........................................................................................................68
67b Corallum undergoes transverse division, resulting in a free anthocyathus stage
with a basal scar, but with a fixed anthocaulus stage.................................102
68a Corallum conical (ceratoid, trochoid or turbinate).....................................69
68b Corallum bowl-shaped...............................................................................87
68c Corallum cupolate (theca horizontal with no surrounding vertical theca).....91
68d Corallum cuneiform............................................ (Turbinoliidae, in part) 97
68e Corallum globular (pores in base of corallum associated with commensal si-
punculid).................................................................................................101
68f Corallum cylindrical.... [Atl. + IWP + Ant.] Stenocyathus Plate 12, Figures K–L
69b Columella rudimentary or absent...............................................................78
69c Columella fascicular...................................................................................83
69d Columella trabecular..................................................................................86
69e Columella styliform................. [SW Pac.] Turbinolia Plate 13, Figures A–B
69f Columella spongy..........................................................................................
.......... [W. Atl. + IWP] Balanophyllia (Eupsammia) Plate 13, Figures C–D
70a Pali or paliform lobes present.....................................................................71
70b Pali and paliform lobes absent. [IWP] Foveolocyathus Plate 13, Figures E–F
71a Pali before septa of second cycle (P2).........................................................72
71b Pali or paliform lobes before septa of all but last cycle................................77
71c Pali or paliform lobes before septa of third cycle (P3)....................................
.................... E. Atl. + IWP + Ant.] Paraconotrochus Plate 13, Figures G–H
72a Theca bears numerous linear rows of spots, pits or thecal perforations.......73
72b Theca solid, not bearing spots, pits or perforations.....................................75
73a Theca perforate..... [W. Atl. + W. Pac.] Trematotrochus Plate 13, Figures I–J
73b Theca bears linearly arranged spots or pits..................................................74
74a A row of pits occurs in each interseptal space on inner theca; costae granular.
......................................... [W. Pac.] Endocyathopora Plate 13, Figures K–L
74b A row of white spots occurs in each interseptal space on outer theca; theca
smooth (epithecate) or covered with hispid spines.........................................
.......................................... [W. Atl.] Pourtalocyathus Plate 14, Figures A–B
75a Theca bears serrate costae...........................................................................76
75b Theca smooth (epithecate).....[SW Pac.] Lissotrochus Plate 14, Figures C–D
76a Theca covered with twice as many costae as septa...........................................
................................................ [SW Pac.] Pleotrochus Plate 14, Figures E–F
76b Costae and septa of equal number.................................................................
.............................[W. Atl. + W. Pac.] Cryptotrochus Plate 14, Figures G–H
77a Pali discrete, pairs of P3 fused into chevrons within each system; no parricidal
budding..................................... [W. Pac.] Notocyathus Plate 14, Figures I–J
77b Multiple paliform lobes on all septa; parricidal budding common.................
............................................. [IWP] Thrypticotrochus Plate 14, Figures K–L
78a Theca smooth (epithecate), costae not present...........................................79
78b Theca granular, costae present (twice the number of septa)........................82
79a Rows of thecal spots visible on theca..........................................................80
79b Thecal spots lacking...................................................................................81
80a Twelve contiguous rootlets present in pedicel; parricidal budding absent.......
.........................................[W. Pac.] Pedicellocyathus Plate 15, Figures A–C
80b Rootlets lacking; parricidal budding from parent fragment common.............
...................................................[Atl.] Schizocyathus Plate 15, Figures D–E
81a Calicular edge smooth...................................................................................
.....................[Cosmopolitan] Flabellum (Flabellum) Plate 15, Figures F–G
81b Calicular edge jagged.....................................................................................
....................[Cosmopolitan] Flabellum (Ulocyathus) Plate 15, Figures H–I
82a Theca perforate; septa hexamerally arranged in 3 or 4 cycles..........................
.................................................... [IWP] Conocyathus Plate 15, Figures J–K
82b Theca imperforate; only10 septa (6+4)...........................................................
............................................[SW. Pac.] Holcotrochus Plate 15, Figures L–M
83a Pali before septa of penultimate cycle (usually P3).....................................84
83b Pali absent......................[E. Pac.] Pseudocyathoceras Plate 16, Figures A–B
84a Thecal edge spines or crests present............................................................85
84b Thecal edge spines and crests absent.................................................................
.[Cosmopolitan] Caryophyllia (Caryophyllia)(in part) Plate 16, Figures C–D
85a Base of corallum usually open, as though broken from parent through asexual
budding........................... [Atl. + IWP] Premocyathus Plate 16, Figures E–F
85b Base of corallum intact..................................................................................

................ [W. Pac.] Caryophyllia (Acanthocyathus) Plate 16, Figures G–H
86a Theca costate; septal notch absent..................................................................
........................................... [Atl. + IWP] Dasmosmilia Plate 16, Figures I–J
86b Theca smooth; septal notch present...............................................................
.......................... [E. Atl. + IWP + Ant.] Aulocyathus Plate 16, Figures K–M
87a Paliform lobes on septa of all cycles; septal edges smooth...........................88
87b Pali before septa of all but last cycle; septal edges smooth..........................90
87c Pali before septa of third cycle (P3); septal edges smooth...............................
.............................................. [W. Pac.] Ericiocyathus Plate 17, Figures A–B
87d Pali and paliform lobes absent; septal edges coarsely dentate..........................
................. [W. Atl. + IWP] Anthemiphyllia (in part) Plate 17, Figures C–D
88a Lower outer edge of corallum bears tubercles or spines on the C1 or C1-2.89
88b Tubercles and spines absent...........................................................................
.....[Atl. + IWP] Stephanocyathus (Stephanocyathus) Plate 17, Figures E–F
89a Six long C1 spines on lower outer edge of corallum.......................................
...................[IWP] Stephanocyathus (Acinocyathus) Plate 17, Figures G–H
89b Twelve to 18 short spines or tubercles on lower outer edge of corallum ........
.... [W. Atl. + IWP] Stephanocyathus (Odontocyathus) Plate 17, Figures I–J
90a Six C1 spines on lower outer edge of corallum...............................................
....................[W. Pac.] Trochocyathus (Aplocyathus) Plate 17, Figures K–M
90b Costal spines absent....... [Atl. +Pac.] Deltocyathoides Plate 18, Figures A–C
91a Costae alternate in position with septa; higher cycle septa increase by bifurca-
tion; thecal base perforate.................................................(Micrabaciidae) 92
91b Costae continuous with septa; higher cycle septa increase by adding addi-
tional cycles; base imperforate....................................................................95
92a Septa rudimentary, composed of a series of tall spines....................................
.......................................[Cosmopolitan] Leptopenus Plate 18, Figures D–E
92b Septa lamellar............................................................................................93
93a Marginal shelf present; columella spongy...................................................94
93b Marginal shelf absent; columella solid............................................................
............................................. [IWP] Stephanophyllia Plate 18, Figures F–H
94a Septa highly porous.................... [IWP] Letepsammia Plate 18, Figures I–K
94b Septa essentially imperforate, porous only at points at which septa bifurcate..
............................................[IWP] Rhombopsammia Plate 18, Figures L–N
95a Synapticular platelets absent; corallum robust; upper septal edges smooth.....
............................................[Atl. + IP] Deltocyathus Plate 18, Figures O–Q
95b Synapticular platelets brace adjacent septa; corallum fragile; upper septal edg-
es bear slender elongate spines...................................................................96
96a Corallum with five cycles of septa (96 septa)...................................................
.[Atl. + IWP. + Ant.] Fungiacyathus (Fungiacyathus) Plate 19, Figures A–C
96b Corallum with four cycles of septa (48 septa).................................................
............. [Cosmopolitan] Fungiacyathus (Bathyactis) Plate 19, Figures D–F
97a Thecal edge crests present..........................................................................98

97b Thecal edge crests absent..........................................................................100
98a Pali absent.................................................................................................99
98b Pali present, before septa of all but last cycle..................................................
..............................................[IWP] Tropidocyathus Plate 19, Figures G–H
99a Twice as many costae as septa....[W. Pac.] Alatotrochus Plate 19, Figures I–J
99b Equal number of costae and septa....[IWP] Platytrochus Plate 19, Figures K–M
100a Columella lamellar; pali absent......................................................................
...................................... [Atl. + Pac.] Sphenotrochus Plate 19, Figures N–O
100b Columella papillose; pali before septa of all but last cycle...............................
.................................................[IWP] Cyathotrochus Plate 20, Figures A–B
101a Theca imperforate (although septa may be perforate).....................................
................................ [IWP + W. Atl.] Heterocyathus* Plate 20, Figures C–E
101b Theca and septa perforate...............................................................................
.............................. [IWP] Heteropsammia* (in part) Plate 20, Figures F–H
102a Columella papillose.................................................................................103
102b Columella spongy....................................................................................108
102c Columella a solid fusion in center of calice..............................................109
102d Columella absent.....................................................................................110
102e Columella fascicular.................................................................................111
102f Columella lamellar.........................[IWP] Placotrochus Plate 20, Figures I–J
102g Columella trabecular......[Atl. + IWP] Placotrochides Plate 20, Figures N–O
103a Pali before septa of all but last cycle.........................................................104
103b Pali before S1-2 (P1-2)............................................................................106
103c Pali before S2 (P2)................[SW Pac.] Kionotrochus Plate 21, Figures A–B
103d Pali absent.[W. Atl. + IWP] Anthemiphyllia (in part) Plate 21, Figures C–E
104a Six long C1 spines on lower outer edge of corallum.......................................
..........................................[W. Pac.] Bourneotrochus Plate 21, Figures F–H
104b Thecal spines absent, although corallum may bear thecal edge crests........105
105a Corallum small (calicular diameter usually less than 5 mm); higher cycle sep-
ta bend toward and fuse with adjacent lower cycle septa................................
...................................[Atl. + W. Pac.] Peponocyathus Plate 21, Figures I–K
105b Corallum larger (calicular diameter usually over 10 mm); higher cycle septa
independent...................................................................................................
...[W. Pac.] Trochocyathus (Trochocyathus) (in part) Plate 21, Figures L–N
106a Septa alternate in position with costae; thecal spots absent.......................107
106b Septa correspond to costae; lines of thecal spots present.................................
............................................ [W. Pac.] Temnotrochus Plate 21, Figures O–P
107a Corallum cuneiform in shape, sometimes with basal thecal spurs (fish-tail
morphology)............................. [W. Pac.] Idiotrochus Plate 22, Figures A–C
107b Corallum (anthocyathus) discoidal to bowl-shaped, without thecal spurs......
..............................................[W. Pac.] Dunocyathus Plate 22, Figures D–E
108a Septa arranged in a Pourtalès Plan; thecal edges often crested........................

.........................................................[IP] Endopachys Plate 22, Figures F–G
108b Septa normally arranged; thecal edges rounded..............................................
................................................[SW Pac.] Notophyllia Plate 22, Figures H–J
109a Multiple paliform lobes on septa of all but last cycle; thecal spots absent; cor-
allum discoidal to cupolate in shape...............................................................
............................................[Indian] Australocyathus Plate 22, Figures K–L
109b Pali and paliform lobes absent; linear rows of thecal spots present; corallum
compressed-cylindrical.....[W. Pac.] Truncatoguynia Plate 22, Figures M–N
110a Buds propagate from thecal edges of corallum...............................................
............................................ [W. Pac.] Blastrotrochus Plate 23, Figures A–B
110b No budding from thecal edges.......................................................................
..................[E. Atl. +IWP + Ant.] Truncatoflabellum Plate 23, Figures C–F
111a Pali before third cycle septa; theca costate.........................................................
.[Cosmopolitan] Caryophyllia (Caryophyllia) (in part) Plate 23, Figures G–H
111b Paliform lobes before second cycle septa; epithecate.......................................
...................................... [SW Pac.] Falcatoflabellum Plate 20, Figures K-M
Acknowledgements
We wish to thank Anna Maria Addamo for showing us the need for writing such a
paper, and Helmut Zibrowius for supplying images of the genus Cladangia. MVK
is also thankful to Philippe Bouchet, Bertrand Richer de Forges, and MNHN and
IRD- Nouméa staff and collaborators for their great effort in collecting and preserving
deep-water scleractinians used to illustrate many genera in the present study. MVK is
supported by the São Paulo Research Foundation (FAPESP).
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Glossary
Anthocaulus: See Transverse Division.
Anthocyathus: See Transverse Division.
Apozooxanthellate: Species that have facultative symbiotic relationships with unicel-
lular photosynthetic dinoflagellates (Symbiodinium spp.).
Axial Corallite: See Corallite.
Axial Septal Margin: See Septum.
Azooxanthellate: Species that do not have symbiotic relationships with unicellular
photosynthetic dinoflagellates (Symbiodinium spp.).
Base (Figure 1): The lower several millimeters of a solitary corallum, usually solid and
composed of an accretion of thecal layers (a monocyclic base) (e.g. Plate 24, Fig.
B), but in some genera composed of concentric rings of partitioned chambers,
called a polycyclic base (e.g. Plate 24, Fig. C) (Cairns 1978). The base usually
expands basally as a thin layer facilitating adhesion to the substrate.
Budding: The process of asexual reproduction that adds new mouths (or polyps) to a
corallum, often resulting in a colony. Intratentacular budding adds new polyps
to the oral disc inside the ring of tentacles surrounds its mouth (e.g. Plate 1, Fig.
K–L). Extratentacular budding adds new polyps outside the ring of tentacles
(e.g. Plate 2, Fig. H). (The third form of asexual reproduction is transverse divi-
sion - Cairns 1988b).
Calice (Figure 1) (pl. Calices): The skeletal analog of the polyp, cupping the polyp
from below, and consisting of the septa, and, if present, the columella and pali
(e.g. Plate 1, Figs C, E, G; Plate 9, Figs A, C, E, G, I, K).
Ceratoid Corallum: See Solitary Corallum.
Coenosteum: The skeletal structure found between the individual corallites of a colo-
nial corallum, including the costae, and various kinds of dissepiments; sometimes
called peritheca (e.g. Plate 24, Fig. E).
Colonial Corallum: See Corallum.
Columella (Figure 1): An axial structure of diverse shape and composition that pro-
jects from the center of a calice. If in the shape of a single lamella (called lamellar)
(e.g. Plate 20, Fig. I), if a maze of interconnected lamellae (labyrinthiform) (e.g.
Plate 8, Fig. A), if a set of twisted lamellae (fascicular) (e.g. Plate 20, Figs C, E),
if a simple rod (styliform) (e.g. Plate 13, Fig. A), if a group of rods (papillose)
(e.g. Plate 8, Fig. K), if a fine porous mass (spongy) (e.g. Plate 12, Fig. G), and
if an irregular group of twisted elements (trabecular) (e.g. Plate 16, Fig. K).
Conical Corallum: See Solitary Corallum.
Corallite: The vertical, usually cylindrical, structure produced by an individual polyp,
consisting of endothecal dissepiments and the calice at the upper end (e.g. Plate
1, Fig. F). If a corallite occurs at the tip of a colony’s branch, it is termed an axial
corallite (e.g. Plate 24, Fig. F).
Corallum (Figure 1) (pl. Coralla): The aragonitic calcium carbonate skeleton of a scle-
ractinian coral. If the coral has only one mouth (or calice), it is termed solitary (e.g.
Plate 10, Figs A–M; Plate 17, Figs A–M), if polystomatous (or more than one
calice), then a colonial (e.g. Plate 2, Fig. A–L; Plate 3, Figs A–L).
Costae (Figure 1) (sing. Costa; adj. Costate): Continuation of a septum on the out-
side of the corallite wall, often as a ridge or low linear mound (e.g. Plate 24, Figs
A, D, G, M).
Crest: See Edge Spine.
Crown (Figure 1): See Palus.
Cuneiform Corallum: See Solitary Corallum.
Cupolate Corallum: See Solitary Corallum.
Cycle: See Septum.
Cylindrical Corallum: See Solitary Corallum.
Discoidal Corallum: See Solitary Corallum.
Dissepiments: Thin horizontal (tabular dissepiments, e.g., Plate 2, Fig. K) or blis-
ter-like plates that form within a corallite (endothecal) or beneath the coenosteum
outside corallites (exothecal), which separate the polyp from the lower part of the
corallum that it no longer occupies.
Distomodeal Budding: A mode of intratentacular budding in which two polyps (or
calices) develop within the common tentacular ring (e.g., Plate 1, Figs I–K).
Edge Spine/Crest/Spur: The external thecal edges of a solitary coral, those associ-
ated with the principal septa, sometimes bears a low thin crest, or a series of
hollow spines. If the crest is limited to the basal portion of the corallum and
project outward in the shape of a fish tail, they may be called spurs (e.g. Plate
24, Figs H–M).
Edge Zone: The fold of the polyp body that extends over the edge of the theca (e.g.
Plate 25, Fig. A).
Endothecal: See Dissepiments.
Epitheca: Thin, external, smooth or wrinkled, non-trabecular sheath surrounding in-
dividual corallites, formed by centripetal (inward) growth (e.g. Plate 10, Figs I,
K; Plate 11, Figs D, I; Plate 12, Fig. F). Tectura is very similar in outward ap-
pearance by originates by centrifugal (outward) growth (e.g. Plate 25, Fig. B)
(Stolarski 1995).
Exothecal: See Dissepiments.
Extratentacular Budding: See Budding.
Fascicular Columella: See Columella.
Free: An unattached corallum (e.g. Plate 14, Figs A–L; Plate 17, Figs A–M; Plate
18, Figs A–Q).
Globular Corallum: See Solitary Corallum.
Imperforate Theca: See Theca.
Intratentacular Budding: See Budding.
Labyrinthiform Columella: See Columella.
Lamellar Columella: See Columella.
Marginal Shelf: A low rim encircling a solitary corallum composed of greatly reduced
septa and costae, or costal spines (Cairns 1989) (e.g. Plate 25, Fig. D).
Menianes: Short ledge-like features on septal faces formed by aligned lateral exten-
sions of trabeculae (e.g. Plate 25, Fig. C).
Monocyclic Base: See Base.
Monostomaeous/Monostomatous: A single-mouthed corallum, i.e., a solitary form
(e.g. Plate 8, Figs A–F).
Normal Arrangement of Septa: Arrangement of septa within a calice in which the
septa are independent and all aligned with the center of the calice. (See Pourtalès
Plan) (e.g. Plate 8, Fig. E; Plate 9, Fig. E).
Paliform Lobes: Small, flattened lobes on the axial septal edge of various cycles, often
more than one per septum, and part of the septum to which they are attached
(e.g., Plate 17, Fig. I).
Palus (Figure 1) (pl. Pali): Small flattened lobes on the axial septal edge of various
cycles, always one per septum, and not part of the septum to which it is attached
but ontogenetically different. Groups of pali occurring on the same cycle of septa
and thus stand at the same distance from the center of the calice are called crowns
of pali (e.g. Figure 1).
Papillose Columella: See Columella.
Parricidal Budding: A mode of intratentacular budding in which new polyps are
generated from the inner surface of a fragment of a parent corallum that has lon-
gitudinally split apart (e.g. Plate 16, Fig. J, L–M).
Pedicel (Figure 1): The stem-like region of a solitary coral just above the base and
below the calicular surface.
Perforate Theca: See Theca.
Peritheca: See Coenosteum.
Polycyclic Base: See Base.
Polystomaeous/Polystomatous: See Corallum.
Pourtalès Plan: A form of septal arrangement in which the axial edges of pairs of
higher cycle septa bend in front of and unite before their adjacent lower cycle sep-
tum (e.g. Plate 25, Fig. E). See Cairns (1994: fig. 2).
Reptoid Budding: A type of extratentacular budding in which polyps are asexually
generated from a thin, reticulate, encrusting ribbon (similar to stoloniferous bud-
ding) (e.g. Plate 6, Fig. B; Plate 7, Fig. F).
Scolecoid Corallum: See Solitary Corallum.
Septum (pl. Septa): Radially arranged longitudinal partitions of a corallite (Figure 1),
usually arranged in hexameral symmetry. Septa are added in cycles, the first cycle
composed of 6 septa, the second also of 6, the third of 12, the fourth of 24, the
fifth of 48, etc. resulting in corallites consisting of 6, 12, 24, 48, or 96, etc. septa
per calice. Septa can bear smooth, dentate, or laciniate axial margin (Figure 1).
Solitary Corallum: Solitary coralla exist in a variety of shapes, the shape being one of
the criteria used to differentiate genera and species. Many solitary coralla are shaped
as an inverted cone (conical), and may be attached and straight or free and usually
curved. If the edges of the cone diverge at a hypothetical basal angle of 10–40°, this
corallum is called ceratoid (e.g. Plate 10, Figs H–I), if the angle is 40–60°, then
trochoid (e.g. Plate 8, Fig. F), if the angle is 60–80°, turbinate (e.g. Plate 13, Fig.
H), and if the angle is 160–180° and the corallum is low, discoidal (e.g. Plate 18,
Fig. P). Coralla may also be cylindrical (e.g. Plate 22, Fig. N), and if the cylinder
is irregular in shape, scolecoid (e.g. Plate 12, Fig. D). Others are wedge-shaped
(cuneiform) or bowl-shaped (e.g. Plate 17, Figs F, H, J, M). Still others have a
flat base with a convex upper surface (cupolate) (e.g. Plate 19, Figs A–F) and oth-
ers are simply onion-shaped or irregular (globular) (e.g. Plate 18, Fig. B).
Spongy Columella: See Columella.
Stereome: A general term for thick calcareous deposits, generally thickening various
parts of the corallum.
Stoloniferous Budding: A type of extratentacular budding in which polyps are asexu-
ally generated from a thin, elongate, encrusting coenenchymal ribbon, the con-
necting ribbon often obscured by encrusting organisms (e.g. Plate 5, Fig. B; Plate
6, Fig. D).
Styliform Columella: See Columella.
Synapticular Plate: Ribbons of calcium carbonate linking adjacent fungiacyathid sep-
ta, first appearing as vertical rods midway between septa, later bifurcate, the two
ends fusing to adjacent septal faces (T- or Y-shaped) (Cairns 1989a) (e.g. Plate
25, Fig. F).
Trabecular Columella: See Columella.
Tabular Endothecal Dissepiment: See Dissepiment.
Tectura: See Epitheca.
Theca (Figure 1): The skeletal sides, or walls, of solitary coralla or corallites of colonial
corals, that enclose the polyps. If the theca is solid, it is termed imperforate (e.g.
Plate 9, Figs D, L); if the theca is porous, perforate (e.g. Plate 22, Figs I–J).
Thecal spots/pores: Some genera have longitudinal series of small pores aligned with
the interseptal spaces of various cycles, termed thecal pores. In other genera, these
analogous structures do not penetrate the theca but are visible only as slightly dif-
ferently colored spots of a constructional consistently different from the remaining
theca, these termed spots (e.g. Plate 11, Figs F–G; Plate 22, Fig. N).
Thecal spur: See Edge Spine.
Trabecular Columella: See Columella.
Transverse Division: One of the three main methods of asexual reproduction among
the Scleractinia (Cairns 1988b). In this method an attached sexually produced
solitary corallum (called the anthocaulus - e.g. Plate 25, Figs H, J) transversely
divides it corallum usually along a crescent-shaped line of thecal weakness, pro-
ducing an asexually generated distal corallum (the anthocyathus - e.g. Plate 25,
Figs G, I), which falls to the substrate as an unattached (free) corallum, the base of
which retains a scar of previous attachment (e.g. Plate 25, Figs I–J).
Trochoid Corallum: See Solitary Corallum.
Turbinate Corallum: See Solitary Corallum.
Zooxanthellate: Species that have symbiotic relationships with unicellular photosyn-
thetic dinoflagellates (Symbiodinium spp.).
Figure and plates
Figure 1. Cutaway diagram of a species of Caryophyllia illustrating the basic morphological features of an
attached, solitary scleractinian (Modified from Cairns 1994). Small circular photos are from different sclerac-
tinian species and are intended to illustrate basic morphological characters used in the taxonomy of the group.
Plate 1. Heteropsammia cochlea A (USNM 97652) and B (USNM 73772): Calicular and lateral view
respectively; Anomocora gigas (MNHN uncatalogued, Terrasses stn. CP3091) C and D Calicular and
lateral view respectively; Eguchipsammia fistula (USNM uncatalogued, Norfolk 2 stn. 2024) E and F Cali-
cular and colony view respectively; Lophelia pertusa (USNM 1071877) G and H Calicular and colony
view respectively; Dichopsammia granulosa (USNM 15847, holotype) I and J Calicular and colony view
respectively; Solenosmilia variabilis (USNM 47426) K and L Distal branch and colony view respective-
ly. Scale bars: blue = 1 mm; white = 5 mm; green = 50 mm. Bold face indicates type species for the genus.
Plate 2. Madracis asperula A (SEM, USNM 99068) and B (USNM 99056): Calicular and colony
view respectively; Cladopsammia sp. (USNM uncatalogued, Norfolk 2 stn. 2023) C and D Calicular
and colony view respectively; Dendrophyllia alcocki (USNM uncatalogued, Norfolk 2 stn. 2135) E and
F Calicular and colony view respectively; Enallopsammia rostrata (USNM uncatalogued, Norfolk 2 stn.
DW 2056) G and H Calicular and colony (calicular and acalicular side) view respectively; Tubastraea
coccinea (USNM 46973) I Calicular view; Tubastraea diaphana (USNM 83677) J Colony view; Gonio-
corella dumosa (USNM 47505) K and L Calicular and colony view respectively. Scale bars: blue = 1 mm;
white = 5 mm; green = 50 mm. Bold face indicates type species for the genus.
Plate 3. Pourtalosmilia anthophyllites A (USNM 1174947) and B (USNM 117494): Calicular and
colony view respectively; Confluphyllia juncta (USNM 97316, paratype) C and D Calicular and colony
view respectively; Coenosmilia arbuscula (USNM 97312) E and F Calicular and colony view respectively;
Sympodangia albatrossi (USNM 97308, holotype) G (SEM) and H Calicular and colony view respectively;
Madrepora oculata (MNHN uncatalogued, Halipro 2 stn. BT104) I and J Calicular and colony view respec-
tively; Cladocora debilis K (USNM 10452, SEM) and L (USNM 62351): Calicular and colony view respec-
tively. Scale bars: blue = 1 mm; white = 5 mm; green = 50 mm. Bold face indicates type species for the genus.
Plate 4. Petrophyllia rediviva (USNM 82696) A and B Distal branch and colony view respectively;
Oculina virgosa (MNHN uncatalogued, SMIB 5 stn. DW101) C and D Calicular and colony view re-
spectively; Cyathelia axillaris (USNM 92665) E and F Calicular and colony view respectively; Bathelia
candida (USNM 47512) G and H Calicular and colony view respectively; Sclerhelia hirtella (MNHN
Michellin collection) I and J Calicular and colony view respectively; Madrepora minutiseptum (MNHN
uncatalogued, SMIB 5 stn. DW101) K and L Calicular and colony view respectively. Scale bars: blue = 1
mm; white = 5 mm; green = 50 mm. Bold face indicates type species for the genus.
Plate 5. Madracis pharensis A (USNM 96676, SEM): Colony view; Madracis sp. B (MNHN uncata-
logued, New Caledonia): Colony view; Rhizopsammia sp. (USNM uncatalogued, Bathus 4 stn. DW
941) C and D Calicular and colony view respectively; Astroides calycularis (USNM 78767) E and
F Calice detail and colony view respectively; Tubastraea coccinea (USNM 46973) G and H Calicular
and colony view respectively; Thalamophyllia tenuescens I (MNHN uncatalogued, Bathus 3 stn. CH802),
J and K (Norfolk 2 stn. 2095): Calicular and corallum views respectively; Hoplangia durotrix (AU 6097)
L (SEM) and M (SEM): Calicular and lateral view respectively. Scale bars: blue = 1 mm; white = 5 mm;
green = 50 mm. Bold face indicates type species for the genus.
Plate 6. Colangia immersa (USNM 73917) A and B Calicular and colony view respectively; Culicia
stellata (MNHN uncatalogued, New Caledonia) C and D Calicular and stolon connection view respec-
tively; Oulangia bradleyi (USNM 92371) E and F Calicular and lateral view respectively; Astrangia danae
(USNM 78507, SEM) G Calicular view; Astrangia poculata (USNM 80350, neotype) G and H and
colony view respectively; Cladangia exusta (YPM 1359, syntype ?) I and J Calicular and colony view re-
spectively; Rhizosmilia sagamiensis (USNM uncatalogued, Norfolk 2 stn. 2124) K and L Calicular and lat-
eral view respectively. Scale bars: blue = 1 mm; white = 5 mm. Bold face indicates type species for the genus.
Plate 7. Polycyathus sp. (MNHN uncatalogued, Beryx 11 stn. DW11) A and B Calicular and colony
view respectively; Phacelocyathus flos (USNM 46077) C and D Calicular and colony view respectively;
Phyllangia americana (USNM 80881) E and F Calicular and colony view respectively; Bathycyathus
chilensis (USNM 100711) G and H Calicular and colony view respectively; Coenocyathus anthophyl-
lites (USNM 48694) I and J Calicular and colony view respectively; Nomlandia californica (SBMNH
35560, holotype) K Calicular view (after Cairns 1994). Scale bars: blue = 1 mm; white = 5 mm; green =
50 mm. Bold face indicates type species for the genus.
Plate 8. Labyrinthocyathus limatulus (USNM uncatalogued, Bathus 4 stn. DW 936) A and B Calicular
and lateral view respectively; Oxysmilia corrugata (USNM uncatalogued, Norfolk 2 stn. DW2125) C and
D Calicular and lateral view respectively; Monohedotrochus circularis (USNM uncatalogued, Norfolk 2
stn. DW2124) E and F Calicular and lateral view respectively; Lochmaeotrochus oculeus (USNM un-
catalogued, Musorstom 6 stn. DW394) G and H Calicular and “aggregation” view respectively; Para-
cyathus sp. (MNHN uncatalogued, Ebisco stn. DW2555) I and J Calicular and lateral view respectively;
Trochocyathus efateensis (USNM uncatalogued, Bathus 4 stn. DW818) K and L Calicular and lateral view
respectively. Scale bars represent 5 mm. Bold face indicates type species for the genus.
Plate 9. Vaughanella concinna (USNM uncatalogued, Norfolk 2 stn. DW2070) A and B Calicular and
lateral view respectively; Caryophyllia (Caryophyllia) diomedeae (USNM uncatalogued, Norfolk 2 stn.
DW2086) C and D Calicular and lateral view respectively; Crispatotrochus rubescens (USNM uncata-
logued, Bathus 3 stn. CP833) E and F Calicular and lateral view respectively; Desmophyllum dianthus
(USNM uncatalogued, Halipro 1 stn. CP877) G and H Calicular and lateral view respectively; Dacty-
lotrochus cervicornis (USNM uncatalogued, SMIB 10 stn. DW208) I and J Calicular and lateral view
respectively; Javania insignis (USNM uncatalogued, Norfolk 2 stn. DW2023) K and L Calicular and
lateral view respectively. Scale bars represent 5 mm. Bold face indicates type species for the genus.
Plate 10. Rhizotrochus typus (USNM uncatalogued, Norfolk 2 stn. DW2024) A and B Calicular
and lateral view respectively; Polymyces wellsi C (MNHN uncatalogued, Ebisco stn. DW2618) and
D (MNHN uncatalogued, New Caledonia): Calicular and lateral view respectively; Monomyces pyg-
maea (USNM 48561) E and F Calicular and lateral view respectively; Stolarskicyathus pocilliformis
(MNHN uncatalogued, Ebisco stn. DW2573) G, H, and I Calicular, oblique, and lateral view respec-
tively; Gardineria hawaiiensis (USNM uncatalogued, Norfolk 2 stn. DW2086) J and K Calicular and
lateral view respectively; Concentrotheca laevigata (USNM 80748) L and M Calicular and oblique view
respectively. Scale bars: blue = 1 mm; white = 5 mm. Bold face indicates type species for the genus.
Plate 11. Ceratotrochus magnaghii (USNM 48780) A and B Calicular and lateral view respectively;
Tethocyathus virgatus (USNM uncatalogued, SMIB 10 stn. DW205) C and D Calicular and lateral view
respectively; Guynia annulata (MNHN uncatalogued, Biogeocal stn. DW253) E, F and G Calicu-
lar, lateral, and oblique view respectively; Conotrochus funicolumna (USNM uncatalogued, Bathus 4 stn.
CP967) H and I Calicular and lateral view respectively; Balanophyllia (Balanophyllia) laysanensis (MNHN
uncatalogued, Musorstom 6 stn. DW407) J and K Calicular and lateral view respectively; Thecopsammia
socialis (USNM 61828) L and M Calicular and lateral view respectively. Scale bars: blue = 1 mm; white
= 5 mm. Bold face indicates type species for the genus.
Plate 12. Trochopsammia infundibulum (USNM 46722) A and B Calicular and lateral view re-
spectively; Pourtalopsammia togata (USNM 91792) C and D Calicular and lateral view respectively;
Bathypsammia tintinnabulum (USNM 14569) E and F Calicular and lateral view respectively; Endop-
sammia philippensis (USNM 83006) G and H Calicular and lateral view respectively; Leptopsammia
stokesiana (USNM 78603) I and J Calicular and lateral view respectively; Stenocyathus vermiformis
(USNM uncatalogued, Norfolk 2 stn. ?) K and L Calicular and lateral view respectively. Scale bars: blue
= 1 mm; white = 5 mm. Bold face indicates type species for the genus.
Plate 13. Turbinolia stephensoni (USNM 80014) A (SEM) and B (SEM): Calicular and oblique view re-
spectively; Balanophyllia (Eupsammia) carinata (MNHN uncatalogued, Chalcal stn. D22) C and D Cali-
cular and lateral view respectively; Foveolocyathus parkeri (MNHN uncatalogued, Musorstom 5 stn. 280)
E and F Calicular and lateral view respectively; Paraconotrochus zeidleri (USNM 85677, paratype)
G and H Calicular and lateral view respectively; Trematotrochus corbicula (USNM 46477) I (SEM) and
J (SEM): Calicular and lateral view respectively; Endocyathopora laticostata (USNM 81894) K (SEM)
and L (SEM): Calicular and lateral view respectively. Scale bars: red = 0.25 mm; blue = 1 mm; white = 5
mm; green = 50 mm. Bold face indicates type species for the genus.
Plate 14. Pourtalocyathus hispidus (USNM 61928) A and B Calicular and lateral view respectively;
Lissotrochus curvatus (AM G16745) C (SEM) and D (SEM): Calicular and oblique view respectively;
Pleotrochus venustus (USNM uncatalogued, Norfolk 2 stn. DW 2104) E and F Calicular and lateral
view respectively; Cryptotrochus sp. (MNHN uncatalogued, Ebisco stn. DW2603) G and H Calicular
and oblique view respectively; Notocyathus venustus (USNM uncatalogued, Bathus 4 stn. DW 958) I and
J Calicular and lateral view respectively; Thrypticotrochus petterdi (MNHN uncatalogued, Ebisco stn.
DW2561) K and L Calicular and oblique view respectively. Scale bars: blue = 1 mm; white = 5 mm. Bold
face indicates type species for the genus.
Plate 15. Pedicellocyathus keyesi (USNM 94268, paratype) A, B (SEM), and C Calicular, pedicel
detail, and lateral view respectively; Schizocyathus fissilis (USNM 61747) D (SEM) and E (SEM):
Calicular and lateral view respectively; Flabellum (Flabellum) politum (USNM uncatalogued, Bathus 4
stn. DW933) F and G Calicular and lateral view respectively; Flabellum (Ulocyathus) messum (MNHN
uncatalogued, Bathus 1 stn. DW661) H and I Calicular and lateral view respectively; Conocyathus zelan-
diae (USNM 85713) J (SEM) and K (SEM): Oblique and lateral view respectively; Holcotrochus scriptus
(USNM 85687) L (SEM) and M (SEM): Oblique and lateral view respectively. Scale bars: blue = 1 mm;
white = 5 mm. Bold face indicates type species for the genus.
Plate 16. Pseudocyathoceras avis (USNM 46962, holotype) A and B Calicular and lateral view respec-
tively; Caryophyllia (Caryophyllia) quadragenaria (USNM uncatalogued, PrFO, New Caledonia) C and
D Calicular and oblique view respectively; Premocyathus dentiformis (MNHN uncatalogued, Ebisco
stn. DW2573) E and F Calicular and oblique view respectively; Caryophyllia (Acanthocyathus) grayi
(MNHN uncatalogued, Ebisco stn. DW2559) G and H Calicular and lateral view respectively; Dasmos-
milia lymani (USNM 82997) I and J Calicular and lateral (aggregation) view respectively; Aulocyathus
juvenescens (MNHN uncatalogued, Lifou 2000 stn. DW37) K, L, and M Calicular, lateral, and oblique
view respectively. Scale bars: blue = 1 mm; white = 5 mm. Bold face indicates type species for the genus.
Plate 17. Ericiocyathus echinatus (USNM 97169, holotype) A and B Calicular and lateral view re-
spectively; Anthemiphyllia patera costata (USNM uncatalogued, Norfolk 2 stn. 2066) C and D Calicular
and lateral view respectively; Stephanocyathus (Stephanocyathus) regius (USNM uncatalogued, Bathus 3
stn. 858) E and F Calicular and lateral view respectively; Stephanocyathus (Acinocyathus) spiniger
(USNM uncatalogued, Bathus 3 stn. CP877) G and H Calicular and lateral view respectively; Stephano-
cyathus (Odontocyathus) coronatus (USNM uncatalogued, Bathus 4 stn. CP950) I and J Calicular and
lateral view respectively; Trochocyathus (Aplocyathus) brevispina (MNHN uncatalogued, Musorstom 8 stn.
DW960) K, L, and M Calicular, oblique, and lateral view respectively. Scale bars represent 5 mm. Bold
Plate 18. Deltocyathoides orientalis (MNHN uncatalogued, Bathus 3 stn. DW829) A, B and
C Calicular, lateral, and base view respectively; Leptopenus discus (SIO Co-1271) D (SEM) and E
(SEM): Oblique views respectively; Stephanophyllia complicata (USNM uncatalogued, New Caledonia)
F, G, and H Calicular, lateral, and base view respectively; Letepsammia formosissima (USNM uncata-
logued, Norfolk 2 stn. DW2032) I, J, and K Calicular, lateral, and base view respectively; Rhombopsam-
mia niphada (USNM uncatalogued, Norfolk 2 stn. DW2069) L, M, and N Calicular, lateral, and base
view respectively; Deltocyathus rotulus (MNHN-Scl.2008-0004) O, P, and Q Calicular, lateral, and base
view respectively. Scale bars: blue = 1 mm; white = 5 mm. Bold face indicates type species for the genus.
Plate 19. Fungiacyathus (Fungiacyathus) paliferus (USNM uncatalogued, Bathus 3 stn. DW 887) A, B and
C Calicular, lateral, and base view respectively; Fungiacyathus (Bathyactis) variegatus (MNHN uncatalogued,
Lagoon NO stn. DC933) D, E, and F Calicular, lateral, and base view respectively; Tropidocyathus les-
soni (MNHN uncatalogued, Musorstom 8 stn. DW1105) G and H Calicular and lateral view respectively;
Alatotrochus rubescens (USNM uncatalogued, Bathus 4 stn. DW 908) I and J Calicular and lateral view
respectively; Platytrochus hastatus (MNHN uncatalogued, Ebisco stn. DW2559) K, L, and M Calicular
and lateral (GCD and LCD) views respectively; Sphenotrochus hancocki (MNHN uncatalogued, Ebisco stn.
DW2617) N and O Calicular and oblique view respectively. Scale bars: blue = 1 mm; white = 5 mm. Bold
Plate 20. Cyathotrochus pileus (USNM uncatalogued, Bathus 3 stn. CP833) A and B Calicular and
lateral view respectively; Heterocyathus aequicostatus (USNM uncatalogued, Bathus 4 stn. DW933)
C, D, and E Calicular, lateral, and base view respectively; Heteropsammia cochlea (USNM uncata-
logued, Bathus 3 stn. DW894) F, G, and H Calicular, lateral, and base view respectively; Placotro-
chus laevis (USNM 81989) I and J Calicular and lateral view respectively; Falcatoflabellum raoulensis
(MNHN uncatalogued, Ebisco stn. DW2603) K, L, and M Calicular, lateral, and oblique view respec-
tively; Placotrochides scaphula (MNHN uncatalogued, Chalcal stn. DW75) N and O Calicular and
Plate 21. Kionotrochus suteri (NZOI F915) A (SEM) and B (SEM): Calicular and lateral view re-
spectively; Anthemiphyllia dentata (USNM uncatalogued, Bathus 4 stn. DW914) C, D, and E Calicu-
lar, lateral, and base view respectively; Bourneotrochus stellulatus (USNM uncatalogued, Bathus 3 stn.
DW877) F, G, and H Calicular, lateral, and base view respectively; Peponocyathus folliculus (MNHN
uncatalogued, Norfolk 1 stn. DW1697) I, J, and K Calicular, lateral, and base view respectively; Trocho-
cyathus (Trochocyathus) discus (MNHN uncatalogued, Biocal stn. DW46) L, M, and N Calicular, lateral,
and base view respectively; Temnotrochus kermadecensis (MNHN uncatalogued, Musorstom 5 stn.
DW328) O and P Calicular and lateral view respectively. Scale bars: blue = 1 mm; white = 5 mm. Bold
Plate 22. Idiotrochus emarciatus (MNHN uncatalogued, Ebisco stn. DW2632) A, B, and C Calicular,
oblique, and lateral view respectively; Dunocyathus parasiticus (USNM 85697) D (SEM) and E (SEM):
Oblique and lateral view respectively; Endopachys grayi (USNM uncatalogued, Norfolk 2 stn. DW2158)
G and H Calicular and lateral view respectively; Notophyllia recta (USNM 85752) H, I, and J Calicular,
oblique, and lateral view respectively; Australocyathus vincentinus (USNM 85699) K and L Oblique
views respectively; Truncatoguynia irregularis (USNM uncatalogued, Norfolk 2 stn. DW2117) M and
N Calicular and lateral view respectively. Scale bars: blue = 1 mm; white = 5 mm. Bold face indicates type
species for the genus.
Plate 23. Blastotrochus nutrix (USNM 97553)– A and B Calicular and lateral view respectively; Trun-
catoflabellum sp. (MNHN uncatalogued, Concalis stn. DW2934) C, D, E, and F Calicular, oblique, lat-
eral, and basal scar view respectively; Caryophyllia (Caryophyllia) abrupta (MNHN-Scl.2009-0067) G and
H Calicular and lateral view respectively. Scale bars: blue = 1 mm; white = 5 mm. Bold face indicates type
species for the genus.
Plate 24. A and B Caryophyllia ralphae (MNHN-Scl.2009-0077, A lateral view and B pedicel section):
Green and red arrows indicating costae and monocyclic base respectively C and D Rhizosmilia robusta
(USNM uncatalogued, Norfolk 2 stn. DW2114 C pedicel section and D lateral view): Red and green
arrows indicating polycyclic base and costae respectively E Astroides calycularis (USNM 78767, colony
view): Blue arrows indicating coenosteum F Dendrophyllia ijimai (USNM uncatalogued, Bathus 4 stn.
DW933, lateral colony view): Yellow arrow indicating the axial polyp G Sphenotrochus hancocki (MNHN
uncatalogued, Ebisco stn. DW2617, oblique view): Green arrows indicating costae H Trochocyathus hasta-
tus (MNHN uncatalogued, Ebisco stn. DW2497, oblique view): White arrow indicating costal spines
I Truncatoflabellum vigintifarium (MNHN uncatalogued, Ebisco stn. DW2578, oblique view): White
arrow indicating lateral edge spines J Idiotrochus emarciatus (MNHN uncatalogued, Ebisco stn. DW2632,
lateral view): White arrow indicating lateral edge spines (fish tail) K Tropidocyathus lessoni (MNHN uncat-
alogued, Musorstom 8 stn. DW1105, lateral view): White arrow indicating alate edge crests L Caryophyl-
lia unicristata (MNHN-Scl.2009-0094, oblique view): White arrow indicating very sinuous lateral crest
M Stephanocyathus weberianus (MNHN uncatalogued, Musorstom 5 stn. DW313, lateral view): White
and green arrows indicating tubercles and costae respectively; Scale bars: blue = 1 mm; white = 5 mm.
Plate 25. A Indeterminate Caryophylliina (lateral view of live specimen, Roatan, Honduras ~200 m
deep): Green arrow indicating the edge zone B Javania sp. (USNM uncatalogued, Norfolk 2 stn. CH 2115,
broken pedicel section): Blue arrow indicating tectura layers C Leptoseris gardineri (JCU uncatalogued,
Australia, septal detail [SEM]): Yellow arrow indicating meniane D Letepsammia franki (MNHN uncata-
logued, Musorstom 6 stn. CP464, oblique view): Blue arrow indicating marginal shelf E Balanophyllia
carinata (MNHN uncatalogued, Chalcal stn. D22, calicular view): Green diagram indicating a complete
septal system arranged in a Pourtalès Plan configuration F Fungiacyathus sp. (MNHN uncatalogued, Biocal
stn. CP17, oblique view): Synapticular plates highlighted in green G and H Truncatoflabellum candeanun
(CSIRO uncatalogued, SS102005 stn. 170-086, lateral views) G anthocyathus and H anthocaulus I and
J Bourneotrochus stellulatus (MNHN uncatalogued, Musorstom 4 stn. DW162, lateral views) I anthocya-
thus and J a specimen undergoing transverse division. Scale bars: blue = 1 mm; white = 5 mm.

Cairns & Kitahara 2012 Key Scleractinia (Anthozoa)

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Cairns & Kitahara 2012 Key Scleractinia (Anthozoa)

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A peer-reviewed open-access journal

ZooKeys 227: 1–47 (2012)

An illustrated key to the genera and subgenera of

Stephen D. Cairns1, Marcelo V. Kitahara2

1 Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution,

Corresponding author: Stephen D. Cairns ([email protected])

The ready identification of azooxanthellate Scleractinia (determined herein by depth of

Key to the Genera and Subgenera of the Recent Azooxanthellate Scleractinia

12b Corallum large (bushy to arborescent), with multiple successive generations

26a Columella massive ...........................[SE Atl.] Sclerhelia Plate 4, Figures I–J

39a Corallum stoloniferous (reptoid)....[IWP] Rhizosmilia Plate 6, Figures K–L

51a Pali before septa of penultimate cycle.............................................................

61b Septa arranged normally............................................................................63

85b Base of corallum intact..................................................................................

97a Thecal edge crests present..........................................................................98

108a Septa arranged in a Pourtalès Plan; thecal edges often crested........................

Figure and plates

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