On Pseudocorynactis species and another related genus from the Indo-Pacific (Anthozoa:corallimorphidae)

              Rev. Acad. Canar. Cienc, XXI (Nums. 3-4), 9-34 (2009) (publicado en septiembre de 2010)
ON PSEUDOCORYNACTIS SPECIES AND ANOTHER
RELATED GENUS FROM THE INDO-PACIFIC
(ANTHOZOA: CORALLIMORPHIDAE)
O. Ocana', J. C. den Hartog , A. Brito^ & A.R. Bos'
' Departamento de Oceanografia Biologica y Biodiversidad, Fundacion Museo del Mar,
Muelle Canonero Dato s.n, 51001, Ceuta, North Africa, Spain, lebruni@telefonica.net
- J. C. den Hartog, National Museum of Natural History, Postbus 9517, 2300 RA Leiden, The Netherlands.
Deceased (1942-2000).
^ Gmpo de Investigacion BIOECOMAC, Departamento de Biologia Animal, Facultad de Biologia,
Universidad de La Laguna, C/ Astrofisico Sanchez s.n., 38206 La Laguna, Tenerife, islas Canarias. abrito@ull.es
^ Davao del Norte State College, New Visayas, 8105 Panabo, Philippines, arthurrbos@yahoo.com
ABSTRACT
Tentacles development is important for the Corallimorphidae in terms of speciation,
being interesting to discuss about the presence of different evolutionary levels in tentacle
anatomy.
We studied three species of Corallimorpharia belonging two of them to Pseiido-corynactis
and one to the new genus Paracorynactis. The extendible capability of the tenta-cles
is proposed to make difference among the above mentioned genera. The species Pseudo-corynactis
globulifera from the Red Sea is include in this genus for the first time, we also
have described the new species Pseudocorynactis tuberculata from Indonesia and Maldives.
The help control of Paracorynactis hoplites on the crown of thorns sea stars population is
exposed.
Key words: Corallimorpharians, tentacles extendible capacity, new genus, new
species, new combination, Indo-Pacific region.
RESUMEN
El desarrollo tentacular es importante para comprender los procesos de especiacion
en la familia Corallimorphidae y poder discutir acerca de la existencia de niveles evolutivos
relacionados con la anatomia de los tentaculos.
En este articulo estudiamos tres especies de Corallimorpharia pertenecientes dos
al genero Pseudocorynactis y una al nuevo genero Paracorynactis. La capacidad de
extension de los tentaculos es una caracteristica que proponemos para diferenciar los
generos. Incluimos a la especie Pseudocorynactis globulifera en este genero por primera
vez y tambien se describe una nueva especie, Pseudocorynactis tuberculata, a partir de
material procedente de Indonesia y las Maldivas. Es interesante destacar la capacidad de
la especie Paracorynactis hoplites para ejercer cierto control sobre las poblaciones de la
estrella de mar corona de pinchos que tantos estragos causa a los madreporarios en el
Indo-Pacifico.
Palabras claves: Corallimorpharia, capacidad de extension de los tentaculos, nuevo
genero, nueva especie, nueva combinacion, region Indopacifica.
1. INTRODUCTION
The genus Pseudocoiynactis was described by den HARTOG [12] to accommodate a
new group of Corallimoipharians from the Caribbean, with extremely well developed acros-pheres.
A species from the same genus was found in the Canary Islands, and den HARTOG
et al. [15] added some additional characteristics, which increased the knowledge of this
species. After these descriptions, new material with Pseudocorynactis resemblance was col-lected
in the Indo-pacific region and provisional names were added to images of these
species printed in several sea life identification guides (den HARTOG [14]; GOSLINER et
al. [8]; COLIN & ARNESON [6]). Moreover, the name Pseudocorynactis showed up in
other publications dealing with a range of marine subjects (see TOMASCIK et al. [19]; BOS
et al. [2]). Meanwhile, some classical papers keep descriptions of Corallimorpharians from
the tropical Indo-Pacific region, such as the genus Coiynactis (see KLUNZINGER [16];
HADDON & SHACKLETON [11]; HADDON [9]; CARLGREN [4], [5]). Coiynactis glob-uligera
described by Ehrenberg, 1834 (see KLUZINGER, op. cit.) from Red Sea and
Corynactis hoplites by HADDON & SHACKLETON {op. cit.) from Flores Sea, are good
samples of this assertion.
The family Corallimorphidae includes the genera Coiynactis, Pseudocorynactis,
Corallimorphus and also Nectactis and Sideractis (see den HARTOG et al., op. cit.). In the
present paper a new genus, Paracorynactis, is introduced and described, in order to accom-modate
one of the species studied. From our point of view, tentacles development is impor-tant
for the Corallimorphidae in terms of speciation. Morphologically, size, type and quanti-ty
of nematocysts differ among genera, whereas acrosphere development and stalk differen-tiation
may also differ. Following the differences in the acrosphere development degrees, it
is possible to assume the presence of different evolutionary levels in tentacle anatomy which
justifies a newly proposed genus. The genera Corynactis, Paracorynactis and
Pseudocorynactis follow three different steps in tentacle development which relates to both
acrospheres and the stalk. The genus Corynactis has the less acrosphere development in the
family and a limited stalk extension capacity. Paracorynactis shows high acrosphere devel-opment
and has an intermediate capacity of stalk extension; whereas the genus
Pseudocorynactis has a more advanced strategy in extending the tentacles stalk and high
acrosphere development. All Corynactis species known today follow this concept (Ocaiia in
prep.).
The present study deals with material collected in Indonesia (Sulawesi, Sumbawa,
Salayer, Tukang Besi islands, Sumba, Komodo) and Maldives by the National Museum of
Natural History (formally RMNH). Material from Aden, Zanzibar, and Siam was addition-ally
studied.
The macro-anatomical analysis of the used specimens was adequate to detect differ-ences
at species level. Histological slides are stored at the National Museum of Natural
History in Leiden and are available to further study the different genera.
10
2. MATERIAL AND METHODS
The specimens studied come from different locations within the Indo-Pacific region.
The majority was collected by scuba divers, but some were collected with a "van Veen" grab
or a rectangular dredge. Samples were fixed with 8-10% formaldehyde and later stored in
70% alcohol in the collections of the National Museum of Natural History in Leiden, The
Netherlands. All the type material is deposited at the RMNH in Leiden. General morpholo-gy
and anatomy were studied by means of a stereo dissecting microscope. The anatomical
and micro anatomical details were studied using staining in toto. Nematocysts (>1500 cap-sules
measured) were examined with a light microscope equipped with a Nomarski differen-tial
interference contrast optic system. The classification and terminology of nematocysts
follows that of SCHMIDT [18], as adapted by den HARTOG [12: 7-9] and den HARTOG et
al. {op. cit). The surveys of the cnidom are summarized in tables in which the means and
ranges of length and width of nematocysts are included. The following codes are used in the
tables: vc: very common; c: common; re: rather common; uc: uncommon; r: rare.
Between October 2007 and November 2008, life specimens of Paracorynactis
hoplites were observed and measured in Samal Island in the Davao Gulf, the Philippines.
The diameter of the polyps was measured with calipers (0.5 cm accuracy) and their depth
was recorded (1 m accuracy). These observations were done during SCUBA-dives with a
maximum depth of 40 m.
3. RESULTS AND DISCUSSION
Taxonomical key for genera of the Corallimorphidae:
Acrospheres very prominent, clearly differentiated from stalk and contain
special spirocysts (den HARTOG, 1 980 and den HARTOG etal, \992>) 1
Acrospheres conspicuous but usually not very prominent and
differentiated from stalk; the special spirocysts are always absent 2
la. Stalk highly extendible (more than the column length), nematocysts
absent from the stalk Pseudocorynactis
lb. Stalk much less extendible, nematocysts present in the stalk Paracorynactis
2a. Small gregarious forms living in intertidal and shallow waters,
disc tentacles present Corynactis
2b. Small or large forms without disc tentacles, apparently never
recorded gregarious and living in deep-waters 3
3a. Large solitary forms, low developed tentacles and two categories of
spirulae in tentacles Corallimorphiis
3b. Small forms from deep waters with one small spirulae category in
acrospheres 4
4a. Presence of large atrichs in acrospheres and several categories of
nematocysts in tentacles, usually attached to hard susbstratum Sideractis
4b. Nematocysts similar to those of Corallimorphus, but (apparently)
not a sessile animal/species Nectactis
11
Genus Paracorynactis gen. nov.
Diagnosis: Solitary form which may reach a relatively large size (diameter of the oral disc
10 to 15 cm); clones have never been observed; 3 to 5 radial rows of tentacles concentrate
at the disc periphery, this character is much more conspicuous in expanded conditions. The
most developed tentacles are concentrated in the disc periphery and at the margin. Tentacle
stalks with medium to moderate extending capacity; nematocysts of several categories pres-ent
in the stalk. All mesenteries are perfect and provided with restricted enlarged parietal
ridges. Cnidom: there are big special spirocysts (length >100 p.m) in the acrospheres, show-ing
the high development of the acrospheres.
Paracorynactis hoplites (Haddon & Shackleton, 1893)
(Figs. 1-3 and 9)
Corynactis hoplites Haddon & Shackleton, 1893: 118-119, no plates, Torres Strait, original description.
Haddon, 1898: 467-468, plate XXX figs. 1-4. Torres Strait; Haddon & Duerden, 1896: 153, only a brief
reference; Carlgren, 1949, reference and distribution; den Hartog, Ocana & Brito, 1993: 22, 26 & 27,
only references about the lack of information about the species.
Material.- Coll. 31023: Indonesia, N Sulawesi, Selat Lembeh, between Tanjungnanas and Teluk
Kungkungan: 0r28'N 125°14'E; steep rocky shore, rockface down to 16 m, coral covered slope;
against dark wall in shadow of trees, diving; 28.x. 1994. N. Sulawesi Expedition, 1994. 3 specimens.
Images of the habitus of the two specimens. 7 cm x 4.5 cm; 5.5 cm x 4 cm; 8 cm x 3.5 cm.
Coll. 31030: Indonesia, SW Sulawesi, Spermonde Archipelago, NNW of Pulau Badi (=20Km NNW of
Ujungpandang); 4°57'S 1 19°17'E, coralreef, under large, thick plate of dead coral, scuba diving, depth
8m. 14.iv.l997. Buginesia prog. UNHAS-NNM. 1 specimen. Images of the habitus of the specimen.
Colonized partially by fungi. 2.5 cm x 0.5 cm.
Coll. 31031: Indonesia, SW Sulawesi, Spermonde Archipelago, NNW of Samalona (=7.5 Km W of
Ujungpandang); 5°07'S 119°20'E, coral reef, under side of dead coral, scuba diving, depth 5-10 m.
21.ix.l994. Buginesia prog. UNHAS-NNM. 2 specimens, one of them very small. Images of the habi-tus
of the two specimens. 2.3 cm x 1.5 cm; 0.6 cm x 0.2 cm.
Coll. 31036: Sta. S4.114: Indonesia, N of Sumbawa, Bay of Sanggar, 8°19.2'S118°14.4'E, lagoon side
of reef barrier, depth -20 m, snorkelling, scuba diving, 21/22-09-84. "Tyro" Indonesian-Dutch Snellius-
11 Exped. Two specimens. Images of the habitus of the two specimens. 5.5 cm x 2.7 cm; 6 cm x 3.5 cm.
Coll. 31038: Sta. S4. 152: Indonesia, SW Salayer, NW coast of Pulau Guang, 06°21'S 120°27'E, reef
flat, round steep wall in cave, scuba diving, 28/29-09-1984. "Tyro" Indonesian-Dutch Snellius-II
Exped. One specimen. Images of the habitus of the specimen. 2.5 cm x 1.5 cm.
Coll. 31037: Sta. S4. 152: Indonesia, SW Salayer, NW coast of Pulau Guang, 06°21'S 120°27'E, steep
cliff in hole, 4-5 m, snorkeling, 29-09-1984. "Tyro" Indonesian-Dutch Snellius-II Exped. One speci-men.
Images of the habitus of the specimen. 5 cm x 2.5 cm.
Diagnosis (complete anatomical descriptions in HADDON [9]:
Color in the column is often conservative brown, although tentacles and oral disc
should be variable: brown oral disc and translucent tentacles with white longitudinal stripes
in the middle of the tentacles, acrospheres of ochre color; grey oral disc and tentacles with
white longitudinal stripes not very conspicuous. In preserved material there is no color trace
in the acrospheres of the studied specimens.
Irregular base in outline, often spread on substrate. Column well developed with no
distinction between scapus and scapulus noticed, although the upper part of the column
12
seems less corrugate. The diameter in preserved specimens varies from 0.2 to 4.5 centime-tres
(attending to its trophic behavior, some specimens should be larger; BOS et al. [2]), in
live specimens the column can reach 10 cm in height. Short radial rows of 2 to 5 ectacmaceus
tentacles, large specimens may have up to 180 tentacles or even more. Tentacles are assem-bled
at the disc periphery and marginal areas of the disc. Tentacles stalk with medium to
moderate extending capacity, nematocysts of several categories present in the stalk.
Acrospheres with a great development of the ectodermic tissues (see cnidom in this study
and HADDON, op. cit: 467), similar to what was found in the genus Pseudocorynactis (den
HARTOGer^/. [15]).
Apparently, the species does not develop siphonoglyphs. The pharynx presents
numerous folds. Most mesenteries are complete; they can be 90 to 105 in number and pres-ent
some incomplete ones growing in the exocoelic (fig. 2c). Among the directives, a couple
of endocoelic mesenteries were observed too (fig. 2c). Sphincter more or less enlarged and
restricted to the upper part of the column (fig. 2a and b; OCANA [17]). Strong ectodermic
musculature of the tentacles bearing some mesogloeal process but it does not have a brush
like structure (den HARTOG et al, op. cit.). Endodermic musculature from the column con-spicuous
and well developed. Retractors weak, only conspicuous in free mesenteries,
restricted just before the cnidoglandular portion of the mesenteries and developing some
ridges (fig. 2c). Parietobasilar muscles present well developed enlarged restricted
mesogloeal ridges in all the mesenteries (fig. 2c and d).
Cnidom (table I, fig. 3): We analyzed the complete cnidoms of three specimens and,
also partially, the tentacles and the body wall of three other specimens. Once the cnidom of
the species is well known, the cnidom from the tentacles and the body wall are good indica-tors
to distinguish one species from another, P-mastigophores E from the tentacles of
Paracorynactis hoplites are larger than in the other species studied in this paper, although
there are no relevant differences in the size of the cnidae between small and large specimens.
Spirulae from tentacles reflect slight differences between small and large specimens. Big
homotrichs from tentacles are common but often appear broken in two parts making meas-uring
difficult. Small p-mastigophores D from tentacles are not very obvious and easily over-looked.
In the sample Coll. 31031 a second category of homotrich from tentacles was
observed. There are measurement differences linked to specimen sizes (Table I), although
Figura 1.- Features on different specimens oi Paracorynactis hoplites: a) specimen from coll. 31036; b) specimen
from coll. 31038.
13
Figura 2.- a) Sphincter development marked by arrows from one specimen of the coll. 31023; b) idem in another
specimen of the coll. 31023; c) mesenteries arrangement and its retractor development from one specimen of the
coll. 31023; d) mesenteries arrangement and enlarged restricted mesogloeal ridges from one specimen of the coll.
31023.
14
these are relevant in relation to the intra-specific variability but do not significantly affect the
distinction between species. Large specimens usually have longer and wider nematocysts. In
a few occasions the opposite was observed; e.g. the specimen 31038 had larger penicilli D
in the tentacles.
Due to the poor conditions of the tentacle tissue in specimens 31037, it was not pos-sible
to size the homotrichs from the tentacles. There was a second penicilli E category in
the filaments of the specimens 31023 and 31030. Due to their rare presence we included the
measurement range (53-63 |Lim) x (22-26 |Lim). Another very scarce spirulae category was
found in the body wall, but possibly it was produced by contamination. In the tentacles we
observed another homotrich category of smaller size, inconspicuous and scarce but merit to
be studied carefully in order to distinguish the species from other species of the genus
Paracorynactis
.
There are two spirocysts categories, one large (>100 |Lim and <250 |u.m) and a much
smaller as known from other soft bodied coral species. We do not include these spirocysts
measurements.
Table I.- Paracorynactis hoplites. Survey of the cnidom of three specimens from different locaHties. A: Coll. 3 1023:
Indonesia, N Sulawesi; B: Coll. 31030: Indonesia, SW Sulawesi; C: Coll. 31036: Sta. S4.114: Indonesia, N of
Sumbawa; D: Coll. 31031: Indonesia, SW Sulawesi; E: Coll. 31037: Sta. S4. 152: Indonesia, SW Salayer; F: Coll.
31038: Sta. S4. 152: Indonesia, SW Salayer. D, E & F showing the sizes classes present on tentacles and body wall
ORGAN PROCEDENCE NEMATOCYSTS lYPE
MEAN AND RANGE OF LENGTH AND WITH
OF NEMATOCYST CAPSULES IN ^m N FREQUENCY
Tentacles A Spirulae 1 25.5(22-301x3 (2.5-3.5) 10 RC-UC
B 22.2(20-251x3.1 (3-3.51 15 RC-UC
C 25(18-331x3.4(3-3.5) 15 C-RC
D 22.5(17-301x3.1 (2.5-41 10 RC-UC
E 29.5(25-341x3(2.5-3.51 6 UC-RC
F 25.7(23-311x2.8(2-3.5) 10 RC-UC
A Spirulae 2 36.9 (35-40) x 6.5 (6-7.5) 11 RC-C
B 31.3 (30-34) x 6,1 (6-6.5) 10 RC-UC
C 34.8 (30-40) X 6.7 (6-8) 15 C-VC
D 31 (27-351x6.2(6-6.51 2 R
E 32 (27-371 X 5.5 (5-6) 8 UC-RC
F 29.2 (27-32) x 6 (5-7) 4 UC-R
A Penicilli E 220 (185-250) X 18.9 (18-21) 15 C-RC
B 194(170-2101x17.6(16-20) 15 C-VC
C 218 (190-250) X 20 (17-23) 10 RC
D 186(143-2251x19.4(16-221 15 RC
E 246(215-2601x19.3(15-211 15 RC-C
F 217.8(185-2451x18(14-231 15 C-RC
A Penicilli D1 158(150-1651x5(5-5.51 10 C
B 163(155-1701x5.6(5-61 10 C
C 159(150-1751x5.1 (5-61 10 C
D 161 (140-1851x5(5-5.51 10 C
E 164(153-1801x5 15 C
15
ORGAN PROCEDENCE NEMATOCYSTS TYPE
MEAN AND RANGE OF LENGTH AND WITH
OF NEMATOCYST CAPSULES \N^m
N FREQUENCY
F 175(160-1901x5.1 (4-6) 15 C
A Penicilli D2 30,8(28-331x5.1 (4-61 10 RC
B 25 (20-301 X 5 8 RC-UC
c 27.8(25-311x5.1 (5-61 10 RC-UC
D 29.5 (29-301 X 5 5 R
E 30x5 1 R
F 29.5(28-311x5(5-5.51 5 R-UC
A Homotrichsl 132.8(112-1501x5.6(5-71 10 RC-UC
B 139(130-1451x5.5(5-71 10 RC-UC
C 142.6(125-1581x5.7(4-6.51 15 C-VC
D 142.3(127-1701x6.4(5-71 15 C-VC
E 140(135-1451x6 5 RC-UC
F 148.5(136-1601x5.6(5-61 10 RC
D Homotrichs2 82.4(70-901x3.2(2.5-41 10 RC-UC
Pharynx A Penicilli E 92(80-1041x22.3(20-261 15 C
B 77.7(71-831x22.2(20-241 10 C
C 101.6(95-1081x23.7(20-251 10 C
A Homotrichs 39.6 (32-451 x 4.6 (4-61 15 C
B 38.5(35-421x4.5(4-51 15 C
C 40.8 (35-471 X 4.6 (4-51 20 C
Filaments A Penicilli E 101 (88-1101x23.6(15-251 25 C-VC
B 83.6(70-951x26.4(20-271 15 c
C 105(87-1171x25.4(22-301 20 c
A Penicilli D 41.3(37-461x7.4(7-8.51 10 c
B 36.2(33-421x6.6(6-71 10 c
C 41.6(33-471x6.6(5-81 10 c
Body wall A Penicilli A 22.8(20-251x6.5(6-71 10 RC-UC
B 20.2(15-251x6(5-7.51 20 C-VC
C 21.4(18-281x5.9(5-71 15 RC-UC
D 20(15-251x6(5-71 5 UC
E 21.7(20-241x5.9(5-71 10 RC
F 23.2(21-261x6.7(5.5-71 5 UC
A Spirulae 22.8 (20-271 x 6.1 (5-71 10 RC-C
B 19(15-241x5.5(4-71 15 C
C 20.8(16-241x6.3(6-71 15 C
D 21.4(16-251x5,7(5-6.51 15 RC-C
E 20(14-271x5.6(5-71 15 RC-C
F 20.7(17-251x5.7(5-61 15 C
A Homotrich 33.5 (30-401 x 5.9 (4-81 20 C
B 28.7 (24- 321 x 6.2 (5-7.51 15 RC
C 32.3 (28-371 x 6 (5-71 11 C
D 35.5 (30-401 x 7.2 (7-81 5 UC-RC
E 31.4(27-351x6.7(5-8.51 10 C-RC
F 35.1 (28-371x6.8(6-81 10 C-RC
16
Figura 3.- Pictorial survey of the cnidom.
ACROSPHERES
17
STALK PHARYNX
Honiotrichs Penicilli D
V
1
FILAMENTS
18
BODY WALL
Homotrich Spirulae Penicilli A
Biological notes: In total, 70 life specimens of Paracorynactis hoplites were observed dur-ing
27 dives in the Davao Gulf between October 2007 and November 2008. The deepest
observation was a specimen at 28 m depth, whereas specimens were also found in shallow
water up to 2 m. The mean depth of the polyps was 7.3 m indicating that the majority of the
polyps was found in relatively shallow water. The diameter of the polyps ranged from 4 to
17 cm with a mean of 9.6 cm.
A newly discovered predator of the crown-of-thoms starfish is a relevant example of
how as the authors assume Paracorynactis hoplites may help control the crown-of-thoms
population (BOS et al. [2], BOS [1]).
Remarks: The new genus presents intermediate characters between Corynactis and
Pseudocorynactis. Acrospheres very prominent, with special spirocysts, and differentiated
widely from stalk (den HARTOG [12] and DEN HARTOG et al, op. cit.), stalk with nema-tocysts
and less extending capability compared to Pseudocorynactis. The absence of a spe-cial
structure (den HARTOG et al., op. cit.) in the tentacle musculature enforce the assertion.
This group of attributes makes to our present new genus a single one among the other gen-era
belonging to the Corallimorphidae family.
We identified the studied material of the new genus to the species described by HAD-DON
& SHACKLETON [11] on the base of the musculature (sphincter and mesenteries)
showed by the first author (HADDON [9]: 467-468, plate XXX figs. 1-4., figs. 2 and 4) The
color varieties described in both papers underline the previous assertion. They have been
known from some time ago (see den HARTOG [14]; and present color data) and it is a good
way to identify the species from others belonging to Pseudocorynactis genus. Furthermore,
the habitat and geographical area of the material examined in the present study are highly
comparable to those of Haddon & Shackleton's study. We included small specimens (see
studied material) as did HADDON & SHACKLETON (op. cit.). Similarly, Corynactis glob-ulifera
was attributed to Pseudocorynactis a long time ago (den HARTOG et al, op. cit. 27)
and apparently only small specimens have been found.
We assumed that the collection of HADDON & SHACKLETON [op. cit.] was in
Dublin, but K.W. England informed J.C. den Hartog that the collection was with the
Cambridge University. Unfortunately this material was not made available from Zoological
Museum at Cambridge University. A re-examination of Haddon's material is highly desired
19
and may contain two different species {Paracorynactis hoplites and Pseudocorynactis tuber-ciilata).
In the case of no type material from Haddon's collection a new type series (Neotype)
should be established with the material deposited in Leiden.
Coiynactis hoplites was considered by den HARTOG et al. {op. cit.) a possible valid
species, the second Corynactis recorded from tropical areas after Corynactis parvula
Duchassaing & Michelotti, 1860 (den HARTOG et al, op. cit.).
The absence of the Corynactis genus from the tropical Indo-Pacific areas should be
taking into account. The presence of Corynactis parvula in certain areas from the Caribbean
Sea may be an exceptional recent case that needs further study. It seems plausible to think
that Coiynactis is restricted to temperate waters (upper bathial in some cases) with the
exception of C. parvula that can colonize some tropical environments and which merit to be
studied separately. Pseudocorynactis and Paracorynactis are tropical-subtropical genera not
known from temperate waters.
Genus Pseudocorynactis den Hartog, 1980
Additions to the diagnosis of the genus: Pseudocorynactis present the highest degree of ten-tacular
differentiation from the shallow water genera {Corynactis, Paracorynactis and
Pseudocorynactis). The stalk may extend enormously, more than the column length and the
nematocysts are absolutely absent from the stalk. From the histological and microanatomi-cal
point of view, the stalk presents strong developed ectodermal longitudinal musculature
supported by conspicuous, brush-like mesogloeal processes (see den HARTOG et al, op.
cit\?>\). The last comment should be included in the genus diagnosis.
Remarks in relation to the species Pseudocorynactis caboverdensis den Hartog, Ocana &
Brito, 1993: This species studied on the basis of a single specimen is better to be placed into
the genus Corynactis. The species was tentatively included into the genus Pseudocorynactis
(den HARTOG et al., op. cit.) although strikingly resemble C. parvula. So, in the present
paper, we include that species into the genus Corynactis. Meanwhile, much more informa-tion
is needed to focus on both Corynactis parvula and Corynactis caboverdensis.
Corynactis caboverdensis presents nematocysts in the stalk of their tentacles and does
not have large spirocysts in the acrospheres, which are characteristic of Pseudocorynactis.
As we pointed out (see den HARTOG et al., op. cit.) "In several respects Pseudocorynactis
caboverdensis seems closer to species of the genus Corynactis''. In addition, it should be
taken into account that the habitat where the specimen was encountered at Cape Verde
Islands is similar habitats as described for Corynactis spp. in other regions (see den HAR-TOG
et al., op. cit: 21; OCANA, op. cit.: 419).
Pseudocorynactis globulifera (Ehrenberg, 1 834) com. nov.
Ectacmaea globulifera EhrQnbQTg, 1834: 39
Corynactis globulifera Kluzinger, 1877: 73, Taf V. fig. 8.
Corynactis globulifera ? (Carlgren, 1900): 40, no figures, Baui island, Zanzibar. The measurements of
the P-mastigophore E from acrosphere indicate close relation to P. globulifera.
Corynactis globulifera, 1943: 7-8, fig. 2. Short description of the material from Siam, comparative of
nematocysts on the material from Siam, Zanzibar and the Red Sea.
20
Material.- Coll 39561: Corynactis globiilifera, intertidal: attached to stone beneath sand. Ras
Jarshyne, Aden. Collected by: K.W. England, 1966. K.W.England collection. 1 small specimen..
Images of the habitus of the specimen. 1.8 cm x 0.8 cm.
Other material analyzed:- ZMS 144: Corynactis globulifera, (Ehr) Roda havet, Koseir, Kluzinger 1855,
Det. Kluzinger. Fragment van Berlin Museum.
ZMS 145: O. Afrika, Sansibar. Insel Baui, 29/6/1889, Stuhlmann (Fran Hamburg Mus.). Fragment van
Stuhlmaim exemplar.
ZMS 1264: Corynactis globulifera, Siam, of Koh Kut, 15 fms. Three specimens, T. Mortensen leg.,
1900, id. O. Carlgren.
Diagnosis:
In preserved conditions (alcohol) the color is pale greyish, tentacles stalks transpar-ent
and acropheres of ochre color. Base irregular in outline, column marked by numerous
distinct ridges, thick and cartilaginous in texture. Oral disk concave and the texture seems
corrugate (fig. 4). Tentacles arranged in endocoelic radial rows, alternating with simple exo-coelic
ones. 48 endocoelic rows with 3-5 or mostly 4 tentacles each row. They are alternat-ing
with single exocoelic tentacles, the largest. In most cases, the penultimate tentacle is the
largest, but sometimes it is the second large tentacle or about equal to the length of the ulti-mate
tentacle. The total number of tentacles reaches 115. The oldest tentacles are relatively
near to the centre of the oral disc. Tentacles stalk with high extensive capability.
There are 24 pairs of mesenteries with two pairs of directives. The directives separate
a series of 10 to/and 12 pairs of mesenteries but the arrangement is not entirely clear. Primary
and secondary mesenteries are perfect, tertiary cycle mostly imperfect, there are also some
small of low development.
Sphincter endodermic and concentrated in the upper part of the column.
Parietobasilar muscles well developed, enlarged restricted mesogloeal ridges in all the
mesenteries (see fig. 4b).
Figura 4.- a) Specimen oiP. globiilifera from Aden collected by K. England; b) close-up of the parietobasilar ridges.
21
Cnidom (table II; fig. 5): A survey of the cnidom is summarized in the table II, adding
some data from the other material analyzed.
We have analyzed the complete cnidom of one specimen collected by K. England in
Aden. The small length of the P-mastigophores E, the scarcity of P-mastigophores E catego-
Table II.- Pseiidocorynactis globidifera. Survey of the cnidom of the material from Aden collected by K. W.
England: A. Additional information of the other analyzed material ZMS 144: B; ZMS 1264: C.
ORGAN PROCEDENCE NEMATOCYSTS TYPE
MEAN AND RANGE OF LENGTH AND WITH
OF NEMATOCYST CAPSULES IN /jm
N FREQUENCY
Tentacles A Spirulae 1 27(22-311x3.4(2.5-4) 20 UC
B 25.3 (21-28) X 3.1 (3-3.5) 5 UC
c 24 (20-28) X 3.5 (3-4) 20 RC
A Penicilli E 110(70-1351x15(13-20) 50 C-VC
B 120(118-130x17(16-19) 10 C-VC
c 119 (110-130) X 17.4 (15-19) 10 C-VC
A Penicilli D1 87.4(75-1001x4.5(4-51 15 C-VC
B 101 (95-1151x6.4(6-7) 20 C-CV
c 98(85-1101x6(5-6.51 15 C-VC
A Honnotrichsl 90.4(77-1051x6(4-7) 30 C-VC
c 99.6 (90-110) X 6.5 (6-7) 20 C-CV
A Honnotrichs2 56 (50-60) X 4.5 (4-5) 5 UC
Pharynx A Penicilli E 57 (54-60) X 20 2 R
C 51.2(45-551x17.3(16-201 10 UC
A Homotrichs 32.6 (32-401 X 5.3 (4-61 20 RC-C
C 30(25-331x5.5(5-61 25 C
Filannents A Penicilli El 115(100-1301x28(19-301 35 C-VC
B 118(105-251x26.5(25-30) 10 RC
C 100 (95-105) X 30 3 UC
A Penicilli E2 65 (55-75) X 21.6 (16-26) 25 C-VC
B 63.4 (60-70) X 20.6 (20-221 10 RC-UC
C 62.7(58-701x20(17-251 22 C-VC
A Penicilli D 24.2 (20-301 X 6.2 (5-71 20 VC-C
B 27.2 (24-341 x 7 (6-81 20 VC-C
C 26.3 (23-301 X 7 (6-81 25 vc
A Spirulae 11.2(10-141x3.5(3-41 20 c
C 11.5(10-151x3.5(3-41 15 c
Body wall A Penicilli A 25.8(21-311x6.6(5-71 30 vc
B 25.6(21-281x6.1 (5.5-71 20 c
C 24.5(18-301x6.5(5-71 30 vc
A Spirulael 10.3(7-121x3.4(3-3.51 20 UC-RC
C 9.5(8-111x3.7(3-4) 15 RC
A Spirulae2 20.1 (16-23) x 5.5 (4-6) 20 UC
B 20.4 (15-22) X 6.1 (5-7) 15 RC
C 15.5(13-181x5.5(4-61 10 UC
A Homotrich 37.4(32-401x8.2(7.5-91 10 UC
22
Figura 5.- Pictorial survey of the cnidom.
ACROSPHERES
ry in the pharynx and the presence of small categories of spirulae are distinctive characters.
We have not noticed any differences concerning to the size of the specimens studied.
Nematocysts absent from the tentacle stalks. Spirulae 2 from tentacles very scarce.
There are two spirocysts categories, one large (>100 |am and <150) )Lim) and the other
smaller than usually found in other soft bodied coral species. We do not include these spiro-cysts
measurements.
The homotrichs 2 have only been found in the specimen from Aden, although it is an
uncommon nematocyst. In the pharynx of the specimen from Aden we found small and spo-radic
penicilli D. The pharynx from the specimen B was not studied due to poor conserva-
23
PHARYNX
BODY WALL
24
FILAMENTS
25
tion of this particular tissue. In this specimen, spirulae from the pharynx were not observed.
Possibly, due to the poor conservation, spirulae of small size and homotrich from the body
wall, also in the specimen C, were not seen.
Notes about other material analyzed:
The material from Siam (see CARLGREN [4]) was on an eroded fragment of
Xenophora-Caryophyllid coral. Base 10x6.5 mm, oral disc semi-contracted 13x10 mm,
height of column 12 mm. Column with distinct prominent ridges corresponding to endo-coeles
(ectoderm partly macerated). 24 rows of 3-5 endocoelic tentacles alternating with 24
singleton exocoelics. Mesenteries arrangement of one half showed two pairs of directives
and 11 pairs of mesenteries divided into perfect and imperfect ones. The color in preserved
conditions is greyish with dirty ochre acrospheres. An abstract of the nematocyst measure-ments
is shown in table II.
The material from the Red Sea, studied by Kluzinger, is not fully retracted but tenta-cles
and acrospheres protrude. Diameter of the base 1 3 mm, somewhat spread, height in
semi-contracted state 8 mm. Color in preserved conditions greenish brown with yellowish-brown-
green acrospheres.
Remarks: CARLGREN (op. cit.) studied the nematocysts of the material from Siam
(Mortensen Expedition) and additionally examined the cnidom of the material from
Zanzibar, previously studied by himself (see CARLGREN [3]), and from the Red Sea, this
last material had already been studied by KLUZINGER [16]. After his examinations,
Carlgren concluded that all the material belonged to Coiynactis globidifera. We had the
opportunity to examine the same material as Carlgren (see material) and we have found out
that the material present the same characters showed by Carlgren paper of 1943, especially
concerning the nematocysts measurements. According to this, the length of the Penicilli E
from acrospheres and the width of the homotrichs from the column are the main character-istics
to diffemtiate Pseudocorynactis globidifera from other species of Psendocorynactis
and the species of the new genus Paracoiynactis.
The color features described by KLUZINGER {op. cit.) are also coincident with our
observations about the material from Aden.
The absence of the nematocysts in the stalks and its extension capacity makes clear
that the species belongs to the genus Pseudocoiynactis.
Nevertheless, examination of more material is needed in order to establish the distri-bution
of this species along the Indo-Pacific.
Pseudocorynactis tuberculata n.sp.
(Figs. 6-8)
Type material: Coll. 31035: 84.106: Indonesia, NE of Komodo; 8°26.9'S119°37.9'E; depth 80 m; cal-careous
stones and nodules with Porifera, Bryozoa, rectangular dredge; 20-ix-1984. "Tyro" Indonesian-
Dutch Snellius-II Exped. Two specimens. Image of the habitus of the specimen. 2.5 cm x 1 cm; 2.8 cm
X 1.6 cm. complete specimen attached on the stone Holotype, sectioned specimen Paratype.
Coll. 31032: S4.016: Indonesia, Tukang Besi islands, Banda Sea, Kaledupa reef, E of entrance, 5°56'S
123°48'E, scuba diving & snorkelling, gently stoping reef, 1-10 m, 06/08-09-1984. "Tyro" Indonesian-
Dutch Snellius-II Exped. One specimen. Images of the habitus of the specimen. 2.2 cm x 0.9 cm. Paratype.
26
Coll. 31033: S4.045: Indonesia, NE coast of Sumba, E of Melolo; 09°54.2'S120°43'E; coarse sand
with shell gravel, some calcareous stones, depth 48-57 m; van Veen grab; 13-ix-1984. "Tyro"
Indonesian-Dutch Snellius-II Exped. One specimen. Images of the habitus of the specimen. 2.5 cm x
1 cm. Paratype.
Coll. 31034: S4.051: Indonesia, NE coast of Sumba, E of Melolo; 09°53.5'S120°42.7'E; depth 75-90
m; calcareous stones, rectangular dredge; 13-ix-1984. "Tyro" Indonesian-Dutch Snellius-II Exped.
One specimen. Images of the habitus of the specimen. 3.8 cm x 1.5 cm. Paratype.
Coll. 31039: Maldives, Kartu Atoll, Villingili, depth 12 m, night dive. 24-iii-1989. M. van der Knaap.
One small specimen sectioned at the middle. Image of the habitus of the specimen. 1.3 cm x 0.7 cm.
Paratype.
Coll. 31040: Maldives, Kanifmolhu, depth 10 m, night dive. 22-vi-1989. M. van der Knaap. One small
specimen. Image of the habitus of the specimen. 0.8 cm x 0.4 cm. Paratype.
Diagnosis:
Base generally spreaded on substrate, upper part of the column marked by conspicu-ous
tubercles without ectoderm, not with foreign particles adhered to mucous coat. It seems
that tubercles are connected by narrow channels (fig. 7a). Definitely, there are specimens
much more tuberculate than others. Color of column orange-red. Tentacles with developed
acrosphere and a high extension capacity, arranged in endocoelic radial rows alternating with
simple exocoelic ones. 24 endocoelic rows with 3-5, mostly 4 tentacles each, are alternating
with single exocoelic tentacles, the largest. Nearly always the penultimate tentacle is the
largest, and sometimes the second tentacle is as long as the ultimate tentacle, or the same
length. The total number reaches 120-150 tentacles. Oldest tentacles are relatively near to the
centre of the oral disc.
Figura 6.- Tubercles in two different specimens.
There are 24 pairs of mesenteries, directives should be present but were not recog-nized.
All the mesenteries are perfect, but the third cycle present pairs of less development
(especially because the slender parietobasilar ridge), giving the impression of imperfect ones.
27
Figura 7.- a) Sphincter and connection channels of tubercules; b) Parietobasilar mesogloeal ridges.
Sphincter endodermic and concentrated in the upper part of the column (fig. 7a).
Parietobasilar muscles well developed and encircled, restricted mesogloeal ridges in all the
mesenteries (fig. 7b).
Cnidom (fig. 8): A survey of the cnidom is summarized in the table III. The presence
of two penicilli D categories in the column, one common and the other uncommon or spo-radic,
is characteristic from the body wall. Normally there are size differences of the spiru-lae
from tentacles among specimens, also seen in P. caribbeorum material. Nematocysts
absent from the tentacles stalk. Fortunately, we have not noticed relevant differences con-cerning
the size of the specimens studied. In the small specimens from Maldives there are
some nematocyst size differences compared to the other studied material. Although these dif-ferences
have to be understood in relation to the intra-specific variability and they do not sig-nificantly
affect, in order to distinguish the species properly.
Table III.- Pseudocorynactis tuberculata. Survey of the cnidom of two specimens from different locahties. A: Coll.
31039: Maldivas; B: Coll. 31033: Indonesia, C: Coll. 31034: Indonesia; D: Coll. 31032: Indonesia; E: Coll. 31035
Indonesia. C, D & E showing only the sizes classes present on tentacles and body wall.
ORGAN PROCEDENCE NEMATOCYSTS TYPE
MEAN AND RANGE OF LENGTH AND WITH
OF NEMATOCYST CAPSULES IN /jm
N FREQUENCY
Tentacles A Spimlae 1 35(30-451x4.6(3.5-5.5) 30 RC-C
B 49.7 (40-55) X 4.3 (4-5) 20 RC-UC
C 45.6 (40-50) X 4.2 (4-5) 10 RC-UC
D 41 (40-45) X 4.1 (4-4.5) 5 UC
E 52x4 1 UC
28
ORGAN PROCEDENCE NEMATOCYSTS TYPE
MEAN AND RANGE OF LENGTH AND WITH
OF NEMATOCYST CAPSULES IN ^m N FREQUENCY
A Penicilli E 156.4 (125-190) X 20.6 (12-28) 30 C-RC
B 161 (145-180) X 20.6 (12-25) 10 RC-UC
C 175 (165-180) X 20.3 (20-21) 5 RC
D 158.7 (145-170) X 18.2 (17-20) 5 RC-UC
E 166 (160-175) X 18.6 (18-20) 5 RC-UC
A Penicilli D1 117.2 (98-140) X 6.1 (5-8) 30 C-RC
B 129.2 (100-145) X 5.1 (5-6) 20 C-RC
C 114.6 (105-127) X 5.5 (5-6) 5 RC
D 117.5 (105-130) X 5 (4-6) 5 RC
E 135 (130-140) X 5.5 (5-6) 5 RC
A Homotnchsl 96.6 (90-100) X 5.3 (5-6) 5 R
B 106 (95-130) X 5.7 (5-6) 10 UC
C 105 (100-1 10) X 5.7 (5-6) 5 R
D 110x6 1 R
E 100x5.5 1 R
A Homotnchs2 65x4 1 R
B 55x3.5 1 R
C 50x3 1 R
Pharynx A Penicilli E 64.3 (54-73) X 24.5 (19-31) 30 C-RC
B 77.2 (65-90) X 27 (20-32) 20 C-RC
A Homotrichs 32.3 (25-38) x 6 (5-7) 30 RC-C
B 30 (25-35) X 5.2 (4-6) 15 UC-RC
Filaments A Penicilli El 84.2 (70-100) X 37 (28-48) 40 C-RC
B 82.3 (75-90) X 33 (25-43) 25 VC-C
B Penicilli E2 64 (63-65) X 20.5 (20-21) 2 R
B Homotrichs 29.6 (27-33) X 4.6 (4-6) 10 UC
A Penicilli D 24 (20-28) X 7 (5-8) 40 RC-C
B 20.3 (20-26) X 6.2 (5-8) 15 RC
Body wail A Penicilli D1 37.8 (27-46) X 10.5 (8-12) 30 UC-R
B 43.6 (35-51) X 10.6 (8-13) 5 R
A Penicilli D2 22.5 (15-26) X 6.4 (5-8) 60 RC
B 22.3 (17-27) X 4.7 (3.5-7) 20 RC
C 18,3 (16-22) X 4.3 (4-5) 5 UC-RC
D 19 (17-24) X 5.7 (5-7) 5 UC-RC
E 19.3 (16-22) X 6.6 (5-8) 5 UC-RC
A Spimlae 1 23.2 (20-26) X 6.8 (5-8) 30 UC
B 22 (19-25) X 5 (4-6) 5 UC-R
A Spirulae 2 12 (11-14) X 3.8 (3-4) 15 RC
A Homotrich 1 42 (33-50) X 17 (15-20) 45 C
B 38 (32-45) X 16 (13-20) 25 C-RC
c 39 (35-40) X 14.5 (14-15) 5 RC
D 37.5 (35-40) X 16 (15-17) 5 RC
E 36.5 (36-37) X 12 5 RC
A Homotnch 2 19.5 (19-20) X 4.5 (4-5) 2 R
29
Figura 8.- Pictorial suney of the cnidom.
ACROSPHERES
30
PHARYNX
FILAMENTS
31
BODY WALL
Spirulae 1 Spirulae 2 Penicilli D 1 Penicilli D 2 Homotrichs 1
Homotrichs 2
In the filaments the peniciUi E2 and the homotrichs have only been noticed on B spec-imen.
Perhaps this is due to the contamination process with the pharynx. In the body wall
from C, D and E spirulae were not observed. Homotrichs 2 are sporadic in the specimen D
so it is not important to the taxonomical concern.
Remarks: The presence of tubercles (absolutely absent from P. caibbeorum) and some
other size differences among nematocysts suggest to us to maintain these two close species
separate {Pseudocorynactis caribbeorum and Pseudocorynactis tuberculata) . P. caribbeo-rum
is distributed in both Atlantic tropical areas, so it is plausible to suppose that P. tuber-culata
may be widespread in the Indo-pacific region. The scarce divergence between both
species may be tentatively explained due to the short period of time that both, the Atlantic
and the Indo-pacific populations, have been geographically separated. After all, 1.8 millions
years ago, the Panama isthmus was closed (CRONIN & IKEYA [7]).
Tubercles are usually evident but sometimes, this character can be less conspicuous.
Fortunately, there are other nematocyst evidences to enforce the distinction of both species
too. The presence of narrow homotrichs 1 in the body wall oiP. caribbeorum makes the most
significant difference in terms of nematocysts measurement. Other morphological characters
of P. caribbeorum can be consulted (den HARTOG et al. [15]; OCANA [17])
According to the ecological data available to us, the nocturnal predator is also appli-cable
to this new species. As usually happens in the case oiP caribbeorum the body wall of
P. tuberculata is also hidden in a crevice while the tentacle crown forms a net ready to prey.
32
Figura 9.- Paracorynactis hoplites color and habitats, specimens recorded at Davao Gulf (Samal and Talikud Islands).
4. ACKNOWLEDGEMENTS
We would like to thank Leen van Ofwegen and Bert Hoeksema, curators at RMNH
(Naturalis at Leiden), for the consulting facilities the material studied for the paper.
Francisca Serrais made the general edition and Leopoldo Moro and J. J. Bacallado made sug-gestions
regarding the scientific edition. Pauline Byram checked the English language.
33
5. BIBLIOGRAPHY
[I] BOS, A.R. 2010. Crown-of-thoms Outbreak at the Tubbataha Reefs UNESCO World
Heritage Site. Zoological Studies, 2009: 124.
[2] BOS, A.R., G. S. GUMANAO & F. N. SALAC. 2008. A newly discovered predator of
the crown-of-thoms starfish. Coral Reefs, 27:581.
[3] CARLGREN, O. 1900. Ostafrikanische Actinien. Jb. Hamb. Wiss. AnsL, 17. Beiheft 2:
21-144, pis. 1-7.
[4] CARLGREN, O. 1943. East-Asiatic Corallimorpharia and Actiniaria. KunglSvenska
Vetensk. Akad. Handl (3) 20 (6): 1-43, figs. 1-32, pis. 1-2.
[5] CARLGREN, O. 1949. A Surrey of the Ptychodactiaria, Corallimorpharia and
Actiniaria. Kungl Svenska Vetensk. Akad. Handl, ser. 4, 1 (1): 1-121, pis. 1-4.
[6] COLIN, RL. & C. ARNESON. 1995. Tropical Pacific Invertebrates. Coral Reef Press,
California.
[7] CRONIN, T. M. & N. IKEYA. 1990. Tectonic events and climatic change: opportuni-ties
for speciation in cenozoic marine ostracoda. In: Causes of evolution. A paleonto-logical
perspective. Ross, R. M. & AUmon, W. D. ed.: 210-248, 4 figs.
[8] GOSLINER, LM., D.W. BEHRENS & GC. WILLIAMS. 1996. Coral reef animals of
the Indo-Pacific. Sea Challengers, USA.
[9] HADDON, A. C. 1 898. The Actiniaria from Torres Strait. Scient. Proceed. R. Dub. Soc.
(2) 6 (16): 393-498, pis. 22-32.
[10] HADDON, A.C. & J.E. DUERDEN. 1896. On some Actiniaria from Australia and other
Districts. Sc. Trans. R. Dublin Soc, (2) VI., 172 pp., plates 7-10
[II] HADDON, A. C, & A. M. SHACKLETON. 1893. Description of some new species of
Actiniaria from Torres Straits. Scient. Proceed. R. Dub. Soc. (N.S.), 8 (1): 116-131.
[12] HARTOG, J.C. den, 1980. Caribbean shallow water Corallimorpharia. Zool. Verh.
Leiden, 176: 1-83, figs. 1-20, pis. 1-14.
[13] HARTOG, J. C. den. 1994. Sea anemones of the Seychelles. In: Oceanic Reefs of the
Seychelles (Netherlands Indian Ocean Programme). Chapter 6.2, volume 2: 76. Edited
by J. van der Land, published by National Museum of Natural History.
[14] HARTOG, J. C. den. 1997. Sea anemones. In: A guide to the seashores ofEastern Africa
and the Western Indian Ocean islands. Published by Sida/Departament for Research
Cooperation, SAREC, Edited by D. Richmond.
[15] HARTOG, J. C. den, O. OCANA & A. BRITO. 1993. Corallimorpharia collected dur-ing
the CANCAP expedition (1976-1986) in the south-eastern part of the North
Atlantic. Zoologische Verhandelingen, 282: 1-76.
[16] KLUNZINGER, C. B. 1877. Die Korallthiere des Rothen Meeres, Pt. I, Die
Alcyonarien und Malacodermen: 1-95, pis. 1-8. Berlin.
[17] OCANA, O. 1994. Anemonas (Actiniaria y Corallimorpharia) de la Macaronesia
Central: Canarias y Madeira. Tesis Doctoral, Universidad de La Laguna, 2 Vol., 485
pp., 166 lam.
[18] SCHMIDT, H. 1972. Prodromus zu einer Monographic der mediterranen Aktinien.
Zoologica, Stuttgart, 121: 1-146,36 figs.
[19] TOMASCIK, T.A., J. MAH, A. MONTH & M. K. MOOSA. 1997. The Ecology ofthe
Indonesian Seas. Part one. The Ecology of Indonesia Series. Vol. VII. Periplus Editions.
34