ON PSEUDOCORYNACTIS SPECIES AND ANOTHER
RELATED GENUS FROM THE INDO-PACIFIC
O. Ocaña1, J. C. den Hartog2, A. Brito3& A.R. Bos4
1Departamento de Oceanografía Biológica y Biodiversidad, Fundación Museo del Mar,
Muelle Cañonero Dato s.n, 51001, Ceuta, North Africa, Spain. firstname.lastname@example.org
2J. C. den Hartog, National Museum of Natural History, Postbus 9517, 2300 RA Leiden, The Netherlands.
3Grupo de Investigación BIOECOMAC, Departamento de Biología Animal, Facultad de Biología,
Universidad de La Laguna, C/ Astrofísico Sánchez s.n., 38206 La Laguna, Tenerife, islas Canarias. email@example.com
4Davao del Norte State College, New Visayas, 8105 Panabo, Philippines. firstname.lastname@example.org
Tentacles development is important for the Corallimorphidae in terms of speciation,
being interesting to discuss about the presence of different evolutionary levels in tentacle
We studied three species of Corallimorpharia belonging two of them to Pseudo-
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
Key words: Corallimorpharians, tentacles extendible capacity, new genus, new
species, new combination, Indo-Pacific region.
El desarrollo tentacular es importante para comprender los procesos de especiación
en la familia Corallimorphidae y poder discutir acerca de la existencia de niveles evolutivos
relacionados con la anatomía de los tentáculos.
En este artículo estudiamos tres especies de Corallimorpharia pertenecientes dos
al género Pseudocorynactis y una al nuevo género Paracorynactis. La capacidad de
extensión de los tentáculos es una característica que proponemos para diferenciar los
géneros. Incluimos a la especie Pseudocorynactis globulifera en este género por primera
vez y también 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
Rev. Acad. Canar. Cienc., XXI (Núms. 3-4), 9-34 (2009) (publicado en septiembre de 2010)
estrella de mar corona de pinchos que tantos estragos causa a los madreporarios en el
Palabras claves: Corallimorpharia, capacidad de extensión de los tentáculos, nuevo
género, nueva especie, nueva combinación, región Indopacífica.
The genus Pseudocorynactis was described by den HARTOG  to accommodate a
new group of Corallimorpharians 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.  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 ; GOSLINER et
al. ; COLIN & ARNESON ). Moreover, the name Pseudocorynactis showed up in
other publications dealing with a range of marine subjects (see TOMASCIK et al. ; BOS
et al. ). Meanwhile, some classical papers keep descriptions of Corallimorpharians from
the tropical Indo-Pacific region, such as the genus Corynactis (see KLUNZINGER ;
HADDON & SHACKLETON ; HADDON ; CARLGREN , ). Corynactis 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 Corynactis, 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 (Ocaña in
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-
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.
2. MATERIALAND 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 , 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; rc: 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, 1980 and den HARTOG et al., 1993) ................... 1
Acrospheres conspicuous but usually not very prominent and
differentiated from stalk; the special spirocysts are always absent ........................... 2
1a. Stalk highly extendible (more than the column length), nematocysts
absent from the stalk ............................................................... Pseudocorynactis
1b. Stalk much less extendible, nematocysts present in the stalk ............ Paracorynactis
2a. Small gregarious forms living in intertidal and shallow waters,
disc tentacles present ......................................................................
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 ................................................................... Corallimorphus
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 ................................................................
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 µm) 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, Ocaña & 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: 01º28’N 125º14’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); 4º57’S 119º17’E, coralreef, under large, thick plate of dead coral, scuba diving, depth
8m. 14.iv.1997. 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); 5º07’S 119º20’E, coral reef, under side of dead coral, scuba diving, depth 5-10 m.
21.ix.1994. 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, 8º19.2’S118º14.4’E, lagoon side
of reef barrier, depth -20 m, snorkelling, scuba diving, 21/22-09-84. “Tyro” Indonesian-Dutch Snellius-
II 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, 06º21’S 120º27’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, 06º21’S 120º27’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 :
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
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. ), 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
HARTOG et al. ).
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; OCAÑA ). 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 of Paracorynactis hoplites: a) specimen from coll. 31036; b) specimen
from coll. 31038.
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.
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 µm) × (22-26 µm). 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
There are two spirocysts categories, one large (>100 µm and <250 µm) and a much
smaller as known from other soft bodied coral species. We do not include these spirocysts
TentaclesA Spirulae 1 25.5 (22-30) x 3 (2.5-3.5)10 RC-UC
B 22.2 (20-25) x 3.1 (3-3.5) 15RC-UC
C 25 (18-33) x 3.4 (3-3.5)15 C-RC
D 22.5 (17-30) x 3.1 (2.5-4)10RC-UC
E 29.5 (25-34) x 3 (2.5-3.5)6 UC-RC
F 25.7 (23-31) x 2.8 (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)10RC-UC
C 34.8 (30-40) x 6.7 (6-8) 15C-VC
D 31 (27-35) x 6.2 (6-6.5)2R
E 32 (27-37) x 5.5 (5-6)8 UC-RC
F 29.2 (27-32) x 6 (5-7)4UC-R
A Penicilli E 220 (185-250) x 18.9 (18-21)15C-RC
B 194 (170-210) x 17.6 (16-20)15 C-VC
C 218 (190-250) x 20 (17-23)10RC
D 186 (143-225) x 19.4 (16-22) 15RC
E246 (215-260) x 19.3 (15-21) 15 RC-C
F217.8 (185-245) x 18 (14-23) 15C-RC
APenicilli D1158 (150-165) x 5 (5-5.5)10C
B163 (155-170) x 5.6 (5-6) 10C
C159 (150-175) x 5.1 (5-6) 10C
D161 (140-185) x 5 (5-5.5) 10C
E 164 (153-180) x 5 15C
ORGANPROCEDENCE NEMATOCYSTS TYPEN FREQUENCY
MEAN AND RANGE OF LENGTH AND WITH
OF NEMATOCYST CAPSULES IN µm
Table I.- Paracorynactis hoplites. Survey of the cnidom of three specimens from different localities. A: Coll. 31023:
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
F 175 (160-190) x 5.1 (4-6) 15C
A Penicilli D2 30.8 (28-33) x 5.1 (4-6) 10 RC
B 25 (20-30) x 58 RC-UC
C 27.8 (25-31) x 5.1 (5-6)10 RC-UC
D 29.5 (29-30) x 55R
F29.5 (28-31) x 5 (5-5.5)5 R-UC
A Homotrichs1132.8 (112-150) x 5.6 (5-7) 10RC-UC
B139 (130-145) x 5.5 (5-7) 10 RC-UC
C142.6 (125-158) x 5.7 (4-6.5) 15C-VC
D 142.3 (127-170) x 6.4 (5-7) 15C-VC
E 140 (135-145) x 65 RC-UC
F 148.5 (136-160) x 5.6 (5-6) 10 RC
D Homotrichs282.4 (70-90) x 3.2 (2.5-4)10RC-UC
PharynxA Penicilli E 92 (80-104) x 22.3 (20-26) 15C
B77.7 (71-83) x 22.2 (20-24) 10C
C101.6 (95-108) x 23.7 (20-25) 10C
A Homotrichs 39.6 (32-45) x 4.6 (4-6)15C
B38.5 (35-42) x 4.5 (4-5) 15C
C40.8 (35-47) x 4.6 (4-5)20C
FilamentsA Penicilli E101 (88-110) x 23.6 (15-25) 25C-VC
B 83.6 (70-95) x 26.4 (20-27) 15C
C105 (87-117) x 25.4 (22-30) 20 C
APenicilli D41.3 (37-46) x 7.4 (7-8.5) 10C
B 36.2 (33-42) x 6.6 (6-7)10C
C 41.6 (33-47) x 6.6 (5-8)10C
Body wallA Penicilli A22.8 (20-25) x 6.5 (6-7) 10 RC-UC
B 20.2 (15-25) x 6 (5-7.5)20C-VC
C21.4 (18-28) x 5.9 (5-7) 15RC-UC
D 20 (15-25) x 6 (5-7)5 UC
E21.7 (20-24) x 5.9 (5-7) 10RC
F23.2 (21-26) x 6.7 (5.5-7)5 UC
ASpirulae 22.8 (20-27) x 6.1 (5-7) 10RC-C
B 19 (15-24) x 5.5 (4-7)15 C
C 20.8 (16-24) x 6.3 (6-7)15C
D 21.4 (16-25) x 5.7 (5-6.5)15 RC-C
E 20 (14-27) x 5.6 (5-7)15 RC-C
F 20.7 (17-25) x 5.7 (5-6)15C
A Homotrich33.5 (30-40) x 5.9 (4-8) 20C
B 28.7 (24- 32) x 6.2 (5-7.5) 15RC
C 32.3 (28-37) x 6 (5-7) 11C
D35.5 (30-40) x 7.2 (7-8)5 UC-RC
E 31.4 (27-35) x 6.7 (5-8.5)10C-RC
F 35.1 (28-37) x 6.8 (6-8)10C-RC
ORGAN PROCEDENCENEMATOCYSTS TYPENFREQUENCY
MEAN AND RANGE OF LENGTH AND WITH
OF NEMATOCYST CAPSULES IN µm
Figura 3.- Pictorial survey of the cnidom.
Penicilli D 2
Penicilli E Homotrichs 1 & 2Penicilli D 1Spirulae 1
Penicilli E 1 Penicilli E 2 Penicilli D
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-thorns starfish is a relevant example of
how as the authors assume Paracorynactis hoplites may help control the crown-of-thorns
population (BOS et al. , BOS ).
Remarks: The new genus presents intermediate characters between Corynactis and
Pseudocorynactis. Acrospheres very prominent, with special spirocysts, and differentiated
widely from stalk (den HARTOG  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  on the base of the musculature (sphincter and mesenteries)
showed by the first author (HADDON : 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 ; 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
Homotrich Spirulae Penicilli A
and may contain two different species (Paracorynactis hoplites and Pseudocorynactis tuber-
culata). 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.
Corynactis 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 Corynactis 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:31). The last comment should be included in the genus diagnosis.
Remarks in relation to the species Pseudocorynactis caboverdensis den Hartog, Ocaña &
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; OCAÑA, op. cit.: 419).
Pseudocorynactis globulifera (Ehrenberg, 1834) com. nov.
Ectacmaea globulifera Ehrenberg, 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.
Material.- Coll 39561: Corynactis globulifera, 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
ZMS 1264: Corynactis globulifera, Siam, of Koh Kut, 15 fms. Three specimens, T. Mortensen leg.,
1900, id. O. Carlgren.
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 of P. globulifera from Aden collected by K. England; b) close-up of the parietobasilar ridges.
Cnidom (table II; fig. 5): Asurvey 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-
TentaclesA Spirulae 1 27 (22-31) x 3.4 (2.5-4)20UC
B 25.3 (21-28) x 3.1 (3-3.5)5 UC
C24 (20-28) x 3.5 (3-4) 20 RC
A Penicilli E110 (70-135) x 15 (13-20)50 C-VC
B 120 (118-130) x 17 (16-19)10 C-VC
C 119 (110-130) x 17.4 (15-19)10C-VC
A Penicilli D1 87.4 (75-100) x 4.5 (4-5)15 C-VC
B 101 (95-115) x 6.4 (6-7)20 C-CV
C 98 (85-110) x 6 (5-6.5) 15 C-VC
A Homotrichs1 90.4 (77-105) x 6 (4-7) 30C-VC
C99.6 (90-110) x 6.5 (6-7) 20C-CV
A Homotrichs2 56 (50-60) x 4.5 (4-5)5 UC
PharynxA Penicilli E 57 (54-60) x 202R
C51.2 (45-55) x 17.3 (16-20) 10 UC
A Homotrichs 32.6 (32-40) x 5.3 (4-6)20 RC-C
C 30 (25-33) x 5.5 (5-6) 25C
FilamentsA Penicilli E1 115 (100-130) x 28 (19-30)35C-VC
B 118 (105-25) x 26.5 (25-30)10 RC
C 100 (95-105) x 303 UC
A Penicilli E2 65 (55-75) x 21.6 (16-26)25 C-VC
B 63.4 (60-70) x 20.6 (20-22)10 RC-UC
C 62.7 (58-70) x 20 (17-25)22C-VC
A Penicilli D24.2 (20-30) x 6.2 (5-7) 20VC-C
B 27.2 (24-34) x 7 (6-8)20 VC-C
C26.3 (23-30) x 7 (6-8) 25VC
A Spirulae11.2 (10-14) x 3.5 (3-4) 20C
C11.5 (10-15) x 3.5 (3-4) 15C
Body wallA Penicilli A25.8 (21-31) x 6.6 (5-7) 30VC
B 25.6 (21-28) x 6.1 (5.5-7)20C
C 24.5 (18-30) x 6.5 (5-7) 30VC
A Spirulae110.3 (7-12) x 3.4 (3-3.5) 20UC-RC
C 9.5 (8-11) x 3.7 (3-4)15 RC
A Spirulae2 20.1 (16-23) x 5.5 (4-6)20UC
B 20.4 (15-22) x 6.1 (5-7)15 RC
C 15.5 (13-18) x 5.5 (4-6) 10UC
AHomotrich 37.4 (32-40) x 8.2 (7.5-9)10 UC
ORGAN PROCEDENCENEMATOCYSTS TYPEN FREQUENCY
MEAN AND RANGE OF LENGTH AND WITH
OF NEMATOCYST CAPSULES IN µm
Table II.- Pseudocorynactis globulifera. 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.
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 µm and <150) µm) and the other
smaller than usually found in other soft bodied coral species. We do not include these spiro-
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-
Figura 5.- Pictorial survey of the cnidom.
Spirocysts Penicilli EHomotrichs 1 & 2 Penicilli D 1Spirulae 1
Penicilli E Penicilli DHomotrichs
Spirulae 2 Spirulae 1HomotrichsPenicilli D
Penicilli E 1Penicilli E 2 Penicilli D
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 ) 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 13 mm, somewhat spread, height in
semi-contracted state 8 mm. Color in preserved conditions greenish brown with yellowish-
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 ), and from the Red Sea, this
last material had already been studied by KLUZINGER . After his examinations,
Carlgren concluded that all the material belonged to Corynactis globulifera. 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 differntiate Pseudocorynactis globulifera from other species of Pseudocorynactis
and the species of the new genus Paracorynactis.
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 Pseudocorynactis.
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.
Type material: Coll. 31035: S4.106: Indonesia, NE of Komodo; 8º26.9’S119º37.9’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, 5º56’S
123º48’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.
Coll. 31033: S4.045: Indonesia, NE coast of Sumba, E of Melolo; 09º54.2’S120º43’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; 09º53.5’S120º42.7’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.
Coll. 31040: Maldives, Kanifinolhu, 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.
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.
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.
Figura 6.- Tubercles in two different specimens.
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.
Figura 7.- a) Sphincter and connection channels of tubercules; b) Parietobasilar mesogloeal ridges.
TentaclesA Spirulae 135 (30-45) x 4.6 (3.5-5.5) 30RC-C
B 49.7 (40-55) x 4.3 (4-5)20RC-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 52 x 41 UC
ORGAN PROCEDENCENEMATOCYSTS TYPEN FREQUENCY
MEAN AND RANGE OF LENGTH AND WITH
OF NEMATOCYST CAPSULES IN µm
Table III.- Pseudocorynactis tuberculata. Survey of the cnidom of two specimens from different localities. 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 TYPEN FREQUENCY
MEAN AND RANGE OF LENGTH AND WITH
OF NEMATOCYST CAPSULES IN µm
A Penicilli E 156.4 (125-190) x 20.6 (12-28) 30C-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 D1117.2 (98-140) x 6.1 (5-8) 30 C-RC
B 129.2 (100-145) x 5.1 (5-6)20C-RC
C 114.6 (105-127) x 5.5 (5-6)5 RC
D 117.5 (105-130) x 5 (4-6)5 RC
E135 (130-140) x 5.5 (5-6)5 RC
A Homotrichs1 96.6 (90-100) x 5.3 (5-6)5R
B 106 (95-130) x 5.7 (5-6)10 UC
C 105 (100-110) x 5.7 (5-6)5R
D 110 x 61R
E 100 x 5.51R
A Homotrichs2 65 x 41R
B 55 x 3.51R
C 50 x 31R
PharynxA Penicilli E 64.3 (54-73) x 24.5 (19-31)30C-RC
B 77.2 (65-90) x 27 (20-32) 20C-RC
A Homotrichs 32.3 (25-38) x 6 (5-7)30RC-C
B 30 (25-35) x 5.2 (4-6) 15UC-RC
FilamentsA Penicilli E1 84.2 (70-100) x 37 (28-48)40 C-RC
B82.3 (75-90) x 33 (25-43) 25VC-C
B Penicilli E264 (63-65) x 20.5 (20-21)2R
B Homotrichs29.6 (27-33) x 4.6 (4-6) 10UC
A Penicilli D 24 (20-28) x 7 (5-8)40RC-C
B 20.3 (20-26) x 6.2 (5-8)15 RC
Body wallA Penicilli D137.8 (27-46) x 10.5 (8-12)30UC-R
B43.6 (35-51) x 10.6 (8-13)5R
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
C18.3 (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 Spirulae 123.2 (20-26) x 6.8 (5-8)30 UC
B 22 (19-25) x 5 (4-6)5 UC-R
A Spirulae 212 (11-14) x 3.8 (3-4)15 RC
AHomotrich 142 (33-50) x 17 (15-20) 45C
B38 (32-45) x 16 (13-20)25 C-RC
C39 (35-40) x 14.5 (14-15)5 RC
D37.5 (35-40) x 16 (15-17)5 RC
E 36.5 (36-37) x 125 RC
A Homotrich 219.5 (19-20) x 4.5 (4-5)2R
Figura 8.- Pictorial survey of the cnidom.
SpirocystsPenicilli E Homotrichs 1 & 2Penicilli D 1 Spirulae 1
Penicilli E 1Penicilli E 2Penicilli D
In the filaments the penicilli 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 ).
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 of P. caribbeorum makes the most
significant difference in terms of nematocysts measurement. Other morphological characters
of P. caribbeorum can be consulted (den HARTOG et al. ; OCAÑA )
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 of P. caribbeorum the body wall of
P. tuberculata is also hidden in a crevice while the tentacle crown forms a net ready to prey.
Spirulae 1Spirulae 2Penicilli D 1 Penicilli D 2Homotrichs 1
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.
Figura 9.- Paracorynactis hoplites color and habitats, specimens recorded at Davao Gulf (Samal and Talikud Islands).
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