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Wien, März 2011
Tridacna ( Chametrachea ) costata R oa-Q u iaoit, K ochzius,
J an tzen , A l-Z ibd ah & R ic h te r from the Red Sea,
a junior synonym of Tridacna squamosina S tu ran y, 1899
(Bivalvia, Tridacnidae)
M. Huber* & A. Eschner**
A b strac t
Analysis of the material of Tridacnidae collected during the Pola Red Sea Expeditions 1895/96
and 1897/98 revealed that the recently described Tridacna (Chametrachea) costata, R oa Quiaoit, K ochzius, Jantzen , A l-Z ibdah & R ichter in R ichter, R oa -Q uiaoit , Jantzen , A lZibdah & K ochzius , 2008 is identical with T. elongata var. squamosina S turany , 1899. For
more than one hundred years the material has been kept in the Mollusca collection of the Natural
Histoiy Museum in Vienna without proper identification. During the analysis of the material it
was possible to identify the relevant specimens with the consequence that T. costata has to be
considered a junior synonym of T. squamosina. With this newly designated lectotype the southemmost occurrence of T. squamosina is recorded.
Key Words: Red Sea, Bivalvia, Cardioidea, Tridacna, squamosina, costata, Sturany, Pola type
material.
Z usam m en fassu n g
Bei Untersuchung der Tridacnidae, die während der Pola Expeditionen ins Rote Meer 1895/96
und 1897/98 aufgesammelt wurden, zeigte sich, dass die jüngst beschriebene Tridacna (Chame­
trachea) costata, R oa -Q uiaoit, K ochzius, Jantzen , A l-Z ibdah & R ichter in R ichter, R oa Quiaoit, Jantzen , A l-Z ibdah & K ochzius , 2008 mit T. elongata var. squamosina S turany ,
1899 identisch ist. Über 100 Jahre war das Expeditionsmaterial in der M olluskensammlung des
Naturhistorischen M useum in W ien aufbewahrt worden, ohne eingehende Identifizierung.
Während der Bearbeitung des Materials konnten die entsprechenden Exemplare identifiziert
werden, mit der Konsequenz, dass T. costata als Juniorsynonym von T. squamosina anzusehen
ist. Mit dem neu designierten Lectotypus wird gleichzeitig auch die südlichste Verbreitung doku­
mentiert.
Dr. M arkus H uber, U niversity Zürich, Zoological M useum, W interthurerstrasse 190, 8057 Zürich,
** Switzerland. - markus.huber@ access.uzh.ch
Mag. A nita Eschner, N aturhistorisches M useum W ien, 3. Zoologische A bteilung, B urgring 7, 1010
Wien, Austria, - [email protected]
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Introduction
Giant clam s are am ong the m ost spectacular marine bivalves. H uge size, ease o f recognition, peculiar feeding m echanism and shallow r e ef habitats are w ell know n features of
this group. Tridacna B ruguiere , 1797 en co m p a sses the h ea v iest and after K uphus
G uettard , 1770 the seco n d largest recent b iv a lve. N um erous tridacnids have been
described. M ost m odern authors consider currently seven extant Tridacna as valid; in
one case, however, the validity is disputed. For Tridacna rosewateri S irenko & S carlato,
1991 it is unclear i f this species is a habitat-variant - as indicated by B enzie & W illiams
(1998), or a distinct species endem ic to the M ascarene Plateau, in the sense o f N ewman
& G omez (2000). L ucas et al. (1990, 1991) described a new species from Tonga and Fiji
w hich tum ed out to be a junior synonym o f T. mbalavuana L a d d , 1934 from the Upper
Tertiary o f Fiji. P aleon tological studies, w h ich tried to reconstruct the evolution and
radiation o f Tridacnidae im plied that modern lineages originated in the P aleogene and
early N eo g en e o f the East African-Arabian Province (see H arzhauser et al. 2008).
D ifficulties in the m orphological classification o f tridacnids have led to modern DN A based genetic studies. T hese m olecular genetic analyses tried to clarify the Situation o f
giant clam populations in the northem part o f the Red Sea and concluded that there live
at least three sp ecies o f Tridacna (R o a -Q uiaoit 2005; M ohamed et al. 2006). In fact
M ohamed et al. (2006) even indicate the possibility o f five extant R ed Sea species. Fur­
thermore, there are doubts, w hether the species com m only identified as T. squam osa
L am arck , 1819 from the R ed Sea is ind eed L am arck’s true T. squam osa (J.J. ter
Poorten, pers. com m ., Oct. 2010). D efinitely, the number and identity o f the living Red
Sea tridacnids is currently unresolved. Consequently, the respective Red Sea species is
here termed T. cf. squam osa.
Recently, an 8 ^ Tridacna species has been described from the northem Red Sea. Tri­
dacna costata R o a -Q uiaoit K ochzius , Jantzen , A l-Z ibdah & R ichter , 2008 presents
clear m orphological, habitat and genetic diagnostics com pared to the other tw o well
know n tridacnids occurring in the R ed Sea, T. m axim a (R öding , 1798) and T. cf.
squam osa L amarck , 1819.
In 1899, Sturany published the results o f the "Pola"-Expedition to the Red Sea. He m en­
tioned three Tridacna taxa found during this expedition o f w hich one was new.
The m aterial from the Pola expeditions forms a significant part o f the scientific collections in the N aturhistorisches M useum W ien, A ustria (NH M W ). In the course o f 7 expe­
ditions - all carried out by SM Ship Pola - clearly defined areas o f the ocean w ere system atically explored. B etw een 1890 and 1894, the eastern M editerranean, and from
1895 to 1898, the R ed Sea w ere sam pled. C om pared to larger expeditions o f other coun­
tries, these expeditions by the A ustro-H ungarian m onarchy tum ed out to be highly efficient and m inutely planned. Thanks to the use o f m odern oceanographic instm m ents, the
eastem M editerranean and the R ed Sea becam e the m ost thoroughly explored areas o f
sea at that time. For m ore details concem ing the expeditions, their scientific results and
the history o f oceanography, see S chefbeck (1991).
M olluscs alone collected during the Pola expeditions in the Red Sea am ount to 1300
series o f gastropods and bivalves (S tagl et al. 1996). As the curator responsible at the
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155
k u.k. Hofm useum , R udolf Sturany w orked on the m ollusc m aterial gathered during the
Pola expeditions. Sturany published a series o f papers on gastropods (S t u r a n y 1896;
|900a, b; 1904) and bivalves ( S t u r a n y 1896; 1899). Unfortunately, his w orks either
remained disregarded or were inaccessible to English speaking researchers.
Sturany’s publications are m andatory reading for every Red Sea bivalve specialist. His
report on the Red Sea "Lam ellibranchiaten" dating from 1899 was first published separately as a "Sonderdruck" and 1901 reprinted as part o f the "Berichte der Com m ission
für oceanographische Forschungen" w hich w ere published in the "D enkschriften der
m ath em atisch -n atu rw issenschaftlichen C lasse d er k aiserlichen A kadem ie der W is­
senschaften, Wien"
Sturany described m any rare deepw ater species dredged during the Pola Expedition.
Included were also m any shallow w ater species collected near or on shore.
M aterial
The com plete dry and w et m aterial o f the Pola Expeditions - including the syntypes o f
T elongata var. squam osina - was originally inventoried at the M ollusca C ollection
under the num ber N H M W Moll. 38076.
Sturany did not individually label each specim en. However, Sturany stated the respective Red Sea stations in his report and the Station data are available for each o f the spec­
imens. With very few exceptions the w hole w et and dry m aterial o f Sturany was avail­
able and conform ed to the stations indicated.
Six T. squam osina syntypes w ere collected in the G u lf o f A qaba, in D ahab (4 speci­
mens) and in Sharm el Sheikh (2 specim ens). The seventh and largest specim en originated in the southem Red Sea at K am aran Island, o ff Yemen. A ltogether Sturany studied
7 squamosina specim ens ranging in size from 102 to 190 mm.
Almost 30 Pola specim ens conform to T. maxima. M ore than 30 Pola specim ens conform to T. cf. squam osa. C ontrary to the Situation nowadays, T. cf. squam osa was around
1900 the m ost com m only collected Tridacna in the R ed Sea.
Results
U nfortunately, m any authors including R osew ater (1965) and Z u s c h in & O liv er
(2003) as w ell as R o a -Q uiao it et al. (2008) and very recent w orks o f R u sm o r e - V il laum e (2008) and K n o p (2009) have overlooked Sturany’s publication. Only O liver
(1992) m entioned his paper and he assigned S turany’s T. rudis Rve. to T. squam osa
L a m a r c k , 1819.
Sturany sorted the well over 60 tridacnids collected by the Pola expedition and recognized 3 taxa. Two Red Sea species - T. m axim a and T. cf. squam osa - were found com ­
m only in various lots w ith approxim ately 30 specim ens each, one species in ju st 3 lots
with 7 specim ens. This is in accordance w ith m odern findings.
S t u r a n y (1899) briefly described the nov. var. T. elongata squam osina:
hingegen
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eine Varietät besonders hervorzuheben, die system atisch zur Tr. squam osa Lm. hinüber­
fuhrt. Diese m it squam osina nov. var. zu bezeichnende Form liegt von den Localitäten
12, 14 und 43 in m ehreren E xem plaren vor " He pointed out the characteristic "
gegen den U nterrand blättrig aufgestellte Q uerschuppen der R ippen "
R o a - Q u ia o it (2005: table 9) elaborated precisely the characteristics o f T. costata - shell
asym m etrical, hinge length < h a lf shell length, scutes crow ded and w ell spaced, wide
byssal orifice, and 5 -6 deeply triangular radial folds (Table 1).
A close m orphological com parison o f Sturany’s T. squam osina w ith the new ly described
T. costata left no doubt that these tw o species are identical. This assessem ent was con­
firm ed by J. J. ter Poorten, a w ell know n cardioid specialist. In addition, tw o o f Stu­
ran y ’s three collecting stations conform to the type locality o f T. costata in the G ulf of
Aqaba.
A lthough Tridacna squam osina S t u r a n y , 1899 was not found cited in other publications, Sturany’s species was validly proposed and is recognizably characterized. Seven
syntypes are unam biguously available. Sturany’s nam e is not preoccupied.
Consequently, a lectotype o f T. squam osina is herein designated and T. costata formally
synonym ized.
W ith respect to the description o f the habitat, spawning, genetics o f T. squam osina and
com parisons w ith other species o f Tridacna, the w orks o f R o a - Q u ia o it et al. (2008) and
esp ecially the thesis o f R o a - Q u i a o i t (2005) provide an excellent overview . These
aspects are not discussed in this article.
However, two facts m erit being added. M ore than 100 years ago T. squam osina was also
the species least com m only collected. The locality o f Station 43 further indicates that T.
squam osina m ay be found throughout the Red Sea and not ju st in its northem part.
Taxonom y
Tridacna squamosina
S t u r a n y , 1899
(Figs. 1-6)
Tridacna elongata var. squamosina S t u r a n y , 1899: 283
Tridacna nov. sp. R o a - Q u i a o it , 2005: 67, 72, 75-77 (with an extended characterization o f nov. sp. and a
comparison o f the other Red Sea species)
Tridacna costata R o a -Q u i a o it , K o c h z iu s , J a n t z e n , A l -Z ib d a h & R ic h t e r in R ic h t e r , R o a -Q u ia o it ,
Ja n t z e n , A l - Z ib d a h & K o c h z i u s , 2008: 1349-1354 (syn. nov.)
Type m aterial. Lectotype (Figs. 1-6) selected herein: largest specimen selected from the material Sturany
described as Tridacna elongata var. squamosina. - Red Sea, Yemen, Kamaran Island, ca. 15° 17’ N; 42° 37’
E, shallow water, Pola expedition leg. F. Steindachner & F. Siebenrock 1.-3. Nov. 1897 [Loc. 43], (NHMW
Moll. 107075); paralectotypes: Egypt, Dahab, shallow water, Pola expedition leg. F. Steindachner & F.
Siebenrock 6. Apr. 1896 [Loc. 12] (4 specimens, N HM W Moll. 107076; Figs. 7, 10); Egypt, Sharm el
Sheikh [Sherm Sheik], shallow water, Pola expedition leg. F. Steindachner & F. Siebenrock 1. Apr. 1896
[Loc. 14] (2 specimens, N HM W Moll. 107077).
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Table 1: D iag n o stic criteria in tridacnids after R o a -Q u ia o it (2005) and H u b e r (2010)
T. squamosina________ T. squamosa__________ T. maxima
Shell symmetry
Hinge length
Scutes
Radial folds
asymmetrical
< half shell length
crowded, well spaced
5-6 deeply triangular
Byssal orifice
Mantle patterns
Incurrent tentacles
Maximum size
Distribution
Habitat
wide
subdued
distinct
320 mm (Red Sea)
Red Sea only
to 2 m, byssally
weakly attached
symmetrical
= half shell length
large, well spaced
4-6 pointed to
bluntly rounded
narrow
subdued mottled
distinct
476 mm (Tonga)
Indo-Pacific
to 25.5 m,
byssally attached
asymmetrical
< half shell length
low, crowded
usually 5 sharply
triangular
moderate to wide
colored
indistinct
500 mm (India)
Indo-Pacific
to 32 m,
usually embedded
C om parative m aterial. Tridacna cf. squamosa (Figs. 8, 11) - Red Sea, Egypt, G ulf o f Aqaba, Nawibi,
Pola expedition leg. F. Steindachner & F. Siebenrock 9.-10. Apr. 1896 [Loc. 10] (NHM W Moll. 38076);
Tridacna maxima (Figs. 9, 12) - Red Sea, Egypt, Shadwan Island, Pola expedition leg. F. Steindachner & F.
Siebenrock 18.-20. Feb. 1896 [Loc. 16] (NHM W Moll. 38077).
D iscussion
Earlier w orkers treated tridacnids as their ow n superfam ily (e.g. K een 1969;
Starobogatov 1992). H ow ever, m ost m odern analyses, including S chneider (1998),
B raley & H ealy (1998), S chneider & F oighil (1999), G iribet et al. (2002) and M atsumoto (2003) demonstrate a clo se relationship betw een tridacnids and cardiids, either
to Fragum R öding , 1798 or even to N em ocardium (K eenaea) H abe , 1951 (M atsumoto
2003). C onsequently, m any m odern researchers attribute tridacnids subfam ilial status
within Cardiidae. From an exclu sive phylogenetic point o f view , this placem ent m ay be
favored.
A superfam ily T ridacnoidea L am arck , 1819 is certainly no longer valid. Tridacnids
must be understood as firm portion o f C ardioidea L am arck , 1809, see B ieler et al.
(2010). However, the restricted biogeography o f tridacnids, their peculiar habitats and
mode o f life, their unique anatomy, together w ith their size and w eight nonetheless justifies a special treatm ent w ithin Cardioidea. As such, a fam ilial treatm ent as Tridacnidae
L amarck , 1819 is here preferred and T. squam osina is understood as m em ber o f this
small bivalve family.
Iredale (1937) started to divide Tridacna and proposed five additional genera for Australian and Pacific species. He even created for the m ost variable tridacnid, T. maxima,
two distinct genera (i.e. Vulgodacna for T. fo s s o r and Sepidacna for T. throughtoni,
which today are both considered synonym ous to T. maxim a). B ut it is doubtful if Persikima is genetically distinct from Tridacna s.s. (R oa -Q uiaoit 2005; S chneider & Ö
F oighil 1999). In addition, the subgenus Chametrachea M örch , 1853 was earlier used
by H errmannsen (1846), w hich is based on Chama aspera R um phius , 1705. Taking
further into account the characteristic features o f each species and the sm all num ber o f
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Figs. 1—6. Tridacna squamosina —lectotype, NHMW Moll. 107075 from Kamaran. 1, right valve
inside; 2, left valve inside; 3, right valve outside; 4, left valve outside; 5, byssal orifice; 6, radial
folds. Scalebar: 2 cm. Photographs: A. Schumacher.
8 valid tridacnids, a forced differentiation o f various subgenera w ithin Tridacna adds lit­
tle benefit. M ore im portant are the real differences betw een species regarding habitats,
m odes o f life, genetics and detailed m orphology. This was excellently w orked out by
R o a -Q u ia o it (2005). C onsequently, the subgeneric level is here abandoned and T.
squam osina is placed as Tridacna.
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Figs. 7-12. Comparison of the three Tridacna species from the Red Sea in the collection of the
NHMW; top: radial folds; bottom: byssal orifice. 7, 10, Tridacna squamosina - paralectotype,
NHMW Moll. 107076a from Dahab; 8, 11, Tridacna cf. squamosa - NHMW Moll. 38076 from
Nawibi; 9, 12, Tridacna maxima - NHMW Moll. 38077 from Shadwan. Scalebar: 2 cm. Photographs: A. Schumacher.
S t u r a n y (1899) reco gnized three R ed Sea taxa. He clearly used the m onograph o f
R eeve (1862) on Tridacna as an identification guide. He identified elongata correctly.
Sturany’s elongata conform s to T. elongata L a m a r c k , 1819. But in the 1950s and 1960s
it was recognized that L am arck’s elongata is a junior synonym o f Tridachnes maxima
R ö d in g , 1798. Tridacna maxima (R ö d in g , 1798) is today recognized as valid, earliest
name for this w idely distributed species.
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The second com m on Red Sea species identified by Sturany was T. rudis. R eeve (1862)
described T. rudis from the Philippines. Indeed, R eeve’s pl. 5 fig. 4 a, b, c superficially
resem bles the m aterial identified by Sturany. B ased on R osew ater’s treatm ent o f this
family, m odern authors today accept R eeve‘s rudis as a peculiar form o f maxima. Consequently, this second species was erroneously identified by Sturany as rudis. This was
recognized by O liv e r (1992), w ho identified it as L am arck’s T. squam osa, here termed
T. cf. squamosa. It is not excluded that Tridachnes imbricata R öding, 1798 was the ear­
lier nam e for L am arck’s fam ous species. Large portions o f the B olten collection, on
w hich R öding based his imbricata, are still to be found in M useum der N atur at Gotha,
Germany. However, a critical review o f the respective m aterial at G otha in September
2010 led to the follow ing conclusion: "Overall, due to m issing descriptions and specific
m arks in Röding, due to lack o f original num bers and labels, due to the peculiar suppressing and renam ing m ethod o f Schm idt and finally due to the curational condition of
the collection we could not unam biguouslv identifv anv B olten/R öding Bivalve type."
(H u b e r & t e r P o o r te n in prep.). Thus, Tridachnes im bricata R ö d in g , 1798 is best
treated as nom en dubium
T he third, the least com m on species w as described by Sturany as squam osina. R oaQ uiaoit (2005) stated the closest m orphological affm ities for T. costata are to T. max­
ima. S turany (1899) cam e to the sam e result in considering squam osina closer to elon­
gata 0=maxima) than to squam osa. From a m olecular phylogeny based on m itochondrial
16 rDN A, R oa -Q uiaoit (2005) concluded that T. costata's closest relative is T. maxima,
w hereas low er affm ities w ere found betw een T. squam osa and T. crocea. For further
d etailed inform ation on T. squ a m o sin a w e refer to R o a -Q uiaoit et al. (2008 as T.
costata) and especially to R o a -Q uiaoit (2005 as T. nov. sp.).
As such, Tridacna squam osina S turany , 1899 is the 8 ^ m em ber o f the genus Tridacna
w ithin the fam ily Tridacnidae. Currently, it is only know n from the R ed Sea, likely living throughout. It is the least com m on o f the described three R ed Sea tridacnids. The
know n habitat is v ery shallow w ithin approxim ately 5 m eters and the m axim al size
reported is 320 mm.
Acknowledgements
Thanks are due to: Prof. Tony Wilson for editorial support, Dr. Peter C. Dworschak for fruitful
discussions and valuable help with the plates, Dr. Mathias Harzhauser for help with paleontological
literature and to Alice Schumacher for the photographs of the material. For their precise remarks
and additions, the authors are deeply indebted to one anonymous reviewer and J.J. ter Poorten,
The Netherlands.
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