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A
review of the Bistolida-complex of Africa's north-eastern,
eastern and southern coastlines and the adjacent Indian Ocean islands.
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Author : Ronnie
Watt, South Africa, 2005
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The
author invites comments from readers. Send e-mail to: clumber@telkomsa.net
(The following
text was adapted from the original publication in Xenophora, October
2005) |
Original
Text can be donwloaded here (.doc file)
| The
sp. and ssp. of the Bistolida-complex relevant to this discussion
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Bistolida stolida
stolida stolida (LINNAEUS 1758)
stolida
clavicola (LORENZ
1998)
stolida
brianoi (LORENZ
2002)
stolida
uvongoensis nov.
sp. (MASSIER 2004)
Bistolida diauges
(MELVILL 1888)
Bistolida erythraeensis
(SOWERBY 1837)
Bistolida hirundo hirundo (LINNAEUS
1758)
hirundo
francisca (SCHILDER
& SCHILDER 1938)
Bistolida kieneri kieneri (HIDALGO
1906)
Bistolida owenii owenii (SOWERBY 1837)
owenii
vasta (SCHILDER & SCHILDER 1938)
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INTRODUCTION
This study document
reviews the Bistolida-complex with specific reference to the
sp. and ssp. along the north-eastern, eastern and southern coastlines
of Africa and the adjacent islands.
Some
three decades ago in 1977, C.M. Burgess (1977) questioned the validity
of some Bistolida sp. on the grounds of conchological similarities
or apparent integrades (such as hirundo and owenii; brevidentata
and stolida; erythraeensis and stolida; hirundo
and neglecta). Much water has flowed under the bridge since
then. One of the Bistolida sp. has since been split: stolida
diauges gave rise to the sp. diauges and the ssp. stolida
clavicola. Two new Bistolida ssp. were described: stolida
brianoi and stolida uvongoensis.
As
regards the Bistolida of Africa and its adjacent islands,
several new localities have been identified which extend the peviously
known distribution ranges. Possible integrades, forms and variants
of the sp. and ssp. are now available for illustration and study.
The status of the Indian Ocean's far north-western stolida specimens
as true stolida stolida or as a ssp. or form of it, could
be debated.
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Bistolida
(COSSMANN 1920)
Shape:
Short-cylindrical to elongate-oval.
Teeth: Rather strong, often numerous,
often extending across the base.
Sides: Usually finely spotted.
Dorsum: Nearly always with a bluish tint
and irreglar blotches, patches or banding.
Range: Red Sea and Indo-Pacific.
Animal: The mantle is whitish, densely
covered with dendritic, short, class 4 papillae. The siphon extremity
is fringed. The protoconch with 3 - 4 whorls indicate a planctotrophic
larval development. Radula is of the "stolida" type.
The bursa copulatrix is present.The teeth extend onto the base. The
fossula is marked and grooved. |
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stolida
stolida (LINNAEUS 1758)
The sp. has
an extensive presence in the Pacific region.[Plate
1] Though the bulk of specimens released onto the shell collector
market originate in the Philippine Islands, the range is far more
comprehensive and includes some far-flung localities such as the Line
Islands (1,610 km to the south of Hawaii), Okinawa Island in Hawaii
(in the channel between the main island and a small coral island off
Torii Station), Vietnam, various reefs in the South China Sea, the
R.O.C. (Taiwan), Hainan Island in China, Fiji, Tonga, Vanuata, New
Britain, Guam, the Solomon Islands, the Chagos Archipelago, the Mortlock
Islands in Papua New Guinea, the Andaman Sea off Western Thailand,
South West Thailand, Aru Island in the Moluccas, Ceram in Indonesia,
the Bidari Group in Malaysia, the Majuro and Ponape Atolls in the
Eastern Caroline Islands, North West Australia, the reefs of Queensland
in Australia and the island group of Wallis and Futuna lying about
a third of the way from New Zealand to Hawaii.
The
range was originally given by Lorenz & Hubert (2000) as the
Indo-Pacific but Lorenz subsequently restricted it to the Western
and Central Pacific. More recently he referred to the "range
of the Pacific stolida and the western clavicola"
as being interrupted by stolida rubiginosa (GMELIN 1791)
in the north-eastern Indian Ocean.
However,
stolida (whether it be s. stolida or stolida
var.) is known to occur in the Indian Ocean region. [Plate
2] [Plate
3] In the Red Sea, specimens were found in a shipwreck off the
Sinai Penninsula. Heiman is doubtful that the Sinai Peninsula is
a true locality for Bistolida stolida. He maintains that
the finds of specimens are sporadical and doubtful which indicate
a mere possibility that the sp. is really living in the Gulf of
Aqaba. On the far northern African coast it is found at Masirah
Island in Oman and Massawa in Eritrea. It has also been found on
Mahe Island in the Seychelles and Helengeli Atoll in the Maldives.
Liltved recorded the most southern locality as Scottburgh in the
KwaZuluNatal Province of South Africa but this might refer to either
diauges, s. clavicola or the recently described s.
uvongoensis. This misidentification might also apply to other
finds of purported s. stolida as reported in Hawaiian Shell
News: Hudhuveli and Kanifinolhu Islands in the Maldives, Fumba on
Zanzibar, Tanzania and at Shimoni in Kenya.
The
habitat of s. stolida is the intertidal zone up to 25 m depth.
The habitat appears to be invariably a hard reef with stone and
coral heads. Saltzgaver studied the s. stolida of Guam and
found deep-water specimens in a habitat of dome-shaped flat-bottom
coral heads on a gently sloping bottom or in piles of smooth round
rocks on a hard sloping surface. The bottom cover is usually short
algal growth. In shallow-water he found specimens in protected areas
such as bays and harbours, usually beneath slabs of dead coral,
the bottoms of which are covered with sponge growths and other material.
A
photo of a live specimen in New Caledonia shows the animal has a
pale yellow mantle. In Guam it shows a thin mantle of light tan
to grey colour that reveals the shell. There is a big variation
in colour of the animal itself: lemon to tan (Manilla), grey (Subic
Bay in the Philippine Islands), lemon-pulp yellow (Dampier Archipelago),
whitish-grey (northern Queensland) and red-brown (Kwajalein). Burgess
describes the papillae as closely spaced, some spike-like with rose-thorn
type branches. The siphon is fringed with long fingerlike processes
and the tentacles are dull yellow-orange.
The
shell of s. stolida is oval to cylindrical, with a flattened
dorsal dome and rather callous and ranges in length from 15 - 36
mm. The teeth are strong. The columellar teeth are long, pale to
brownish in colour. The dorsum is bluish and the base is yellowish.
As for markings, the margins and tips are spotted with 4 blotches
on either side. The marginal blotches usually connect with the compact,
dark dorsal blotch.
By
comparison, the s. stolida and stolida var. of the
Indian Ocean might or might not have marginal blotches and when
present, these blotches frequently do not connect with the dorsal
blotch. The immaculata pattern variation with an absence
of dorsal and marginal blotches have been collected in the Seychelles
and Oman. In some specimens the teeth are distinctly short and thick.
The shape is sometimes quite angular (reminescent of s. crossei)
and sometimes pyriform like that of diauges.
Amongst
the specimens that have been collected in the Maldives and Eritrea
[Plate 4]
are some with red-stained teeth and dorsum patterns resembling s.
rubiginosa. These are not the true s. rubiginosa. Similar
s. rubiginosa-look-alikes, specifically as regards the red-stained
teeth, can be found amongst s. stolida populations in the
Philippine Islands and Guam.
The
Indian Ocean specimens deserve further study to determine whether
they are merely interesting forms or variations of s. stolida
(as has been recorded elsewhere, e.g. those resembling s. crossei
amongst the s. stolida population of the Philippine Islands)
or could be acknowledged as a ssp. of s. stolida. It would
also be of interest to compare specimens taken in shallow water
with specimes from deeper water (10 m +) to note whether differences
in size and shape of shells could be related to water depth and
bottom conditions as Saltzgaver(1974) suggests. These Indian Ocean
shells are not commonly found and to gather a sufficient number
from the region for a detailed study, will be difficult.
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stolida
clavicola (LORENZ 1998)
Despite the
similarities of the animals, the differences in the shells offered
sufficient justification for Lorenz & Hubert (2000) to split diauges
to create a new Bistolida ssp. alongside it: stolida clavicola.
The ssp. is the conchological link between the east African diauges
and s. stolida from the Philippine Islands.
The
shell ranges in length from 22.8 - 34.9 mm, is oval and callous
with a rounded dorsal dome. The teeth are not tinted and abruptly
shorten on the anterior columellar side. The base is plain white.
The dorsal blotch appears perforated and is seldom connected with
the marginal blotches. The marginal blotches can be substantial,
stretching far along the margins. When the dorsal and marginal blotches
do connect, the dorsal pattern is inevitably bold and dramatic.
An immaculata form specimen has been collected in South Africa.
[See specimen
#2475 on Plate 8]
One
diver reports that when a live specimen is taken, the colouration
on the posterior of the dorsum is so deep that it will obscure the
blotch. Out of water, the colour fades rapidly to reveal the dorsal
marking.
The
locus typicus is Nungwi Village in the north of Zanzibar, Tanzania.
Specimens were subsequently reported from western Mauritius, Mombasa
in Kenya, Nosy Be in Madagascar, Massawa in Eritrea and Mogadishu
in Somalia. Specimens were also taken off the coast of the KwaZuluNatal
Province of South Africa on the Aliwal Shoal (off Park Rynie) and
off Pumula.
[Plate 5]
[Plate 6]
[Plate 7]
[Plate 8]
[Plate 9] [Plate
10]
The
preferential habitat is narrow crevices in exposed, shallow reef
areas formed by colonies of the scleractinian coral Pavona clavus.
In the Seychelles, Jarret (2000) observed that s. clavicola
favour sandy areas, usually under pieces of coral on sandy parts
of reef. Along the coast of South Africa, specimens have been found
2 km off the shore at 20 m and as deep as 44 m on a low profile
reef. On Aliwal Shoal the ssp. has been found along with scurra
scurra (GMELIN 1791).
Little
has been published on the animal. Jarret (2000) describes the mantle
as being extremely thin and pale fawn-grey in colour with the foot
having a similar colour. Smith observed the mantle to be near translucent.
A live specimen was photographed at Kosi Bay by dr. Dai Herbert,
Head of the Department of Mollusca at the Natal Museum, Pietermaritzburg.
[Plate 11]
The Herbert specimen (Natal Museum NMSA S1748) was found on
a coral reef at a depth of 20 - 24 m. The photo shows a mantle that
is fawn-grey and it is densely covered with thin, short papillae.
The foot is pale yellow. The tentacles are yellow, tapered from
base to tip. The siphon is fawn-grey and appears to have fine brown
speck and is fringed with short, thick processes.
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stolida
brianoi (LORENZ 2002)
Though strongly
reminescent of brevidentata deceptor (IREDALE 1935) in size,
shape, colour and the character of marginal spotting but because of
its geographic proximity to s. clavicola, dr. Lorenz has provisionally
assigned s. brianoi to the Bistolida. It is the conchologial
intermediate between stolida and brevidentata.
The
shell is oval and lengths of 22.2 - 27.9 mm have been recorded.
The extremities are slightly rostrate with the posterior extremity
elegantly margined and solid and the anterior extremity fragile,
margined on both sides and pointed. The margins are white and reveal
numerous, very small and distant brown spots. Each margin has two
stripes and these divide the shell into three equal proportons.
The labral margin is low and rounded whereas the columellar margin
is callused and bent up. The narrow aperture is curved and equally
wide throughout. The teeth are pronounced on both side. On the columellar
side the teeth are sligtly finer and restricted to the aperture.
The labral teeth extend across the middle of the lip. The plain
white base is callused and convex. The tips show pale brown terminal
spots. The dorsum is a greenish-cream colour with a square-shaped
brown blotch. [Plate
12]
The
animal has not been studied and described. A very limited number
of shells, usually found dead, have been collected from south-eastern
Madagascar. The habitat is probably the upper sublittoral zone.
It is not known if s. brianoi occurs sympatric with s.
clavicola and diauges found in western Madagascar.
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stolida
uvongoensis (MASSIER 2004)
Massier (2004)
has described a new Bistolida ssp. unique to the KwaZuluNatal
coast of South Africa, known specifically from the mouth of the Tugela
River in the northern part of the province to the central Transkei
region in the Eastern Cape Province. Uvongo/Shelly Beach is the locus
typicus. Beach specimens have been collected at Mtwalumi, Aliwal Shoal
and Durban Bay by Rina Matthee and at Richards Bay (the furthest northern
locality) by prof. Douw Steyn. Live specimens were found off Pumula
and Scottburgh by Valda Fraser and Martin Wallace.
Though
only 16 specimens (only one of them taken live) were available to
Massier for study, the material presented a shell that differs significantly
from its relatives s. clavicola, s. brianoi and
diauges. More beached specimens, a few fresh-dead specimens
and only three known to be live-taken, have since been identified.
Because of restrictions on taking any live specimens along the coast
of the KwaZuluNatal Province, it is highly unlikely that many fresh
specimens of the ssp. showing full gloss and markings, will become
available to collectors.
In
s. uvongoensis the shell is elongate-cylindrical with the
dorsum gently and evenly curved, showing no dorsal plateau. The
lengths of the holotype and paratypes vary between 22.8 - 29.6 mm.
The beige to brown anterior and posterior terminals are well margined
on both sides and slightly rostrate. [Plate
13]
The
labral margin is bent up along the entire length and visible when
the shell is viewed dorsally. The columellar margin is not well
produced and is rounded. The umbilicus which is partly covered by
a brown blotch, is deep and completely covered with callous. The
aperture is almost straight, gently curved towards the posterior
end and of the same width throughout. The base has a white to off-white
colour and is covered with callous in adult specimens.
The
columellar teeth are a distinctive feature. They are short along
the entire length of the aperture and continue only a few mm onto
the base. They also continue into the aperture, covering the well
produced fossula. The labral teeth are stronger than the columellar
teeth and slightly cover more than half of the labrum. A few teeth
towards the posterior terminal reach the margin. The teeth are not
stained.
Another
distinctive feature is the marginal spotting. The spots are extremely
small and densely set. The spotting continues onto the dorsum but
does not reach the centre. The dark colour of the spots gives the
margins, especially the labral margin, a much darker appearance
than any of the other ssp.
The
shell colour is unique greenish-beige with a very large, dark brown,
center blotch. The two labral blotches are very small and are positioned
at one third and two thirds of the margin and these do not connect
with the dorsal blotch.
A
live specimen was photographed in 2004 by Fraser. [Plate
14] The photo shows a mantle that is fawn-grey with numerous
dendritic papillae. The foot is of the same colour, somewhat transparent.
The tentacles are lemon-yellowish, tapered from base to tip. The
photos do not reveal the siphon nor the animal.
The
Fraser specimen was found under a loose stone in an area of scattered
reef 5 km off shore at a depth of 28 m. Another live specimen was
seen by Fraser and her diving partner Wallace along a ridge quite
close to shore at the same depth. The second specimen was under
a piece of broken smooth-horned type coral. Fraser and Wallace observed
that other Cypraeidae found in the same localities included
chinensis violacea (ROUS 1905), ziczac misella (PERRY 1811),
cribraria comma (PERRY 1811) and owenii vasta.
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diauges
(MELVILL 1888)
The shell is
distinctively elongate pyriform with rather long, untinted teeth throughout.
The colour of the dorsum is light blue to grey to greenish with fine
brown spots. Two pale bands are revealed. The dorsal blotch is compact.
The marginal blotches are greatly reduced and rarely connect with
the dorsal blotch. The base colour is yellow-ish. [Plate
15] [Plate
16] [Plate
17] [Plate
18] [Plate
19]
The
east African habitat is intertidally in muddy water with seaweed.
In South Africa it occurs in deeper water. Woodcock collected live
specimens at Mafamete Island in Mozambique at a depth of 1 m under
large, dead coral slabs at the edge of a minor reef. Lorenz (2002)
reports the sympatric occurrence of s. clavicola and diauges
at Pemba Island off Zanzibar and the mainland Tanzania coast.
No
detailed notes on the animal have been published. Lorenz says it
has a grey, fluffy mantle and a white foot with a black line on
top. Woodcock reports short, small, dense papillae on a closed mantle
with pale, dull, lemon-coloured tentacles. He also reports that
the animal does not appear tmid in the least. After harvesting the
very dark shell soon fades to the typical specimen colour.
The
distribution of diauges ranges from the southern Red Sea
to the Western Indian Ocean. The figured syntype in Melvill's description
and the other syntypes in the Melvill-Tomlin Collection of the National
Museums and Galleries of Wales, most certainly all identifiable
as diauges, are said to have originated in Mauritius. [Plate
20] This is confusing because no specimens are known from Mauritius.
The accuracy of Melvill's typicus locus must hence be considered
suspect. The attribution by Allan (1956) of Natal (South Africa)
as typicus locus for diauges is also not accurate. The sp.
is known to occur on the KwaZuluNatal coast of South Africa but
finds of specimens are rare. The description by Lorenz & Hubert's
(2000) of the typicus locus as extending from deep water in South
Africa to the (southern) Red Sea, is accurate.
Specimens
with no dorsal blotch and marginal blotches are frequently collected
in Tanzania. [Plate
21] they merely show the fine brown spotting on the dorsum.
Some specimens reveal a vestige of the marginal blotches on the
base of the shell. This is the immaculata form of diauges.
In the past this form of diauges was described as s. fluctuans
(IREDALE 1935) because of the supposed similarity with the Australian
brevidentata in shape, colour and dentition. (Lorenz &
Hunert (2000) maintain these similarities consitute a link between
the Pacific Ocean sp. and Indian Ocean ssp. Woltz and Belcher (undated)
noted that the shell is slightly more ovate than in diauges,
has a length range of 17.8 - 25.4 mm and has a rare occurrence at
Tiles and Fungu Mkadya in Tanzania. This form has also been reported
from Diani Beach in Kenya. The fluctuans description should
be restricted to the brevidentata. The correct description
of these African coast specimens is diauges form immaculata.
Melvill
also described stolida var. gelasima (MELVILL 1888)
with typicus locus Ceylon (Sri Lanka). [Plate
22] He mentions the delicate pale brown spotting, absence of
the dorsal blotch and pale fawn extremities which are known features
amongst the immaculata forms of both diauges and s.
clavicola. Melvill notes that the teeth of gelasima are
as in diauges but the dorsum is flattened and that the pale
olive green dorsum of gelasima matches the colour description
of diauges.
Melvill's
notes which detail important similarities with diauges in
conjunction with photographs of the figured syntype of gelasima,
can leave no doubt that gelasima is nothing other than a
specimen of diauges form immaculata. Melvill's typicus
locus of Sri Lanka is improbable. No diauges has been recorded
to have been collected there. The above justifies that gelasima
be a synonym of diauges rather than s. stolida.
A
third Bistolida description by Melvill is stolida
var. moniontha (MELVILL 1888) and he introduces it as "A
common form, almost a sub-species" and an "
intermediate
between the type [stolida] and C. brevidentata".
[Plate 23]
No locality data was provided for the syntypes but one may assume
the specimens were collected in an Indian Ocean region where diauges
does occur. His description is of a shell that is not humped
but has a rounded dorsal surface. Specimens have a single rounded
blotch that is occasionally absent. The sides are streaked and speckled
and the teeth are not prolonged and white. The length of the largest
specimen studied by Melvill is 3.175 cm.
From
a study of photos of the figured syntype of moniontha, no
evidence can be found that it differs significantly from diauges.
The shape of the shell, the structure of the teeth and the description
of the dorsal and labral markings match those of diauges.
It must therefore be considered a synonym of diauges and
not of brevidentata.
Apparent
integrades of diauges and s. clavicola
In Somalia, diauges occur along with s. clavicola but
it is not clear from the available collection data whether the sp.
and ssp. share the exact localities. The same lack of collection
data of shared localities applies to South Africa where the three
Bistolida sp. and ssp. (diauges, s. clavicola and
s. uvongoensis) occur. Specimes that show apparent integrades
have been collected from Somalia and South Africa. [Plate
24] The shell shape is intermediate between the sp. and spp.
but in colour and markings, more reminescent of diauges.
In the two illustrated specimens from Somalia, the columellar teeth
show the abrupt shortening on the posterior side that is associated
with s. clavicola.
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erythraeensis
(SOWERBY 1837)
Summarised from
descriptions by Lorenz & Hubert (2000) and Heiman (2002). An elyptical
to cylindrical shell. The extremities are acuminate and tinged with
orange to brown. It has a grey-bluish dorsum revealing three transverse
bands (of which the central band is most prominent), dark small spots
and a dark blotch. The blotch is not centered and has an irregular
form and is mostly positioned towards the posterior end of shell.
Specimens with a suffused dorsal pattern are known.
The
base is slightly convex and ranges from white to grey with many
red-brown to orange-brown spots. The margins are high but not callused
and finely spotted with blotches on the tips. The right margin is
slightly angled and visible from the dorsum. There is a complete
absence of the marginal blotches associated with other Bistolida
sp. and ssp.
The
teeth extend far onto the base. The labral teeth are well developed.
The dental interstices are stained and this is especially visible
in fresh specimens. The columellar teeth resemble plications (folds)
and reach more than halfway across the base and are never shorter
in the anterior part (as is the case with s. clavicola).
Heiman notes that the southern Red Sea population (as studied by
M. Schilder, 1967) differs in the number of columellar teeth from
the Siani population.
In
length the shell varies from 10 - 29 mm. Gigantism, with shells
reaching a length of more than 32 mm, occurs.
The
typical Red Sea habitat, according to O'Malley (1971), is in calmer
water, under rock slabs amidst a short, seaweed type of plant growth.
An alternative habitat noted by him is on the hard reef, under the
base of a type of finger coral. The northern Red Sea populations
are found in very shallow water of about 1 m. The southern populations
prefer a depth of 5 - 15 m.
O'Malley
also described the animal characteristics in great detail:
Mantle: Transparent gray-white with numerous white papillae. Eyes:
Black. Area around eyes also black, fading to light gray a few mm
from the stalks. Tentacles: Light orange-red, tapered from base
to tip. Proboscis: Light orange-red, with rounded end. Siphon: Transparent
gray-white bordered with small white filaments about 2 - 5 mm long.
Body: Light creamy-white.
Foot:
Overall colour is dirty white on crawling area. Top of foot has
a distinctive blotch at the anterior and posterior portions. Anterior
portion has a white band or margin approximately 0.5 to 1mm in thickness,
running across the width of the foot. When viewed from the top,
these resemble two white appendages. Posterior portion has a very
distinctive black stripe approximately 0.75 - 1.5 mm thick running
from the edge of the posterior portion of the foot up the body of
the animal. This stripe is made up of small black blotches. According
to Woodcock, the animal is capable of autotomy. A specimen collected
by him shed a 4 - 5 mm portion of the posterior end of the foot.
Autotomy is the ability to cast off a part of the body as a means
of defence. The cast-off part remains wriggling and writhing for
some time, distracting predators and allowing the animal to escape.
The lost part can be regenerated. Amongst sea slugs, Discodoris
lilacina, berthella martensi and Phyllodesmium magnum
are known to practise autotomy.
Supplementary
to the above, Yaron (1978) observed in his study of five living
specimens collected at Na'ama (Marsa el Et) in the Sinai region
of the Red Sea:
"The mantle is thin and the shell dorsal pattern clearly shows
through it
The foot and the serrated siphon are also grayish
white, while the crawling surface of the foot is somewhat lighter
in colour. The tentacles are a bright orange-yellow, and strongly
contrast with the general grayish-white tones of the animal."
The
north-eastern border of its distribution is the northern Red Sea.
Specimens have been collected on the Red Sea coastlines of Israel
and Jordan; at Safaga in Egypt, Jeddah in Saudi Arabia, Sharm el-Shaik
on the Sinai Peninsula, the Dahlak Archipelago off Eritrea, Aden
in Yemen, along most of the coastline of Djibouti, the Gulf of Tadjoura
off Djibouti and the Tiran Straits. Burgess (1985) is of the opinion
that the Dahlak Archipelago is probably the centre of the population.
[Plate 25]
The
most southern locality is Zanzibar, Tanzania where finds have been
scarce. Lorenz (1998) documented three specimens from Zanzibar and
a fourth is in the collection of the author. These have an absence
of dark dorsal markings and the interstices of the teeth are not
stained dark. The accuracy of records of two specimens from the
northern coast of Madagascar and one specimen from Park Rynie on
the KwaZuluNatal coast of South Africa, can not be verified. [Plate
26]
Lorenz
(1992) found fossil eryhthraeensis from the Pleistocene in
the hills behind Magawish Village in Hurghada in Egypt. These include
erythraeensis which are identical with living representatives
and the ssp. erythraeensis cepaformis (LORENZ 1992). The
latter has thin but rostrated extremities, a rather curved aperture,
shorter teeth which extend only slightly towards the base and rounded,
callous margins which are bent up. The dorsum is irregularily blotched
with faint and scarce marginal spotting.
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hirundo
hirundo (LINNAEUS) 1758
The sp.
occurs over a vast range in the Indo-Pacific. There is a claim that
the most northern range is Eilat in the Gulf of Aqaba. This is a doubtful
locality according to Mienis (1974) and Heiman (2002) also questions
the presence in the Sinai region from where no specimens were found
found during the past 30 years. Tanzania appears to be the furthest
southern range. It is known to occur in the Maldives, Mauritius, Seychelles
and Massier has a specimen from Reunion.
The
shell is oval to cylindrical with often pronounced extremities with
all four corners of the extremities marked with a dark blotch. The
teeth are fine and extend over most of the base. The dorsum has
three blue to grey zones and is spotted with brown and often blotched.
The margins are finely spotted. The length varies from 8 - 22mm.
The habitat is in coral reefs, intertidally to 10 m. [Plate
27] In the Seychelles, h. hirundo and k. kieneri
have been recorded to share same habitat near the reef under small
coral fragments with brown algae on top at a depth of 0.5 m.
Burgess
(1985) describes the animal as lemon-pulp yellow. Papillae are either
short, thick and white to bright yellow in colour or long, dendritic
and pure white. The tentacles are coloured a bright, deep orange.
The fringed siphon is speckled with black. The foot is lemon yellow
and discretely studded with black specks.
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hirundo
francisca (SCHILDER & SCHILDER 1938)
This ssp.
is rare throughout its distribution region that extends from the east
coast of Africa to Mauritius, Seychelles and Maldives. Cailliez (1981)
reported a find of five specimens with big brown blotches similar
to saulae saulae (GASKOIN 1843), at Bird Island, north of the
Seychelles, in 1981. Massier has a specimen from Barros Island in
the Amirantes Archipelago, west of Seychelles.
Woodcock
collected the ssp. at Anse aux Pins on Mahe Island in the Seychelles:
"
in shallow water (2 - 3 m) at low tide under large dead
coral blocks on the inside edge of the reef barrier. I first found
four live specimens under one dead coral block. A complete search
of the entire area produced three more specimens found 10 - 20 m
from the original spot. Returning twice over the next few days and
spending up to 5 - 6 hours at a time searching in the same general
area proved unsuccessful, not even a dead shell or fragment could
be found. I suspect that this shallower water location was not the
normal habitat of these shells. In 1998-1999 an Indian Ocean temperature
fluctuation had badly bleached 75% of the Seychelles coral reefs.
Many coral species had been killed with only the largest and hardiest
species partially surviving. These cooler temperature fluctuations
may have forced the animals into the warmer shallower waters inside
the reef barrier."
The
shell, ranging from 10 - 21 mm in lengt, is narrow, oval-cylindrical.
The teeth are fine and long. Allan notes that the labral side is
distinctly margined and the fossula is concave. The absence of the
blue colour component is specific to this ssp. and the dorsum reveals
an orange colour. [Plate
28] (Specimens from the Maldives do show a blueish dorsum.)
It
is possible that the mainland specimens could differ from those
from the Seychelles which is the locus typicus. Lorenz (1998) noted
differences when examining a 22.5 mm specimen from Nungwi, Zanzibar,
Tanzania and a 20 mm specimen from Mombassa in Kenya. The specimens
showed relatively short extremities, reduced marginal spotting and
a complete absence of the typical darker colour zones of hirundo
hirundo. Lorenz lacked a sufficient number of these specimens
to come to any conclusion that there is a consistent difference
between the mainland and Seychelles specimens.
The
animal of Seychelles h. francisca specimens differ from the
mainland h. hirundo populations. Jarret (2000) describes
it as being white nearly al over. The mantle is very thin, pure
white and transparent with fairly numerous branching papillae of
a fine white colour. The siphon and the foot (ventral and dorsal)
are white. The tentacles and proboscis are bright yellow. Jarret's
description, however, could also apply to h. hirundo found
on Seychelles because he discusses colour and pattern variations
of both the sp. and ssp. under the same heading of h. francisca.
To this description Woodcock adds that the colour of the mantle
is milky white rather than pure white, that the small papillae appeared
very slightly longer than usual for a shell of this size and the
orange tentacles had dark eyes at the tips. From observations in
the Seychelles, Woodcock says that h. francisca group together
and that the animals were very timid.
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kieneri
kieneri (HIDALGO 1906)
The sp.
resembles hirundo hirundo but can easily be separated from
the former by the markedly attenuated columellar teeth.
It
is found over much of the Indo-Pacific. Along the coast of Africa,
it has a documented presence in Kenya, Tanzania, Mozambique and
South Africa and it is known from the Seychelles (Praslin, Curieuse
and La Digue Islands), Madagascar, Mauritius and). Bosch et al (1995)
recorded its presence in southern Oman.
The
sp. has a length that ranges from 8 - 24 mm. The shell is elongate-oval.
The extremities are marked with a dark blotch on each side. The
teeth are rather coarse and markedly attenuated on the anterior
columella. The base is white and the margins are densely and darkly
spotted. The dorsum reveals three blueish-green zones. [Plate
29] A variation from the Indian Ocean islands (locus typicus
Seychelles) have more numerous teeth and wide, yellow dorsal zones.
These are sometimes named "form reductesignata"
after Schilder 1924.
From
the description of a specimen from American Samoa the animal is
known to be pearl grey with a mantle so thin that the dorsal pattern
of the sell can be clearly seen. Some of the tufted papillae are
black and cone-shaped while others are white and dendritic. The
long tentacles range in colour from lemon-peel yellow to orange-yellow
and are tipped with the same but darker colour. The siphon is white
and smoothly scalloped. The foot is lemon-pulp yellow, mottled with
cream and covered with fine, evenly distributed, black specks. The
description by Jarret (2000) of specimens from Seychelles has some
similarities as regards the foot (tinged with yellow, covered with
numerous black and white spots) and tentacles (yellow) but he refers
to a transparent yellow mantle and cream and yellow papillae. David
Touitou too confirms a yellow animal in the Seychelles.
The
habitat is shallow water under algae and dead coral slabs in muddy
areas or under stones and in coral on a hard reef and have even
been found completely exposed out of the water at low tide.
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owenii
owenii (SOWERBY 1837)
The distribution
is largely restricted to the Comoros and the Mascarene Islands. southern
Indian Ocean islands of Mauritius and Reunion. Specimens are also
known from Madagascar. Mainland specimens have been collected at Merca
south of Mogadishu in Somalia, Mombassa in Kenya, Zanzibar, Tanzania
and Coffee Bay in the Transkei region of South Africa.
In
size it ranges from 8 - 20 mm. The shell is rounded and depressed.
The strong teeth reach onto the base. The margins are callous and
very densely spotted. The colour of the dorsum is blueish with small
brown spots and larger black blotches. Others show no blotches but
only mottling. In some, colour zones can be distinguished. [Plate
30]
The
habitat is in coral reefs from the intertidal fringe to aproximately
100 m. At Diego Suarez it was found in coral rubble at 25 m.
The
animal is said to resemble that of diauges (a grey, fluffy
mantle and a white foot with a black line on top).
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owenii
vasta (SCHILDER & SCHILDER 1938)
This is
listed as a ssp. by Lorenz & Hubert (2000) but the authors were
of the opinion that it could be a valid species.
The
distribution range is south-east Africa with specimens recorded
from Tanzania, Mozambique, Madagascar and South Africa. Most specimens
are ex pisce from 50 m and deeper. Fraser and Wallace found live
specimens on low profile sandstone reefs with coral, seaweed and
algae at a depth of 15 to 30 m.
Shells
range from 12 - 27 mm in length. The shape of the shell is oval-rhomboidal.
The margins are bent up. The dorsum has small blotches or no blotches
but only confluent mottling. Colour zones can be distinguished.
The columellar teeth are rather thick and dense.
[Plate 31]
Lorenz
(1998) described the animal. The mantle of the animal is yellow,
somewhat transparent and densely covered with 1 mm long papillae.
(Fraser, Wallace and Smith describe the papillae of specimens seen
off KwaZuluNatal as sparse, short and raised.) The very long foot
is transparent white with noticeable black longitudinal stripes.
The proboscis is yellow-orange with a dark ring arund the mouth
opening and the syphon is finely frayed. The tentacles are black
at the base and red at the tips.
The
animal is extremely shy and at the slightest disturbance withdraws
into the shell. The mantle is extremely fraile. Lorenz postulates
that the animal is capable of autotomy. Lorenz has not observed
autotomy in live specimens of owenii vasta, but have noted
amongst dead specimens the separation of part of the foot at a specific
locus.
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owenii
vasta southern form
Callussed specimens
of owenii vasta such as those from Madagascar, Mozambique (Pemba)
and the KwaZuluNatal Province of South Africa (Richards Bay, Durban,
Pumula and Park Rynie) have been dubbed the "southern form"
by Lorenz. Specimens have been taken live (intertidal and dived at
30+ m), beached, trawled (at 80 - 120 m) and ex pisce (at 40 - 60
m). [Plate
32]
The
distinguishing features are that the shells have depressed dorsums,
both margins are heavily callussed and the marginal spotting is
intense, dark and confluent. The four blotches of the extremities
are very dark. In every respect other than the dorsal spotting and
blotch, the features of owenii vasta are exagerated in owenii
vasta southern form.
The
southern form specimens share the same locality but perhaps not
the same habitat with owenii vasta. To eliminate confusion
arising from their presence in the same (southern) region, a more
descriptive form name is merited.
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Concluding
comments
The distribution
ranges of the Bistolida sp. and ssp. in the western Indo-Pacific
region in some cases far exceed the descriptions by their authors.
The original descriptions of the habitats have also been supplemented
with new information.
Specimens
from the extremes of the distribution ranges, often show differing
features that may not be sufficient to justify conchological differentiation
or at most may justify a form name, e.g. owenii vasta southern
form.
Access
to previously unexplored localities or the process of reviewing
specimens in collections, offer the chance to discover and describe
new ssp., e.g. the recently named s. uvongoensis. This might
also apply to the s. stolida-like specimens from the Sinai
Penninsula, Masirah Island, Eritrea and the Maldives but many more
specimens of these shells shells, studies of their live animals
and detailed locality data must be forthcoming to justify this.
The
debate about the sympatric occurrence of Bistolida ssp. and
sp., e.g. s. clavicola and diauges in Tanzania, continues
and has to be resolved. Is there room within the definition of what
constitutes a valid sp. and ssp., specifically as regards locality,
to accommodate sympatric occurrence even if the justification is
based only on different habitats within the same locality?
Dealers,
collectors and divers alike must be encouraged to record comprehensive
data on specimens. Data on the exact locality, descriptions of the
habitats and especially notes on the animals will add to our knowledge
and benefit all.
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Acknowledgements
Several
marine shell specialists provided valuable specimens, comments, information
and research material for this document: Dr. Felix Lorenz jun. (Germany),
Werner Massier (Namibia), Philippe Quiquandon (France), Philippe Poppe
(Philippine Islands), Ed Heiman (Israel), Mike Woodcock (Georgia)
and Val Fraser, Martin Wallace, Rina Matthee, Dawn Brink, prof. Douw
Steyn, Vellies Veldsman, dr. Dai Herbert and Gerald Smith (South Africa).
Special
mention must be made of the kind and unstinting assistance of Harriet
Wood, Collections Manager (Mollusca) at the National Museums and
Galleries of Wales, for providing research material on the Bistolida
diauges, gelasima and moniontha syntypes in the
Melvill-Tomlin Collection.
The
specimens for the accompanying plates, except where indicated otherwise,
were photographed by Stephan Veldsman. Val Fraser and dr. Dai Herbert
(South Africa) respectively photographed the live spcimens of stolida
uvongoensis and stolida clavicola. James Turner, Biological
Photographer (Mollusca) at the National Museums and Galleries of
Wales photographed the Melvill syntypes.
The
author invites comments. You may contact Ronnie Watt by e-mail at:
clumber@telkomsa.net
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|
Tables
of Differences
Sources: Lorenz.,
F. & Hubert, A., 2000: A Guide to Worldwide Cowries - 2nd Revised
and Enlarged Edition - ConchBooks, p.167-168, 170, 172, 174 &
Massier, W., 2004: Description of a new subspecies of Bistolida
stolida (LINNAEUS 1758) (Gastropoda: Cypraeidae)
Schriften zur Malakozoologie, XXI, Cismar, pp. 35-38 |
|
Table
Of Differences 1 (Plate
A)
|
|
stolida
stolida
|
stolida
clavicola
|
stolida
brianoi
|
stolida
uvongoensis
|
diauges
|
erythraeensis
|
|
Shell
shape
|
oval-cylindrical
|
elongate-cylindrical
|
oval
|
elongate-cylindrical
|
elongate
|
elyptical-cylindrical
|
|
Dorsal
colour
|
blueish
|
bluish-grey
|
greenish-cream
|
greenish-beige
|
greenish-grey
|
grey-blueish
|
|
Columellar
teeth
|
long,
pale to brownish
|
extend
to middle of base posteriorly, restricted to aperture anteriorly
|
very
short, restricted to aperture
|
short,
slightly encroaching onto base
|
extend
to middle of base
|
resemble
plications, reach more than halfway across the base, never shorter
in the anterior par
|
|
Dorsal
blotch
|
dark
brown, compact
|
perforate,
generally small, pale to dark brown
|
square-shaped,
brown
|
large,
centered, dark brown
|
compact,
small to large, pale to dark brown
|
dark
brown, irregular, usally towards posterior
|
|
Marginal
blotches
|
2
blotches on each side, connecting with dorsal blotch
|
clearly
defined, seldom connecting with dorsal blotch
|
indistinct
stripes
|
small
and indistinct
|
small
to absent
|
absent
|
|
Marginal
spotting
|
very
variable from feint and sparse to dark and crowded
|
small,
distant, very light beige
|
small,
distant, brown
|
small,
numerous, very crowded, dark brown
|
small,
distant, beige to brown, showing white margins
|
many,
red-brown to orange-brown
|
Table
Of Differences 2 (Plate
B)
|
|
hirundo
hirundo
|
hirundo
francisca
|
kieneri
kieneri
|
|
Shell
shape
|
elongate-cylindrical
|
oval-clindrical,
narrow
|
oval-cylindrical
|
|
Dorsal
colour
|
pale
blue
|
orange
|
greenish
to yellowish
|
|
Teeth
|
fine,
extend over base
|
fine
and long
|
coarse,
extend onto base except at anterior
|
|
Dorsal
pattern
|
3
blue-grey zones often blotched
|
absence
of colour zones, with big, brown blotch
|
3
blue-grey zones sometimes with small blotches
|
|
Marginal
spotting
|
fine
|
few
|
dense,
dark
|
Table
Of Differences 3 (Plate
C)
|
|
owenii
owenii
|
owenii
vasta
|
owenii
vasta southern
form
|
|
Shell
shape
|
oval,
narrow
|
rhomboidal
|
rhomboidal,
callous
|
|
Dorsal
colour
|
blueish
|
dark
green-bluish
|
dark
green-bluish
|
|
Teeth
|
strong,
distant, extend far onto base
|
thick
and dense
|
thick
and dense
|
|
Dorsal
pattern
|
mottling
with or without a blotch
|
dark
blue-grey zones with or without very small blotches
|
dark
blue-grey zones with mottling and/or perforated blotches
|
|
Marginal
spotting
|
dense
|
confluent
|
intense,
dark and confluent, extending deep onto base
|
PLATES
PLATE
1: Selection of stolida stolida (LINNAEUS
1758) from various localities.
PLATE 2:
stolida var. (LINNAEUS 1758)
PLATE 3: stolida
form immaculata (LINNAEUS 1758)
PLATE 4: stolida
var. (LINNAEUS 1758) "red tooth"
PLATE 5: stolida
clavicola form immaculata (LORENZ 1998) - Madagascar
PLATE 6: stolida
clavicola (LORENZ 1998) - Kenya
PLATE 7: stolida
clavicola (LORENZ 1998) - Mauritius and Mozambique
PLATE 8: stolida
clavicola (LORENZ 1998) - Somalia
PLATE 9: stolida
clavicola (LORENZ 1998) - South Africa
PLATE 10: stolida
clavicola (LORENZ 1998) - Tanzania
PLATE 11:
Live specimen of stolida clavicola (LORENZ 1998)
Photography by dr. Dai Herbert, Head of the Department of Mollusca at
the Natal Museum,
Pietermaritzburg, South Africa.
PLATE 12:
stolida brianoi (LORENZ 2002)
PLATE 13: stolida
unvongoensis (MASSIER 2004)
PLATE 14:
Live specimen of stolida uvongoensis (MASSIER 2004)
Photograph by Valda Fraser, South Africa.
PLATE 15:
diauges (MELVILL 1888) - Tanzania
PLATE 16: diauges
(MELVILL 1888) - Kenya
PLATE 17: diauges
(MELVILL 1888) - Madagascar
PLATE 18: diauges
(MELVILL 1888) - Reunion, Seychelles and Somalia
PLATE 19:
diauges (MELVILL 1888) - Mozambique and South Africa
PLATE 20:
Melvill's diagues syntypes (MELVILL 1888)
Two of the syntypes in the Melvill-Tomlin Collection. Photography by James
Turner,
Biological Photographer (Mollusca) at the National Museums and Galleries
of Wales.
PLATE 21:
diauges form immaculata (MELVILL 1888) - Tanzania
PLATE 22:
Melvill's gelasima syntype (MELVILL 1888)
From the Melvill-Tomlin Collection. Photography by James Turner, Biological
Photographer (Mollusca)
at the National Museums and Galleries of Wales.
PLATE 23:
Melvill's moniontha syntype (MELVILL 1888)
From the Melvill-Tomlin Collection. Photography by James Turner, Biological
Photographer (Mollusca)
at the National Museums and Galleries of Wales.
PLATE 24:
Apparent integrades of diauges (MELVILL 1888) and stolida clavicola
(LORENZ 1998)
PLATE 25: erythraeensis
(SOWERBY 1837) - Southern
PLATE 26:
erythraeensis (SOWERBY 1837) - Northern
PLATE 27:
hirundo hirundo (LINNAEUS 1758)
PLATE 28:
hirundo francisca (SCHILDER & SCHILDER 1938)
Photography by Mike Woodcock
PLATE 29:
kieneri kieneri (HIDALGO 1906)
PLATE 30:
owenii oweni (SOWERBY 1837)
PLATE 31:
owenii vasta (SCHILDER & SCHILDER 1938)
PLATE 32:
owenii vasta southern form (SCHILDER & SCHILDER 1938)
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