untitledTHE RAFFLES BULLETIN OF ZOOLOGY 2007
THE RAFFLES BULLETIN OF ZOOLOGY 2007 55(2): 355–362 Date of
Publication: 31 Aug.2007 © National University of Singapore
COSTASIELLA CORONATA, NEW SPECIES, AND A REVISED DIAGNOSIS FOR THE
FAMILY COSTASIELLIDAE
(MOLLUSCA: OPISTHOBRANCHIA: SACOGLOSSA)
C. Swennen Faculty of Science and Technology, Prince of Songkla
University, Pattani 94000, Thailand
Present address: Doolhof 7, 1792CM Oudeschild, The Netherlands
Email:
[email protected]
ABSTRACT. - Costasiella coronata, new species, occurs on its food
alga Vaucheria sp. high in the intertidal zone of mangrove forests
bordering parts of the Gulf of Thailand. It differs from all
previously described Costasiella species by having fl attened
cerata and a large renal shield. The external characteristics and
the internal anatomy of the digestive and reproductive organs are
described and depicted, and notes are given about reproduction and
behaviour. The characteristics of the genus Costasiella Pruvot-Fol,
1951 have gradually been changed in the course of the years and
have become restricted to ceratal-bearing sacoglossans with
admedian eyes and digitiform or auriculate rhinophores without
diverticula of the digestive gland. The described species of
Costasiella are listed. The original characteristics of the family
Costasiellidae, as given by Clark (1984), appear to include specifi
c characteristics of only few species. This inadvertently left out
the type species of the genus Costasiella , which is the type genus
of the family, and several others. Therefore the diagnosis is
shortened and changed to include all species.
KEY WORDS. – Gastropoda, Opisthobranchia, Sacoglossa,
Costasiellidae, Costasiella, Thailand.
INTRODUCTION
All ceratal-bearing sacoglossans with the eyes situated close
together between the rhinophores are usually considered to belong
in the genus Costasiella Pruvot-Fol, 1951, within the monotypic
family Costasiellidae Clark, 1984. Unfortunately the status of the
genus and several of the species it contains is disputable. Luckily
it concerns only 15 taxa, but several of them have been poorly
described. The few described species differ in fundamental details,
which forced Marcus (1982) and Jensen (1990) to suggest that the
genus should be split up, but they did not provide any specifi c
proposal.
This paper gives an extensive description of a Costasiella from the
Gulf of Thailand. The species could not be identifi ed with any of
the previously described species and is considered new to science.
While reviewing the existing descriptions of taxa, it was found
that some descriptions were vague and/or likely to contain
mistakes. A check of the original diagnosis of the family
Costasiellidae, as given by Clark (1984), showed that it contains
characteristics that only hold true for a few Costasiella species.
This excludes several species, among which is even the type of
Costasiella. Therefore, the family diagnosis is adapted to include
all known species.
MATERIALS AND METHODS
Specimens of the new species were collected by hand from the mud
and from their food algae during low tide. A number of specimens
were narcotised with MgCl2 in small Petri dishes in the laboratory.
Specimens for dissection were temporarily stored in a solution of
formaldehyde-seawater with 5% acetic acid. Dissections were
undertaken by hand with the help of fi ne needles. Specimen was
kept in water in a Petri dish under a 4-64x stereo microscope,
notes and drawings of details were made, while some images were
taken with a digital camera. For permanent storage, some specimens
were stored for a few days in formaldehyde and thereafter
transferred into 70% ethanol. Specimens were kept alive for about
1–2 weeks in trays with mud, Vaucheria sp. and local seawater with
and without aeration in October 2003.
Costasiella coronata, new species (Figs. 1, 2)
Material examined. – Bang Tawa, in the mangrove forest (6°51'N
101°09'E), some specimens and egg-strings, 10 Jul.2000. Same site,
some tens of individuals with a length up to 7 mm, also egg-strings
found, Oct.2000. Same site, some tens of individuals with a length
of 4-10.5 mm, also egg-strings found, 13 Oct.2003.
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systeem
Note
Swennen: Costasiella coronata, new species
Fig. 1. Costasiella coronata, new species: A, outline of a
submerged live specimen; B, outline of a live specimen on mud
during low tide; C, different shapes of the anterior part of a live
specimen; D, outline of a cross section through the cardiac area of
a relaxed specimen; E, ventral view of a relaxed specimen showing
organs shining through; F, branch of the digestive gland in ceras;
G, outline of cross section of a preserved ceras; H, branches of
the albumen gland in a series of cerata of different sizes; upper
row dorsal sides, lower row ventral sides; I, dorsal view of the
anterior body of a preserved specimen; the renal shield and cerata
are removed; former position of renal shield is indicated by
slashes; J, the genital apertures at the right anterior side of a
preserved specimen with removed renal shield; K, ventral view of
anterior area showing the mouth in a preserved specimen. Legend: ag
– albumen gland; ap – anus papilla; bl – black line; ce – ceras; dg
– digestive gland; dr – dorsal rim; fo – follicles of ovotestis; lc
– lost cerata; ma – male aperture; mo – mouth; oa – oviducal
aperture; pc – pericardium; rh – rhinophore; rp – renal pore; rs –
renal shield; st - stomach; va – vaginal aperture.
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THE RAFFLES BULLETIN OF ZOOLOGY 2007
Fig. 2. Costasiella coronata, new species: A, eyes; B, radular
tooth and connection; C, pharynx in lateral view; D, digestive
system from mouth to stomach in lateral view; ring of main ganglia
around anterior part of oesophagus is removed. E, branches of the
digestive gland in the body of a preserved specimen; dorsal skin
and branches of the albumen gland are removed; F, hermaphrodite
follicles and ductules from a preserved specimen; G, branches of
the albumen gland in the body of a preserved specimen; dorsal skin
removed; the heart and some organs of the genital complex block the
view in the outlined area; H, penis; I, egg-strings; J, arrangement
of eggs in an egg-string; K, diagram of the genital complex.
Legend: ad – albumen duct; ag – albumen gland; am – ampulla; an –
anus; bc – bursa copulatrix; bw – body wall; cc – central canal; dg
– digestive gland; ey – eyes; gr – genital receptacle; hd –
hermaphrodite duct; in – intestine; ma – male aperture; mu – mucus
gland (large oviduct); oa – oviducal aperture; oe – oesophagus; og
– oral gland; op – oesophageal pouch; ov – oviduct; ph – pharynx;
pr – prostate; sg – salivary gland; st – stomach; va – vaginal
aperture; vd – vas deferens.
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Swennen: Costasiella coronata, new species
Tak Bay, on intertidal mud bordering a small fringe of mangroves
(6°16'N 102°03'E), 2 specimens with a length of 4-5 mm, 23
Apr.2005. Both locations are in southern Thailand along the Gulf of
Thailand, and all specimens were on, or in, patches of the alga
Vaucheria sp.
The description is based on 11 specimens collected in Oct.2003, of
which fi ve were dissected and six selected as types. The holotype
and three paratypes are deposited in the Zoological Reference
Collection (ZRC) of the Raffl es Museum of Biodiversity Research,
National University of Singapore, and three paratypes in the
Zoological Museum, University of Amsterdam (ZMA).
External morphology. – Black eyes close together between
rhinophores (Fig. 1 A, C). Renal shield large, fi xed on dorsal
side of small pericardium, covering dorsum from just posterior of
eyes to two-third of body length; wide, free rim can be held
upwards and pleated (Fig. 1 A). Rim usually covers basal parts of
bordering cerata and often nearly their whole length (Fig. 1 B).
Transparent renal shield contains irregular central channel with
side-branches bordered by dispersed dark dots. Rhinophores smooth,
wide, somewhat fl attened at base, but rounded distally and
tapering into blunt tips. Shape of head varies according to degree
of straightening (Fig. 1 C); frontal part usually kept bilobed.
Lateral areas of back wider than body by thick rims (Fig. 1 D, E).
Each rim fi lled with maze of branches of albumen and digestive
glands. Cerata in a row of up to 22 on dorsal side of each rim,
starting shortly behind rhinophores; not covering central part of
dorsum and posterior end of foot; strongly fl attened, wide, and
smooth (Fig. 1 F, G); fl at sides concave-convex in preserved
specimens (Fig. 1 G). Each ceras containing wide branch with side
branches of digestive gland (Fig. 1 F) and in reproductive slugs
also branches of albumen gland being most dense on dorsal side of
ceras (Fig. 1 H).
Foot somewhat wider than dorsum in live specimens, but the opposite
is true after preservation. Anterior border of foot with blunt
extensions; posterior tip pointed (Fig. 1 A, E). Foot length 4-6
times width in actively crawling specimens (N = 11). Sole
anteriorly with shallow central longitudinal groove continuing
posteriorly as unpigmented line. Anus on small papilla anterior
right of pericardium covered by renal shield (Fig. 1 I). Renal pore
just left of anus papilla. Male aperture below right rhinophore,
vaginal aperture just posterior of male aperture and oviductal
aperture slightly posterior to and below vaginal aperture (Fig. 1
J).
Live colouration. – Colour of body and cerata is mainly determined
by colour of digestive gland: bright green in well-fed individuals,
and pale yellowish brown when not fed recently. Skin largely
transparent with scattered iridescent blue-green dots, white or
yellowish glandular dots, and angular reddish-brown dots, latter
especially on renal shield. These colours only visible under
specifi c light conditions. Moreover, fi ne grey-purple pigment on
lateral and dorsal sides, anterior foot border, and in two
longitudinal bands on sole of foot. White lines on posterior sides
of rhinophores and thin dark pigment on lateral sides. Appearance
of lines
and areas with grey-purple pigment on head largely depends on
degree of expansion of head (Fig. 1 C, I). A small black line
present on both sides of mouth (Fig. 1 K). No pigment around
genital orifi ces and eyes. In preserved specimens content of
digestive gland brownish and purple areas in epidermis grey.
Internal characteristics. – Diameter of eyeball 125–145 µm (N = 4,
length of live specimen 6–8 mm). Eyeballs touching (Fig. 2 A).
Clusters of oral glands besides anterior part of pharynx (Fig. 2
D). Pharynx nearly spherical, diameter 400– 520 µm (N = 5) with
prominent ascus, strong ascus muscle, and well developed dorsal
septate muscle. Wide, internal, longitudinal band of dark pigment
borders pharyngeal space (Fig. 2 C, D). Radular teeth blade-shaped,
smooth and narrow, with slightly curved sharp tips (Fig. 2 B).
Length of leading tooth in large specimens 85–100 µm (N = 5). In
ascending row 5-8 teeth, in descending row and ascus 18–27
progressively smaller teeth (N = 5). Teeth, including 3 preradular
teeth, remain fixed on radular ribbon that curls in a spiral in
ascus. Length of base of fi rst preradular tooth about 4 µm, second
about 8 µm, third with minuscule blade 15 µm in total. On both
sides of short oesophagus, salivary duct between pharynx and
salivary gland below prominent oesophageal pouch on dorsal side of
oesophagus just anterior of stomach (Fig. 2 D). Oesophagus and
pouch dark pigmented; salivary glands and ducts pale. Stomach dark,
ventral in body. From right dorsal side of stomach, pale coloured
intestine ascends to anal papilla (Fig. 2 D). A branch of the
digestive gland starts in both posterior corners of stomach. Both
immediately split into a branch that goes slanting upwards and a
branch that directly goes in posterior direction and then upwards
(N = 5). Both branches knotty, repeatedly branching in respectively
anterior and posterior dorsal parts of body. First branch reaches
to just posterior of eyes, second up into tail far posterior of
origins of posterior cerata (Fig. 2 E). Only some branches of
digestive gland in dorsum connected with a ceras.
Extensive network of albumen gland tubules ramifying in dorsal side
of body close to branches of digestive gland including in cerata
(Figs. 1 H, 2 G). Tubules white and relatively thick in egg laying
specimens, thin and pale in non-reproductive individuals, uniting
into single albumen duct. Hermaphroditic follicles dense (Fig. 2
F), small (diameter single follicles 50-150 µm, length of compound
follicles 200-300 µm), and numerous (>500), on ventral side of
body cavity from near frontal border of foot to below division
between anterior and posterior networks of digestive gland.
Follicles yellow, connected by ductules that unite into short
hermaphrodite duct. Organs of genital complex (Fig. 2 K) packed
together in anterior right half of body cavity. Hermaphrodite duct
widens into strongly coiled ampulla leaving it at anterior end and
splitting into vas deferens and small oviduct. Vas deferens
receives prostate duct coming from about 30 fusiform prostate
glands in anterior part of body below spherical genital receptacle,
then running anteriorly over female system to long, narrow, unarmed
penis (Fig. 2 H). Male system uncoloured, but prostate glands
white. Vaginal duct widens into bursa copulatrix going to posterior
end of
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THE RAFFLES BULLETIN OF ZOOLOGY 2007
solid, ovoid central canal. Small oviduct receives albumen duct and
enters central canal together with vaginal duct. Oviduct leaves
central canal in anterior end and continues as large oviduct via
wide and voluminous mucous gland occupying most of ventral right of
anterior body cavity. Genital receptacle spherical, pale, situated
dorsally slightly right of oesophageal pouch and left of loop of
bursa copulatrix (vaginal duct). Genital receptacle connected to
left side of central canal by narrow tube.
Habitat and behaviour. – Specimens of Costasiella coronata were
found on, and shallowly buried into, soft muddy sediment in and
around tufts of Vaucheria sp. in open spaces within or bordering
mangrove forests high in the intertidal zone. The slugs lived
amphibiously. When the Vaucheria patches had emerged during low
tide, more and more slugs crawled over the algae for feeding. The
contracted rhinophores were fi rmly kept against the back and the
cerata were held largely under the renal shield that seemed to
increase in thickness and relative size (Fig. 1 B). The epidermis
looked dirty by mud suggesting that the ciliary actions were
reduced or had ceased completely. The number of visible slugs
declined again by hiding in mud when the tide was out for some
time. In captivity it could be seen that they fed by sucking the
siphonal cells of Vaucheria sp. both when submerged and when
emerged. In October 2000 the temperature of the water was 35°C and
the salinity 29‰, but salinity fl uctuates strongly over the season
in the coastal zone (Swennen et al., 2001).
The slugs were diffi cult to keep alive in captivity for more than
a few weeks. The main problem was maintaining the algae in a
condition such that they could be consumed by the slugs; replacing
the algae every few days was necessary. A good growth of the
Vaucheria sp. did occur when keeping them with substrate either
permanently emerged in a damp atmosphere, or submerged with some
aeration, but in both cases the slugs either hardly fed or did not
feed at all.
Copulation was not observed in individuals out of the water. When
two submerged slugs met for mating, they erected their rhinophores
and repeatedly touched each other with the swollen anterior parts
of the head for about 15 seconds. Then they aligned their bodies in
opposite directions with the heads just side by side for penal
insertion. The duration of the copulation could not be timed
precisely, but the intimate contact lasted 15 to 30 s. When mating
ended the bodies shook as if the penis had to be redrawn with force
out of the partner. The eggs were in colourless, transparent,
gelatinous strings of a length of 9–15 mm and a width of up to 1.7
mm (Fig. 2 I). The strings contained 2000–4000 eggs in a spiral
chain (Fig. 2 J). The capsules of fresh eggs were touching each
other in the spiral, but not those in neighbouring curves. Fresh,
still uncleaved ova yellowish; developing embryos were pale. The
egg capsules measured 108.2 ± 3.6 µm by 90.2 ± 6 µm (N = 36),
diameters of uncleaved ova 61.1 ± 1.7 µm (N = 22). In captivity,
the egg masses were deposited both by submerged and emerged
individuals on the mud or on Vaucheria sp. The eggs developed to
free-swimming veliger larvae within four days (temperature
28–35°C).
Etymology. – The specifi c name refers to the striking character
that the dorsal side is crowned by the renal shield.
DISCUSSION
Characteristics of C. coronata, new species, agree with the type
species of Costasiella such as the position of the eyes, shapes of
radular teeth and penis, and several details of the genital complex
as described by Marcus (1982). C. coronata differs from all other
described Costasiella species by having fl attened cerata and a
large renal shield, both may be related to its specifi c
habitat.
Species considered to belong to Costasiella with their type
localities are: 1. C. ocellifera (Simroth, 1895); described as Doto
(?)
ocellifera. Transferred into Costasiella by Thompson (1977) after
suggestion of Marcus & Marcus (1960). St. George, Bermuda, UK,
North Atlantic Ocean.
2. C. virescens Pruvot-Fol, 1951. Type species by monotype. Monaco,
Mediterranean Sea.
3. C. formicaria (Baba, 1959); described as Stiliger (Stiliger)
formicarius, but transferred to Costasiella by Baba, (1961, in the
postscript). Amakusa, Japan.
4. C. nonatoi Marcus & Marcus, 1960. Near Ubatuba,
Brazil.
5. C. illa (Marcus, 1965); described as Stiliger (Ercolania) illus
and transferred to Costasiella by Marcus (1982). Ifaluk Atoll,
Caroline Islands, Micronesia.
6. C. lilianae (Marcus & Marcus, 1969); described as Stiliger
(Stiliger) lilianae belongs to Costasiella according to Thompson
(1977), and is a junior synonym of C. ocellifera according to Clark
(1984). Ubatuba, Brazil.
7. C. pallida Jensen, 1985. Hong Kong. 8. C. paweli Ichikawa, 1993.
Miyako, Ryukyu Islands,
Japan. 9. C. usagi Ichikawa, 1993. Ishagika, Ryukyu Islands,
Japan. 10. C. vegae Ichikawa, 1993. Ishagika, Ryukyu Islands,
Japan. 11. C. rubrolineata Ichikawa, 1993. Ishagika, Ryukyu
Islands,
Japan. 12. C. iridophora Ichikawa, 1993. Kuro, Ryukyu
Islands,
Japan. 13. C. kuroshimae Ichikawa, 1993. Kuro, Ryukyu
Islands,
Japan. 14. C. mandorahae Jensen 1997. Mandorah, Darwin
Harbour,
Northern Territory, Australia. 15. C. coronata, new species. Bang
Tawa, Pattani Province,
Thailand.
The majority of the species are found in tropical and subtropical
West Pacifi c waters, with only C. virescens, C. nonatoi and C.
ocellifera occurring in the tropical and subtropical Atlantic
region. Most species live in the subtidal photic zone and seem to
feed on Avrainvillea spp. as far as is known. Only C. coronata, C.
formicaria and C. pallida live in the intertidal zone. The food of
C. formicaria is unknown, whereas both C. coronata and C. pallida
feed
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Swennen: Costasiella coronata, new species
on Vaucheria sp. (Jensen, 1990; this paper) and occupy the same
niche in the tropics as do Alderia modesta (Lovén, 1844) and
Limapontia depressa (Alder & Hancock, 1862) in temperate and
boreal North Atlantic coasts (Engel et al., 1940; Hartog &
Swennen, 1952).
Several of the Costasiella species have been poorly described, but
C. formicaria, C. ocellifera (= C. lilianae), C. paweli, C. usagi,
and C. virescens are not similar to C. coronata because they have
tentacular anterior foot corners, while C. illa and C. nonatoi have
the cerata fi lled with knobby diverticula, and C. iridophora, C.
kuroshimae, C. rubrolineata, C. vegae and C. mandorahae have the
whole dorsum covered with swollen cerata. In 2000 when the fi rst
specimens of C. coronata were found, the author believed that they
belonged to C. pallida mainly because of their association with
Vaucheria sp. The specimens of both samples died in the laboratory
before they could be studied. When they were found again a few
years later, the author prepared a description of the external and
internal characters of the Thai specimens. Comparison of that
description with that of C. pallida showed that the Thai specimens
differed in a number of characters. The original description of C.
pallida is meagre and based on a single juvenile specimen, but
additions and an important correction have been published after
about ten adult specimens were found (Jensen, 1990). Jensen (1985,
1990) noted for C. pallida: (1) scattered bright red spots all over
the body; (2) a median ceras in front of the tail (also pictured);
(3) the tail shaped like a ceras (also pictured in ventral view);
(4) cerata moderately infl ated; (5) eight teeth in the ascending
limb, 14 in the descending and a heap of teeth in the ascus; (6) no
oesophageal pouch observed; (7) stomach lodged between branches of
mucus gland; (8) wide lateral branches of the digestive gland send
diverticula out into the cerata. Several cerata receive tributaries
from one lateral branch (Jensen, 1990: 429, fi g. 13); (9) white
branches of the albumen gland wind around digestive gland inside
cerata (also pictured); (10) penial opening behind the right
rhinophore and female aperture is behind penial opening; (11) dark
pigment is present on penial sheath; (12) sausage-shaped egg masses
contain 483.3 ± 283.9 eggs (N = 11); (13) egg capsules are 132.8 ±
7.9 µm by 107.4 ± 8.6 µm (N = 20); (14) fringe surrounding the
pericardium has an elevated edge that disappears in preserved
specimens; (15) no black stripes occur near oral opening (Jensen,
1985: fi g. 5). While in C. coronata there are: (1) no bright red
spots, but iridescent blue-green ones; (2) no median ceras occurs
in front of tail; (3) tail tapers gradually to a point; (4) cerata
strongly fl attened; (5) teeth in ascus are in a spiral; (6) large
oesophageal pouch is present; (7) stomach and mucous gland are
spatially separated; (8) several side branches of lateral branches
of digestive gland do not contact any ceras, and no side branch
send tributaries to several cerata; (9) branches of albumen gland
do not wind around digestive gland in cerata; (10) penial opening
is below right rhinophore, and there are two separate openings of
female system; (11) there is no dark pigment on the penial sheath
or penis; (12) number of eggs in an egg mass is higher (2000–4000);
(13) egg capsules are relatively smaller, measuring 108.2 by 90.2
µm; (14) fringe on pericardium does not disappear in preserved
specimens and this renal organ
extends more than halfway down the body and also to over more than
the width of the body; it is called a renal shield; (15) two black
lines alongside oral opening. These may be similar as the small
lines of black pigment cells that fl ank the mouths in Costasiella
nonatoi (Marcus & Marcus, 1960: 150), Elysia evelinae (Marcus,
1957: 411), Stiliger evelinae (Marcus, 1959: 23), which are reduced
to a melanophore on each side in C. lilianae (Marcus & Marcus,
1969: 10). The difference in the number and size of the eggs and
details in the coloration do not necessarily indicate specifi c
difference between C. pallida and C. coronata, but most of the
others characters do indicate specifi c differences.
The diagnosis of the genus Costasiella Pruvot-Fol, 1951, and of the
type species, C. virescens Pruvot-Fol, 1951, had a shaky start. The
sample became lost before the internal characters of the type
species could be studied. As striking characters for the genus were
noted the large not rolled, but somewhat fl attened rhinophores,
and anterior corners of the foot tentaculiform and pointed. For the
type species, it was stressed that the eyes are close to the median
line, nearly touching each other and that the anterior part of the
head is concave (Pruvot-Fol, 1951; 1954). The second species placed
in the genus was C. nonatoi Marcus & Marcus, 1960. This species
has rounded anterior foot corners and the rhinophores are grooved
along their whole length, thus the main genus characters were
invalidated in favour of the characters of the eyes of the type
species for including the species in the genus. Marcus (1982: 22)
stuck to the large size of the eyes to validate the genus. She did
not accept the transfer of Stiliger formicarius to Costasiella,
because the eyes are not especially large. This may be based on the
fi gure (Baba, 1959: plate 27, fi g. 3 a) where the eyes are
indicated by simple dots of similar size as in the drawings of
other slug species in the same paper. Baba (1959, 1961) gives no
measurements of the eyes. Jensen (1990) found the eyes of her C.
pallida were relatively smaller than those of C. liliana, but does
not give measurements. She maintained the placement in Costasiella
because of the position of the eyes between the rhinophores and
close together. Since then, all ceratal-bearing sacoglossans with
the eyes close together between the rhinophores have been
considered to belong to the genus Costasiella.
Marcus (1982) and Jensen (1990) stated that the named species of
Costasiella are rather diverse in several characters and the genus
may have to be split up. In the mean time, eight more species have
been named, but the situation remains unclear. The diversity in the
reproductive systems seems large in the few species in which
details are known. Two genital apertures have been noted in C.
virescens according to Marcus (1982), C. pallida according to
Jensen (1990), and C. mandorahae according to Jensen (1997). Three
apertures occur in C. nonatoi according to Marcus & Marcus
(1960) and Marcus (1982), C. lilianae according to Marcus &
Marcus (1969) and Marcus (1982), and C. coronata (this paper).
However, in C. virescens the vaginal duct opens in the male atrium
(Marcus, 1982), which makes it diffi cult to decide whether the
systems principally differs from C. coronata in which the vaginal
aperture is in the body wall
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THE RAFFLES BULLETIN OF ZOOLOGY 2007
just outside the male aperture (Fig. 1 J). An extra aperture has
been described that can be considered as an outlet for excess
seminal fl uids from the genital receptacle (Marcus & Marcus,
1969). It runs via a narrow duct to the dorsum in C. mandorahae
according to Jensen (1997) and to the left body wall in C. lilianae
according to Marcus & Marcus (1969). It may have been
overlooked in other species and does seem of diagnostic value as
far I know.
The vaginal aperture is anterior to the oviducal opening in C.
coronata and C. virescens. A fundamentally different character
seems to occur in C. lilianae, where the vaginal opening is
posterior of the opening of the oviduct (Marcus & Marcus, 1969:
11 and Abb. 25). Contrary to the other Costasiella species, C.
lilianae has a short penial stylet that was unfortunately not fi
gured [“kutikular Mundstück of 13 µm” in Marcus & Marcus
(1969); “a quite short stylet” in Marcus (1982)]. Both differences
suggest that Costasiella lilianae is related to Ercolania
(Limapontiidae). It may be that C. formicaria has the same
characteristics because Baba (1959) noted that the genital orifi ce
in Costasiella formicaria is situated roughly in the same position
as in Stiliger boodleae (Baba, 1938). Later S. boodleae was placed
in Ercolania after a careful description of its anatomy (Baba &
Hamatani, 1970a, b). Note that the presence of a short, pointed
penial stylet in C. pallida mentioned by Jensen (1985) was later
retracted as a mistake (Jensen, 1990).
Up to now, the genital complex has only been described and fi gured
by Marcus & Marcus (1969) for C. lilianae, Marcus (1982) for C.
virescens, and in this paper for C. coronata. The splitting off to
a new genus of C. liliana or placing it in Ercolania has to wait on
a check or redescription of Costasiella liliana because mistakes
cannot be excluded. C. liliana is a junior name for C. ocellifera
according to Clark (1984), but in the same paper is stated that the
ascus is absent in C. ocellifera without any comment, while Marcus
& Marcus (1969) has explicitly described the ascus in C.
lilianae. Clark (1984) also does not refer to the aberrant order of
the genital orifi ces in C. lilianae, which has the oviducal
aperture frontal of the vaginal aperture while the reverse is found
in other Costasiella species.
The reno-pericardial elevation may have diagnostic value. In C.
nonatoi the pericardium lies below the renal tube (Marcus &
Marcus, 1960: 151). In C. pallida the pericardium seems large,
starting at the level of the fi rst row of cerata and extending to
about halfway down the body. It has a fringe in the posterior part
and along the lateral sides, which disappears in preserved
specimens. The fringe contains numerous white branching tubules
that may be dorsal veins or renal ducts (Jensen, 1985; 1990). The
pericardium is small and covered by the renal area that has a
prominent fringe that remains present in preserved specimens of C.
coronata. Nothing about these characters is known for any of the
other species.
Clark (1984) erected the monogeneric family Costasiellidae because
the genus Costasiella was diffi cult to place in any of the known
sacoglossan families. The original family diagnosis included some
species-specifi c characters (e.g. fusiform
non-fl attened cerata, digestive diverticula usually knobby and in
grape-like clusters within the cerata, genital apertures polyaulic,
foot rather narrowly triangular, oesophageal diverticulum absent)
(Clark, 1984). This diagnosis has as remarkable result that the
type species of Costasiella and several other species including C.
coronata do not meet the family description. Therefore, it is
better to simplify the diagnosis of the Costasiellidae into:
ceratiform sacoglossa with the eyes close together; radular teeth
blade-shaped with smooth sides, gradually tapering to the tip;
penis long, thin and unarmed. Jensen (1996) found the family
Costasiellidae unwarranted in her cladistic analysis of the order
Sacoglossa. She placed the genus Costasiella within the
Limapontiidae Gray, 1847 (= Stiligeridae Iredale & O’Donoghue,
1923) and suggested it may deserve the status of a subfamily, thus
Costasiellinae. However, even when the status and phylogenetic
place of the Costasiellidae seems dubious and the various species
have to be placed in different known or still to describe genera
and taxa of higher order, it has an indisputable and practical
function for the moment. Any change has to wait till more species
have properly been described and older descriptions carefully
checked.
ACKNOWLEDGEMENTS
My sincere thanks are due to Mr Sathit Ponkoon, who was my reliable
and tireless assistant in the fi eld and the laboratory during this
study. Most of the fi nest dissections could not have been
conducted without his help. I like to thank Mr Nukul Ruttanadakul
for facilitating all aspects of the work in Thailand. Thanks are
also due to Messrs R. G. Moolenbeek and A. van der Bijl for
suggestions about the draft and their help in obtaining literature
and to two anonymous referees for detailed comments that helped me
to improve the fi nal text.
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<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>
/ENU (Use these settings to create Adobe PDF documents for quality
printing on desktop printers and proofers. Created PDF documents
can be opened with Acrobat and Adobe Reader 5.0 and later.)
>> /Namespace [ (Adobe) (Common) (1.0) ] /OtherNamespaces [
<< /AsReaderSpreads false /CropImagesToFrames true
/ErrorControl /WarnAndContinue /FlattenerIgnoreSpreadOverrides
false /IncludeGuidesGrids false /IncludeNonPrinting false
/IncludeSlug false /Namespace [ (Adobe) (InDesign) (4.0) ]
/OmitPlacedBitmaps false /OmitPlacedEPS false /OmitPlacedPDF false
/SimulateOverprint /Legacy >> << /AddBleedMarks false
/AddColorBars false /AddCropMarks false /AddPageInfo false
/AddRegMarks false /ConvertColors /NoConversion
/DestinationProfileName () /DestinationProfileSelector /NA
/Downsample16BitImages true /FlattenerPreset <<
/PresetSelector /MediumResolution >> /FormElements false
/GenerateStructure true /IncludeBookmarks false /IncludeHyperlinks
false /IncludeInteractive false /IncludeLayers false
/IncludeProfiles true /MultimediaHandling /UseObjectSettings
/Namespace [ (Adobe) (CreativeSuite) (2.0) ]
/PDFXOutputIntentProfileSelector /NA /PreserveEditing true
/UntaggedCMYKHandling /LeaveUntagged /UntaggedRGBHandling
/LeaveUntagged /UseDocumentBleed false >> ] >>
setdistillerparams << /HWResolution [2540 2540] /PageSize
[612.000 792.000] >> setpagedevice