BIOLOGY AND ANATOMY OF THE LIVING FOSSIL CONGERIA-KUSCERI (BIVALVIA, DREISSENIDAE) FROM SUBTERRANEAN RIVERS AND CAVES IN THE DINARIC KARST OF THE FORMER YUGOSLAVIA
B. Morton et al., BIOLOGY AND ANATOMY OF THE LIVING FOSSIL CONGERIA-KUSCERI (BIVALVIA, DREISSENIDAE) FROM SUBTERRANEAN RIVERS AND CAVES IN THE DINARIC KARST OF THE FORMER YUGOSLAVIA, Journal of zoology, 245, 1998, pp. 147-174
The cave-dwelling Congeria kusceri is described and aspects of its uni
que ancestry and biology elucidated in the context of its, for a bival
ve, similarly unique habitat. The species lives in subterranean waters
of the Dinaric karst of the former Yugoslavia. C. kusceri is acutely
heteromyarian in form commensurate upon its lifestyle of byssal attach
ment to the walls of either permanently or periodically inundated kars
t caves. Shells are sometimes covered by travertine, precipitated calc
ium carbonate, and the tubes of the, similarly unique, serpulid Marifu
gia cavatica. Neither species occurs outside such caves and, as a cons
equence, the shell and tissues of C. kusceri are colourless and there
are few sense organs, i.e. no statocysts and light receptors, although
the pallial papillae possess tiny sense cells and there is a supra-br
anchial osphradium. Mantle fusions are extensive and involve the inner
folds posteriorly to create the siphons, but also the inner surfaces
of the middle folds ventrally so that here the mantle is largely cover
ed by a thin (4-5 mu m), fibrous, periostracum and there are, on the s
eparated middle folds around the pedal gape, external ciliary cleansin
g tracts. There are internal cleansing tracts too, but the former have
been identified, hitherto, only in representatives of the Pectinidae.
In terms of general anatomy, Congeria kusceri has large ctenidia and
tiny labial palps as an adaptation to the mineral waters of its habita
t containing little food and, thus, that little is rejected when colle
cted. There are few sorting currents in the stomach either and the int
estine is short. The aragonitic shell comprises two layers, outer and
inner, both with a homogeneous crossed-lamellar structure. Each shell
valve, however, contains many growth checks, represented by thin prism
atic layers. Assuming that a growth check is formed once each year, po
ssibly in late summer when water levels fall in the caves, making them
accessible for investigation, an individual of, for example 12.7 mm s
hell length, was possibly 25 years of age. If so, an individual of 25
mm would be very much older. Because, with age, the species lays down
growth checks at progressively reduced increments, a lifespan of many
decades may be possible. Extreme reduction in shell growth causes marg
inal thickening and incurving. The species is dioecious, i.e. separate
sexes, with females brooding lecithotrophic (70 mu m) eggs in the cte
nidia, the filament bases of which become highly glandular. Mature and
brooding individuals occur when water levels are low in the caves, su
ggesting that internal fertilization takes place at this time too, spe
rm transfer being in the film of surrounding water to prevent loss. Fe
rtilized eggs probably hatch either as larvae with a short planktotrop
hic life or as crawl-away juveniles. Analysis of large samples suggest
s that recruitment is probably low, and there is evidence that the spe
cies is threatened by pollution of its habitat.