A preliminary revision of the living Notocypraea

by Felix Lorenz

The species of the genus of Notocypraea from the southern half of Australia are a confusing groups of cowries. Many authors have avoided dealing with that group and most collectors have simply "given up". Other authors have tried to manage the diversity of its members by introducing a large number of names. Many of these are based on single specimens and the descriptions are not accompanied by sufficient illustrations. Preliminary revisions made by other authors have either increased or reduced the account of taxa, more recent revisions have introduced the methods of DNA barcoding. Another approach included the features of radulae, however, to evaluate the importance of radula features in this genus would rewuire studies dealing with the general variability of radulae within populations, which in the case of cowries has only been done in very few genera (e.g. Zoila and Monetaria).

The present-day taxonomy and systematic arrangement of the genus is based on conchological features and in few cases, DNA features helped to clarify the status of species (in the case of pulicaria, dissecta and subcarnea). The DNA analysis, helpful in the study of genera and subfamilies, has added little that would support the species and subspecies evaluation in the other species. The opposite is the case, some species appear to be split in subspecies not supported by shell features (e.g. angustata), whereas apparently good subspecies (or species) are not different from their allies (e.g. occidentalis and piperita).

In the Pleistocene age, at the end of the last ice age, approx. 10.000 years ago, the ocean levels rose and vast areas of Australian coastline became oceanfloor. These new habitats presented a battlefield of different species competing for food sources, hiding places, ect. Random variability within each immigrant population and natural selection has lead to the formation and manifestation of new features, varieties and finally, species, subspecies, and a large number of variations in the shallow water. The situation in the Notocypraea is comparable to that of the genus Zoila. The archaic ancestral populations from which the shallow water immigrants originated may have looked pretty much the same as those forms we can find in greater depths today. In the deep water, the ecological conditions have hardly changed at the same time, the populations could remain more stable and retain more archaic features.

The species of Notocypraea mostly have wide ranges along the Australian coast, but their occurrences may be scattered. Four provinces seem to exist in which different species or subspecies are found: Western Australian Province (WA), South Australian Province (SA), Victoria and Tasmania (Vic), New South Wales (NSW). Little is known about the occurrence of Notocypraea from the Great Australian Bight. the name euclia suggests a population of Notocypraea in the Eucla area in the middle of the Bight, but I have not seen recent finds of such shells anywhere near Eucla.

The main problem in the conchological characterization of taxa is the enormous variability in every conchological feature, along with the rather frequent occurrence of conchological intermediates, which may be hybrids between nearly all taxa. After the comparison of five thousand specimens of Notocypraea I have found these features to be more important than others: general shape, structure of the fossula, character of the dorsal banding and pattern, size and density of marginal spots. The following species, subspecies and forms seem constant and consistent enough for a consideration:

angustata: no subspecies or forma-names. Synonym: castanea, verconis.

Oval depressed, flat fossula, dorsum mostly unbanded, white to dark brown color, no pattern. Marginal spots small and dense. The labrum is declivous anteriorly. Shells from SA are usually smaller, with larger and less dense marginal spots compared to those from Vic. Shells from Tasmania vary greatly in size and shape, some are black when freshly collected. There is a rare form with a faint dorsal band, perhaps a result of interbreeding with comptonii. Varieties from deeper water, mainly found in the Bass Strait area (Vic) have less marginal spots and sometimes a very pale, almost whitish dorsum. The names emblema, molleri and subcarnea were used for such shells in the past, however, all these names belong to what is now recognized as subcarnea, a valid species mainly found along the coast of Tasmania.

angusta.jpg angustadtas.jpg angutas3.jpg
 angustata, typical, Vic  angustata, typical, Vic  angustata, typical, Tasmania  angustata, banded form, Tasmania

angust2.jpg angust.jpg anguemb2.jpg anguemb.jpg
 angustata, western form, SA   angustata, western form, SA  angustata, var. with reduced spots, Vic  angustata, pale form, Bass Strait, Vic

declivis: no subspecies, forma-names or synonyms

Oval, depressed, flat fossula, dorsum unbanded or faintly banded with a compact transverse band, dorsum covered with minute spots, marginal spots sparse and small. Occasionally albinistic. Found in SA and Vic. Tasmanian specimens seem more solid and shorter than those from other areas.

deci1.jpg deci3.jpg decitas.jpg decitas2.jpg
 declivis, typical, Vic  declivis, albinistic, Vic  declivis, variation from Tasmania   declivis, variation from Tasmania

comptonii: no subspecies. Forma names: casta, mayi, trenberthae. Synonym: wilkinsi

Elongate oval to pyriform. Fossula mostly slightly projecting. Dorsum with a darker transverse zone with is darker along its borders, most shells appear to have two narrow transverse bands which are not interrupted towards the labral side. Dorsum often finely spotted, rarely netted. Marginal spots small and dense, mostly reaching onto base. The labrum is declivous anteriorly.

A very variable species found in WA, SA and Vic, but no subspecies can be recognized based on geography. Therefore, the names formerly listed as subspecies of comptonii are now regarded as mere individual forms (mayi, casta, trenberthae). Specimens from SA and Vic may have a dark brown base (variety trenberthae), there is also a wide range of purely white, pale orange to faintly spotted forms found within populations from SA and Victoria (except Tasmania) Many names exist to use on such shells and dealers have been very inventive in the past, yet the only correct name to use is casta. The callused and large variety mayi is found mainly in Vic, with a yellow to orange shell and sparse or absent dorsal spotting. Narrow specimens of that form from deeper water are also known as wilkinsi. Nearly all forms of comptonii seem to occur alongside in certain areas.

caomp1.jpg compex.jpg comt1.jpg comptre.jpg
 comptonii, typical, WA   comptonii, typical, WA   comptonii, typical, WA   comptonii, typical, WA

comlinc.jpg comtas.jpg compt4.jpg compt6.jpg
 comptonii, typical, SA   comptonii, typical, Tasmania   comptonii, typical, SA   comptonii, typical, Vic

trenb.jpg comter.jpg compdeep1.jpg compdeep2.jpg
  comptonii, var. trenberthae, SA    comptonii, var. trenberthae, SA  comptonii, deep water, Esperance WA   comptonii, deep water, Esperance WA

casta.jpg castaspot.jpg compt3.jpg mayihg3.jpg
 comptonii var. casta, Vic   comptonii var. casta, faint spots, Vic   comptonii intermediate casta / mayi, SA    comptonii intermediate casta / mayi, SA

mayi1.jpg mayi2.jpg comtres.jpg mayitas.jpg
 comptonii var. mayi, SA   comptonii var. mayi, Vic  comptonii var. mayi, Tasmania   comptonii var. mayi, Tasmania

piperita: one subspecies: occidentalis. Forma name: bicolor. Synonyms: reticulifera, leucochroa

Elongate oval to pyriform. Fossula mostly slightly projecting. Dorsum with a darker, usually interrupted transverse zone or two to four interrupted bands. Dorsum often finely netted, often along with larger spots. Marginal spots small, mostly not reaching onto base.

A very variable species found in WA, SA and Vic. Albinistic specimens do not seem to occur in piperita from SA and Vic. Specimens with conspicuous dorsal banding are called var. bicolor, another name formerly in use was dissecta, which is actually the cowry formerly called hartsmithi (see below).

pipesp3.jpg pipesp.jpg pipbanding.jpg piper2.jpg
 piperita, typical, Esperance, WA  piperita, typical, Esperance, WA   piperita, typical, Albany, WA    piperita, typical, Albany, WA

piers3.jpg pipbicol.jpg pip5.jpg bicolo.jpg
 piperita typical, SA  piperita var. bicolor, SA   piperita var. bicolor, SA   piperita var. bicolor, Vic

occidwest.jpg pipesp2.jpg spec1.jpg species2.jpg
  piperita, deep water, Esperance, WA    piperita, deep water, Esperance, WA    piperita variation, Cape Recherche, WA     piperita variation, Cape Recherche, WA

piperita occidentalis: No forma names or synonyms.

More pyriform. Aperture rather narrow, fossula more developed. Pale cream to white, mostly with narrow interrupted bands, sometimes very sparse pattern. From the western coast of WA, Cape Leeuwin area. Sometimes albinistic, then easily confused with pulicaria var. eulica.

Formerly considered a valid species, but similar shells are found among typical piperita from other areas so that this taxon should be considered a western subspecies, at most.

occid1.jpg occid2.jpg occideuc.jpg
 piperita occidentalils, typical, WA   piperita occidentalils, typical, WA   piperita occidentalils, albinistic, WA    piperita occidentalils, albinistic, WA

pulicaria: no subspecies, one forma: euclia. Synonym: candida

Narrow cylindrical, slightly depressed. Fossula distinctly projecting. Dorsum with four narrow interrupted bands, mostly with sparse spots. Marginal spots small and dense. Rarely albinistic (var. euclia). Restricted to WA. A deep water (80-150 m) variation from the Esperance area is more inflated, and a shorter fossula. The name euclia is also misused for dwarf albinistic piperita from the Cape Recherche area. Interestingly, the holotype of pulicaria is distinctly spotted dorsally, but lacks the four transverse bands.

pulica.jpg pulica2.jpg pulicsf.jpg pulics.jpg
 pulicaria, typical, WA   pulicaria, typical, WA   pulicaria, typical, WA   pulicaria, pale, nearly unbanded, WA

pulicque.jpg pulique2.jpg pulics2.jpg spepip.jpg
 pulicaria, deep water, Esperance, WA   pulicaria, deep water, Esperance, WA   pulicaria, deep water, Esperance, WA   pulicaria, deep water, Esperance, WA

pulieucl.jpg puliceu.jpg puliceu4.jpg puliceu2.jpg
  pulicaria var. euclia, Cape Recherche WA    pulicaria var. euclia, Cape Recherche WA      pulicaria var. euclia, Esperance, WA  pip. occidentalis resembling euclia, WA

dissecta: no subspecies or forms, synonym: hartsmithi

Pyriform, mostly rostrate. Fossula slightly to strongly projecting. Dorsum with four indistinct narrow, interrupted bands and no other pattern. Marginal spots large and dense. The prominent fossula along with the large marginal spots make this species unmistakable.

In an important paper published in the Records of the Australian Museum in 1931, Tom Iredale described several subspecies and forms of Eastern Australian cowries. Among them is Notocypraea dissecta from 90 m depth off Twofold Bay, just south of Eden, New South Wales. The illustrations of the holotype are poor, but one feature of the 21 mm shell is visible: large lateral spots. Its geography and this feature had given me the suspicion that dissecta might be an earlier name for the elusive Notocypraea hartsmithi Schilder from New South Wales. At last, I managed to trace the dissecta type with the kind help of Dr. Alison Miller and Dr. M. G. Allen of the Australian Museum, who submitted photographs. The large, sparse marginal spotting, slender shape, umbilicate spire, rostrate extremities and the projecting fossula, the fine dentition and the faint middorsal banding agree and it is apparent that hartsmithi and dissecta are conspecific. Therefore Notocypraea dissecta Iredale 1931 must be given priority over Notocypraea hartsmithi Schilder 1967. That name was described on the basis of a single beachworn specimen collected at Wyargine, Sydney. (Original Reference: Notadusta hartsmithi F. A. Schilder (1967) Arch. Moll., Vol. 96, p. 3941.). F.A. Schilder gave a detailed and accurate analysis of the specimen, along with details of the characteristic fossula. A second specimen was later found on the same spot, largely confirming Schilder's first diagnosis. This specimen is not available to me for study. After these two findings, it became quiet around hartsmithi and dissecta. Both names were virtually forgotten, and listed as synonyms of comptonii, and the name dissecta was also in use for piperita with distinct banding and reduced dorsal pattern.

When I met Lance Moore in his shop at "the Rocks" in the downtown of Sydney in 1982, he gave me a batch of beachworn Notocypraeas of all blends, with the remark: see what you can find out on these. Back in Germany I realized that one specimen was quite different from all the others. It contained a label stating "Hacking River", which is an area near the Sydney harbour, and i had a deja-vu. Some research in my piles of xeroxed cowry-literature finally produced what I had been looking for - the description of the "Sydney-Cowry" hartsmithi. Comparing the poor photo with my shell I had no doubt that I was looking at a hartsmithi. For the next years, I kept searching through all collections and selections of Notocypraeas I could locate, with a couple of "almost- hartsmithi-like" shells showing up, but none was actually the "real thing".

When the work on the "Guide to Worldwide Cowries" began, my hartsmithi remained the only specimen we had at hand. It took a long time to convince my late friend Alex Hubert that even on the basis of this one shell it was necessary to list hartsmithi as valid - in the end he surrendered with more than just reservations... Of course, the rarity of hartsmithi made a lot of people point at me with criticism, I agree it must have appeared a bit forward to draw conclusions from just one shell and a poor xeroxed b/w photo.

During the decades of a split Germany, Schilder's collection with the holotype of hartsmithi was hidden away in chests and boxes in the Museum of East Berlin, inaccessible to us. After the fall of the Berlin wall the holotype finally could be located in what was left of the Schilder-collection. It was photographed for me by Dirk Fehse of Berlin in 1997. Little later, a livecollected hartsmithi appeared in an old Australian collection: much smaller, paler, and from further south (Bass Strait area), but conchologically typical. Now I am able to show another, very typical specimen corresponding exactly with the holotype and my first specimen. It was collected on a beach south of Sydney and is in moderately good condition.

Since then, at least a dozen more specimens of N. dissecta (hartsmithi) were collected, from fishtraps off New South Wales, and by a divers at 15-25 m, in different areas in New South Wales.

dissec1.jpg dissecta.jpg diss3.jpg dissect4.jpg
 dissecta, typical, NSW   dissecta, deep water, NSW   dissecta, typical, NSW coll. Hawke   dissecta, typical, NSW

subcarnea: no subspecies or forms, Synonyms: albata, emblema, molleri

Oval, depressed to inflated, usually large. Flat fossula, ribbed posterior columella. Dorsum whitish to cream, sometimes with very faint narrow interrupted bands. The labrum is gently curved anteriorly, and not declivous as in comptonii and angustata. Recently, studies of the holotype of subcarnea revealed that this name has been misinterpreted as a variety of angustata, which differs by having smaller and denser marginal spots and a posterior columellar ridge instead of ribbing. A comprehensive study was published recently in the Visaya-magazine. It is a valid species supported also by DNA differences. It seems restricted to Tasmania and the Bass Strait.

In a review of the cowries from Tasmania, Beddome (1897) compares numerous variations of Notocypraea collected in Tasmanian waters. He differentiates between comptonii, piperita, declivis and angustata. He assigns four main variations to latter species: angustata typical, var. subcarnea (which he refers to as a taxon suggested by Ancey), var. albata, and var. mayi (which today is assigned to comptonii). In his hand-drawn plates he illustrates these in an easily recognizable way. Hugh Morrison and Simone Pfützner supplied a set of livecollected Notocypraea from the South of Hobart, Tasmania, as well as a large collection of shells gathered on 7 Mile Beach, Port Stanley, Northwestern Tasmania over a period of many years. Among that material, I noticed a strange cowry that recalled me of a shell I had studied in the British Museum in summer 1985: the holotype of subcarnea. That shell (reg. No. 1900.11.14.26) confirms my Tasmanian series as being the true subcarnea. From a single livecollected shell taken by Hugh Morrison at 6 m in the southeast of Tasmania, along with declivis and angustata, a piece of dried animal was recovered and sent to Dr. Christopher Meyer for DNA analysis. He managed to retrieve enough material to sample the specimen and informed me that it did not correspond at all with any of the classic Notocypraea-species but seems closest related to Notocypraea dissecta (formerly hartsmithi, see above) from New South Wales, which had just been included in the database (Meyer 2004). Finally, I found another, apparently live taken, specimen trawled at unknown depth in the north of Tasmania - in my own collection, wedged inbetween boxes in a drawer of "odds and ends".

Notocypraea subcarnea Beddome has a large (21-32 mm), depressed shell with a moderately strong callus along both sides, the spire is covered with callus and deeply umbilicate in juvenile shells. Some specimens are fairly elongate and drop-shaped. A feature of some shells is a transverse ribbing of the shell-surface, already noted by Beddome. The base is convex, the short extremities are callused dorsally. The labrum is convex, its outer margin is angular and bent up towards the dorsum. The dentition is very fine, extending slightly onto the labrum and the base. The fossula is fairly well developed, showing three to six denticles. The aperture is wide and strongly curved throughout. The posterior end of the columellar side is distinctly ribbed. The anterior terminal ridge is very short and declivous. The dorsum is pale yellow-cream, without secondary pattern but mostly with four rather indistinct and narrow interrupted brown bands. The marginal spots are large, sparse, rather pale and mostly situated towards the dorsal part of the margins. The basal aspect rarely shows any spots. The extremities are conspicuously tinted darker. The animal seems to be bright red judging from the remnants found in the dived specimen.

Notocypraea subcarnea differs from angustata by the posterior end of the columellar side being ribbed instead of smooth. N. subcarnea is more depressed and paler, the banding differs by being narrow and interrupted whereas in the rare specimens of angustata that show a banding that is broad and uninterrupted. The anterior terminal ridge of angustata is longer and rather straight, the aperture is less curved. The extremities of angustata are shorter and less callused. The most easy to determine feature separating subcarnea from angustata and all but one species of the genus is the sparser and much larger spotting of subcarnea. That feature is shared only with Notocypraea dissecta which also has a similar dorsal banding, but which is a narrow, elegantly rostrate species with a stronger fossula. It is interesting to note that the formation of the marginal spotting of most Notocypraea supports the genetic relationships. Those species which differ from having coarser and sparse spotting (dissecta and subcarnea) are also closest related and more remote from the rest which all have very fine and dense marginal spots (declivis, comptonii, piperita, angustata and pulicaria). These conchologcial features are consistent within the series studied. A comprehensive revision of the genus, including comprehensive statistical analysis, is in preparation.

The relative rarity of subcarnea has probably supported the fact that it was overlooked for so long. It seems endemic to the shores of Tasmania. Its name has instead been misused for all kinds of pale variations found in Notocypraea, e.g. the white shelled form of comptonii (variation or subspecies casta), or pale specimens of angustata from deeper waters. Incidentally, Beddome's albata (introduced after subcarnea in the same 1897-paper) is also based on a specimen of subcarnea and therefore becomes a synonym of that species.

subca2.jpg subc1.jpg subc3.jpg subc4.jpg
 subcarnea, typical form, Tasmania   subcarnea, typical form, Bass Strait, Vic   subcarnea, large form, Tasmania    subcarnea, large form, Tasmania

In the Conchologists of America Vol.35 No. 2, Don Cram reports the re-discovery of Notocypraea emblema. He illustrates the holotype, which clearly shows all the conchological features of subcarnea, including the characteristic large marginal spots and the ribbing of the posterior columellar canal. The shell is slightly distorted, hence the labrum does not look very characteristic but is untypically curvy. In his most interesting article he illustrates radulae of different species of Notocypraea, including that of the holotype of emblema (hence the first illustration of the subcarnea-radula). He discusses a cowry he calls Notocypraea sp. x cf. emblema on page 16 fig. 2 and 3. These illustrations in my opinion show N. comptonii var. casta with faint labral spots. The fossula region, the small size and density of the marginal spots, the declivous labrum and the smooth formation of a posterior columellar ridge bordering the canal reveal that these shells are not the same species as the shell shown in fig. 1 on that page, whose ribbed posterior columellar canal, the rather evenly curved instead of declivous labrum and larger sparse marginal spots identify it as subcarnea. The radula for comptonii is not shown to compare with the shells labelled N. sp. x cf. emblema, but in Bradner & Kay's Catalogue of Cowry Radulae fig. 210 there are good illustrations which reveal that Cram's shells have radulae most similar to those of comptonii.

Aug. 2007