Bubble-cowries

A Review of the cruickshanki-group - Cypraeidae: Cypraeovula

The fascinating deep water species of Cypraeovula, castanea, cruickshanki, connelli and iutsui have been subject of vast interest by various authors, some of whom went in detail by showing variations and proposing a couple of forma-names. The taxonomy has been debated up and down (especially that of castanea) and the standpoints were made more or less clear. Apparently these four species have, despite their rarity, demanded special interest amongst taxonomists and collectors.

This seems reason enought to have a look at the present state of knowledge on the shells of these beautiful creatures, whose only advantage from our attention is, that once in their lifes - at the very moment they reach the surface - they will see the sunlight.

The person who has done the most detailed study, and who posessed the largest variety of these rare deep water shells was W.E.J. Walles of Durban - a scratchy old genius whom to meet I had the pleasure in 1982. Despite the fact that he was certainly a weird, somewhat confused and extremely difficult personality, his bright intelligence and sense of humor made the day at his place unforgettable. For instance, he showed me a manuscript he had prepared, which was rejected by all magazines he offered it to. This paper contained (amongst other interesting and a lot of "strange" things) first descriptons and photos of the two iutsui variations from the Atlantic (which Walles had known and posessed long before anyone else). And it contained the description and several photos of a shell that is now universally known as connelli, including its variation connelli peelae (this name was morally but not legally introduced by Walles - the merit for peelae should have been given to him - at least I adopted the name when finally introducing the name in 2002.

This site shows a variety of shells of the four species mentioned, it tells my personal story on the subject. I do not claim to show the complete range of variation, but a good deal of it. The specimens belong to different people and some pictures where taken from other sources but my collection. They are abbreviated: (Bu): Burgess, (Ch): Chiapponi, (Ga): Gabrielli, (Li): Liltved, (Ta): Tarr

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Cypraeovula cruickshanki, Paratype 2. This is one of the three original types after which cruickshanki was described. The typical form is plain white, rather large, with spiny prolongations of the anterior and posterior teeth. The labral base is always dull. Range: Durban area, 280-450 m Sizes: 28-34 mm

Cypraeovula cruickshanki, callous variety In callous specimens the left portion of the anterior extremity is produced and visible from dorsal view. The labrum is wider basally, and tends to have a faint orange hue. The spiny processes of the teeth are even more apparent. There is a regular labral flange, not markedly separated from dorsum. Range: Durban area, 350-550 m Sizes: 27-31 mm

Cypraeovula cruickshanki, lightweight variety In subadult or lightweight specimens, the spire is not fully covered by callus, the base is glossy on both sides and hardly calloused. The columellar side of the aperture forms a ridge. Such specimens very often have tiny holes from the spines of rock lobsters and sea urchins amongst which these shells were brought up by the trawlers. Range: Durban area, 250-550 m Sizes: 25-33 mm

Cypraeovula cruickshanki, bulbous variety During a short period of time, a large, bulbous form was found, in which the labral flange is hardly developed and the teeth are rather short, while the fossula is long and produced. The spire is deeply umbilicate. The few shells known were reported to come from further south of the typical spot offshore Durban. No living shells were found. Range: towards Amanzimoti, Natal, about 350 m Sizes: 29-34 mm

Cypraeovula cruickshanki, peach coloured form A beautiful variant shows a uniform orange-brown hue. Morphologically these shells resemble typical cruickshanki but are usually smaller. Range: Durban area, 350-450 m Sizes: 26-29 mm

Cypraeovula cruickshanki, flanged, orange-brown specimen This unique specimen shows a solid and prominent labral flange separated from the dorsum by a step. The labrum is wide, the teeth short. Dorsum and base show a uniform orange-brown colour. Range: Durban area, 450 m Sizes: 33 mm

Cypraeovula cruickshanki, irregularly flanged variety There are a few specimens with extreme development of a labral flange. The teeth are produced and spiny, the shell's outline is irregular. All specimens I have seen were dead taken, but from the typical habitat. Range: Durban area, 350-550 m Sizes: 27-30 mm

Cypraeovula cruickshanki, northern variety Some shells were trawled outside the typical area universally known as "The Gate" offshore Durban. These specimens are bulbous, with elegant extremities and a finer dorsal structure. If one believes the data of the trawlers, these shells were from shallower waters. Only deadcollected shells are known to me. Range: 50 kms north of Durban, 250 m Sizes: 25-29 mm

Cypraeovula cruickshanki, spotted variety A rather rare variety shows distinct, small brown or orange spots, mostly along the labrum (left specimen, Li), and sometimes also middorsally (right specimen). Morphologically, these shells seem to belong to the typical cruickshanki. Range: Durban area, 350-550 m Sizes: 27-32 mm

Cypraeovula cruickshanki, blotched variety Another variation shows larger yellow or orange blotches all over the dorsum. In the specimen on the right, the dorsum was covered completely with dense brown pattern. Unfortunately this shell was dead taken and is decorticated. Range: Durban area, 350-550 m Sizes: 25-33 mm

Cypraeovula sp. 1 Only three specimens of this population are known to me. The shells are depressed, very callous, but the labrum is not separated from the dorsum by a step, it is gently ascending towards the dorsal dome. The labral teeth are very long, the extend across the labral margin throughout. Such shells resemble Trivia splendidissima, but differ by having short columellar teeth. They evidently belong to the Cypraeidae. Range: Durban area, 600 m Sizes: 28-30 mm

Cypraeovula connelli peelaeParatype 1 This small specimen was trawled from over 600 m south of Durban. It is the paratype No. 1 of Walles' peelae. Range: Durban area, over 600 m Size: 23 mm

Cypraeovula connelli peelae The northern deep water population of connelli displays some characteristics of cruickshanki, especially in respect of shape. The specimen shown left is fresh deadcollected (Ch), on the right is a livecollected specimen (Li). Range: Durban area, 450-650 m Sizes: 19-27 mm

Cypraeovula connelli peelae, holotype The holotype of peelae is one of the most globular specimens I have seen. The dorsal pattern is partly preserved. Range: Durban area, 650 m Size: 24 mm

Cypraeovula connelli peelae, spotted variety This livecollected specimen (Li) shows distinct labral spotting occasionally found also in typical connelli, but not in cruickshanki. Range: Durban area, 550 m Size: 24 mm

Cypraeovula sp. 3 ="Crossia kasiae" Walles 1980, holotype (invalid name, the paper was rejected by the publishers of the Venus magazine) This rather worn and decorticate specimen of connelli was amongst the shells that Walles intended to name. Range: off Port Shepstone, Transkei, 250 m Size: 25 mm

Cypraeovula connelli , holotype The typical connelli is a slender shell without spiny dental processes. The extremities are callous and rather blunt. Comparing this shell with those connelli that have subsequently been dredged, one can say that the holotype is not really a typical shell. Range: central Natal, 200 m Size: 25 mm

Cypraeovula connelli peelae Dead specimens have been collected in the Aliwal Shoal area. The specimen shown here has short, blunt extremities and remnants of dorsal pattern, related to deep water peelae found somewhat further north. Range: Aliwal Shoal, 200 m Size: 22 mm

Cypraeovula connelli peelae, intermediate variety A few dead shells have also been dredged offshore Port Shepstone, all of them rather small and in very bad condition Range: Southern Natal, 100 m Sizes: 20-24 mm

Cypraeovula connelli , typical These fresh specimens (Li) show well the variability in shape and pattern. Both were collected in the same area offshore Coffee Bay. Note the degree of rostration in the left specimen. Range: Transkei, 100-150 m Size: 25-29 mm

Cypraeovula connelli , rostrated variety Along the Transkei coast, several differing populations seem to exist. The most spectacular one is a globular, heavy variation with rostrated extremities. It is also a favorite because it is occasionally found in a fresh dead or even livecollected state. Range: Coffee bay area, 80-100 m Size: 24-30 mm

Cypraeovula connelli , rostrated variety This is a livecollected specimen of the rostrated variety, dredged from 150m off Coffee Bay. Many shells show "attempted drill holes" which means a predator tried to drill a hole to inject acid and poison, but the cowry managed to exscape and repair the hole. Range: Coffee Bay, 90 m Size: 29,5 mm

Cypraeovula colligata This rare species is restricted to the northern Transkei. It is heavy, with fine teeth and a smooth base. The peristome is coarsely ribbed. It connects the Crossia-complex with other species of Cypraeovula. It is believed to be the result of hybridization of connelli with immelmani. Range: Msikaba to Port Edward, 90-100 m Size: 20-26 mm

Cypraeovula kesslerorum This rare species resembles colligata and connelli, but differs by a ribbed base. Few specimens have so far been collected. It connects the Crossia-group with Cypraeovula capensis gorda, which is believed to be involved in this hybrid form. Range: East London area, 90- 100 m Size: 20-24 mm

Cypraeovula volvens The small, nearly shpaerical shell of volvens makes it an unmistakable species related to iutsui. It is found in the Port Alfred area only. Range: Fish River, Port Alfred area, 90- 100 m Size: 18-22 mm

Cypraeovula iutsui , typical The holotype of iustui is a moderately large, inflated shell with an umbilicate spire. This speciens (Bu) resembles the holotype closely. The fine and dense dentition not forming spiny processes and the reduced fossula are safely telling iutsui from cruickshanki and connelli, whereas a conchological distinction from castanea is not always possible. Range: False Bay to southern Tanskei, 100-250 m Size: 28-45 mm

Cypraeovula iutsui , typical, but smaller form Most specimens of the typical form are moderate sized, with a distinctly curved aperture and some dorsal patches of brown. The shell is not very callous and usually the basal spotting of the former juvenile shines through. Range: False Bay to southern Tanskei, 100-250 m Size: 28-35 mm

Cypraeovula iutsui , globular variety The most spectacular specimens are those with an almost spherical shape. They usually belong to the typical iutsui. Range: Mossel Bay area, 150-250 m Size: 26-29 mm

Cypraeovula iutsui , brown toothed variety The teeth in iutsui are usually pale cream to white. In some specimens, there is a distinct brown staining of the teeth (Ta). Range: Mossel bay to Cape St. Francis, 150-250 m Size: 24-31 mm

Cypraeovula iutsui , giant variety Occasional giant specimens of more than 40 mm are known. The one depicted here was offered via the internet, but I have no idea where it ended up. It was claimed to measure 45 mm (Ta) Range: somewhere around Cape Agulhas Size: 45 mm

Cypraeovula iutsui , heavy form The lobster trapping in deep water offshore Cape st. Francis has produced many beautiful specimens of iutsui. These are heavy shelled, rather small and very colourful. The aperture is more narrow and less curved than in the larger, lighterweight iustui shown above. Range: Cape Agulhas to Algoa Bay, 80-250 m Size: 22-29 mm

Cypraeovula iutsui , heavy form Very often, the shells from lobster traps have fine cracks from dropping on the deck of the fishing boats. Completely perfect shells with vivid colours are still rare. Range: Cape Agulhas to Algoa Bay, 80-250 m Size: 22-29 mm

Cypraeovula iutsui , small globular variety Also from the Cape St. Francis area there is a small, globular variety, with heavily calloused base and rather rostrated extremities. Range: Cape St. Francis, 120-250 m Size: 18-25 mm

Cypraeovula atlantica, typical The population from moderately shallow waters from the Atlantic is characterized by a rather slender, little callous shell of whitish to pink colour. The dorsal pattern is usually absent. Range: Only known from an area south of Capetown, 40-55 m Size: 25-37 mm

Cypraeovula atlantica, typical Most specimens are heavily flawed. The growth lines are quite obvious in fresh (pink) shells, but almost disappear as the shell fades to cream. Range: Only known from an area called Soetwater, south of Capetown, 40-55 m Size: 25-37 mm

Cypraeovula atlantica, spotted variety Occasionally, densely spotted specimens are found amongst the plain coloured shells. Range: Only known from an area called Soetwater, south of Capetown, 40-55 m Size: 25-37 mm

Cypraeovula iutsui , lightweight variety Rather large, lightweight specimens of iutsui are found throughout the range of the species. They gradually may intergrade into deep water specimens of castanea, as the following pictures will demonstrate. Range: From False Bay to Algoa Bay, 150-300 m Size: 25-42 mm

Cypraeovula iutsui , lightweight variety The specimen (Ga) here already shows a tendency towards a more elongate castanea-shape and colouration. It is now considered a deep water form of castanea. Range: Mossel Bay Size: 38 mm

Cypraeovula levissima Paratype (Ch) Large, inflated and fragile specimens of iutsui have been trawled far offshore in the Atlantic, by trawlers operating from Hondeklip towards the Namibian border. These were described as ssp. levissima by Raybaudi. Range: From Hondeklip (Atlantic) to Algoa Bay, 250-500 m Size: 33-40 mm

Cypraeovula levissima The most extreme, nearly translucent specimens were collected at 500 m offshore Namibia. The illustrated shell shows the "juvenile" character combined with rather rostrate extremities. Range: Offshore the coast of Namibia, 500 m Size: 38 mm

Cypraeovula levissima This specimen was offered for sale on the internet. No locality data is given but the large, flat instead of umbilicate spire resembles a subadult deep water castanea rather than a iutsui. Range: ?? Size: 36 mm

Cypraeovula iutsui / castanea, intermediate This specimen, conchologically identical to Atlantic levissima but also resembles subadult castanea from the Cape St. Francis area. Range: Mossel Bay, 150 m Size: 38 mm

Cypraeovula castanea, fresh specimens Fresh specimens of castanea display a dense dorsal mottling of brown. Left is a subadult, on right one of the first ever livecollected shells (Li) Range: From Cape Agulhas to Ciskei coast, 25-55 m Size: 35-44 mm

Cypraeovula castanea, fresh specimens In some of the fresh shells there is a distinct brown staining of the teeth (there is a form like this in iutsui, too). In some shells, there also brown spots along the margins. Range: From Cape Agulhas to Ciskei coast, 25-55 m Size: 35-44 mm

Cypraeovula castanea, typical Most shells available to collectors are worn and white. This shell resembles the holotype of Higgins. It shows traces of pattern, is rather elongate, with white teeth and a wide aperture. Range: From Begha, Ciskei coast, beached Size: 39 mm

Cypraeovula castanea, typical This shell reflects best the concept of the typical castanea. The teeth are unstaned, the dorsum densely blotched with brown. Range: Cape st. Francis, off 45 m Size: 36 mm

Cypraeovula castanea, Cape Agulhas form - recently described as castanea malani A very poorply known variety is small sized, with short teeth. I have only seen very worn specimens, but perhaps diving in this area might produce fresh shells. Range: Cape Agulhas, only known beached Size: 26-29 mm

Cypraeovula castanea, densely toothed deep water variety A large, solid and heavy variety of castanea was trawled in 100-200 m. It has very dense, long labral teeth, spiny extremities and densely spotted margins. As few specimens are known so far, little is known about their status. Range: Port Alfred area, off 100-200 m. Size: 36-40 mm

Cypraeovula castanea, densely toothed deep water variety Recently, another such specimen was dredged at 40 m off Port Alfred. Also that shell has finer labral teeth. It may be a characteristic of the eastern castanea. Range: Port Alfred area, off 40 m. Size: 40 mm

Cypraeovula castanea latebrosa Solid and heavy specimens of castanea from deeper water off Jeffreysbay have been given the name latebrosa Swarts & Liltved. The margins of such shells may be spotted. Range: Jeffreysbay area, off 100 m. Size: 33-38 mm

Cypraeovula castanea latebrosa intermediate form This specimen was dredged from 80 m at Jeffreysbay. It shows the gradual transition from the extreme latebrosa to the regular shallower water castanea. Range: Jeffreysbay area off 60-80 m. Size: 33-37 mm

Cypraeovula castanea, globular deep water variety (Li) Few specimens of castanea from deep water off Cape Agulhas and Cape St. Francis display intermediate character to the smaller, heavy forms of iutsui. The margins show traces of spots, the teeth are long and dense on labral side, short and indistinct on columellar side. These shells constitute another bridge between the two species. Range: Cape Agulhas to Cape St. Francis, off 150-400 m. Size: 30-37 mm

Discussion:
Many authors are making a distinction between the cruickshanki-group and other members of Cypraeovula. This is a difficult venture, especially when it goes as far as including iutsui to the group often called subgenus Crossia. This name was introduced by Shikama for iutsui. The group iutsui, cruickshanki, connelli at first glance appears well defined by the fact that all these are deep water shells with globular dorsal domes. It is possible to distinguish connelli and cruickshanki from all others by the defined fossula and the spiny teeth. On the other hand, iustui and castanea are so closely linked by their deep water form that it seems impossible to separate them on a higher level. Castanea again resembles other shallow water species such as fuscorubra and coronata. In my opinion, it would make more sense to separate connelli and cruickshanki in one subgenus, and castanea and iutsui in another. Finally, the recently described colligata and kesslerorum (possibly products of hybridism) link connelli with typical Cypraeovula conchologically. Molecular data confirms the recognition of the Crossia group against the other species of Cypraeovula at some level, but certainly not as subgenus.

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First uploaded 2003, updated around 2005, 2012, and again July 2021