Table Of ContentHYM. RES.
J.
Vol. 17(1), 2008, pp. 86-109
Patterns of Usage of Snail Shells for Nesting by Wasps
(Vespidae: Masarinae and Eumeninae) and Bees
(Megachilidae: Megachilinae) in Southern Africa
Sarah K. Gess and Friedrich W. Gess
Department of Entomology, Makana Biodiversity Centre, Albany Museum, Grahamstown, 6139
South Africa; email: [email protected] and [email protected]
—
Abstract. The present contribution to the knowledge of the use of snail shells for nesting by
Hymenoptera in southern Africa adds considerably to the earlier contribution by the authors on
thissubject. Itestablishes thatthestrategyofnestinginsand-filledshellsiscommonlyemployedby
Qiiartinia Ed. Andre (Vespidae: Masarinae) throughout the sandy winter rainfall areas. It records
additional rearings from Quartinia nests of Allocoelia Mocsary (Chrysididae), Tricholabiodes
Radoszkowski and an undescribed genus (Mutillidae), and Apolysis hesseana Evenhuis and
Greathead (Bombyliidae). Nestingby Tachi/sphex hermia Arnold (Crabronidae: Larrinae) in a sand-
filled snailshell is recorded forthefirsttime. Itestablishesthatthedistributionofuseofsnailshells
for nesting by bees coincides with and extends along the south coast east of that of Quartinia. The
bees (Megachilidae: Megachilinae) involved are osmiines, species of Wainia (Caposmia) Peters and
Hoplitis (Antliocopa) Lepeletier and Serville, and anthidiines, including Afranthidium (Afranthidium)
hamaticauda Pasteels, all cavitynesters usingemptysnailshells. The onlyotheraculeaterecorded as
usingempty shells isAlastorricaeGiordaniSoika (Vespidae: Eumeninae). Parasitisminbeenestsin
snail shells is uncommon. Only Chrysis Linnaeus (Chrysididae) and Eupelminae (Chalcidoidea:
Eupelmidae) were found, each at only two sites. The preferred snail shells are those of several
species of Trigonephrus Pilsbry (Dorcasiidae) on the west coast and in the desertic areas and of
Tropidophora ligata (Miiller) (Pomatiidae) alongthe southcoasttotheeast, allindigenous species. In
some areas the habitats of these have been invaded by an exotic snail, Theba pisana (Miiller)
(Helicidae). Shells ofthis snailareusedtoalimited extentbythesmallernestersinsnailshells. This
exotic snail presents a threat to the indigenous snails and to the snail shell nesters. In some areas
ongoing coastal development is also a threat.
The bulk of the know^ledge of the use of dium, from the Mediterranean into Asia
snail shells for nesting by aculeate Hyme- Minor (Extracted from OToole and Raw^
noptera prior to Gess and Gess (1999) vv^as 1991 and Bellman 1995). Minimal records
confined tothenorthernHemispherew^here for southern Africa prior to 1999 were
the use ofemptysnail shellsbybees thatdo observations, not supported by specimens,
not excavate nesting burrows, but search of a ''Eumenid-wasp" and an "Anthidium-
out pre-existing burrows in which to con- bee" nesting in Trigonephrus shells (Hesse
struct their brood-cells, is widely known 1944) and a record, supported by speci-
and is practised in a variety of habitats by: mens, of nesting by Wainia (Caposmia)
in the Nearctic two species of Osmiini elizabethae (as Osmia sp. in Gess and Gess
{Osmia and Ashmeadiella); and in the Pa- 1988 and as Hoplitis sp. in Gess and Gess
laearctic 14 species of Osmiini {Osmia and 1997) in shells of Tropidophora ligata collect-
Hoplitis), three in Britain, five in Central ed by Ed. Callan on coastal sand-dunes at
Europe, three in the Mediterranean, four in Salt\4ei, Port Alfred, Eastern Cape.
Eurasia and one in Japan, and for one The 1999 study investigated the use by
speciesofAnthidiini,a speciesofK/zot^^rz^/2z- wasps, bees and spiders of empty and
Volume 17, Number 1, 2008 87
sand-filled shells of Trigonephrus at 10 sites investigated the use of the shells of other
in seven desertic winter-rainfall areas in land snails-an invasive exotic snail Theba
southern Africa from immediately north of pisana along the coast from Port Nolloth
the Orange River east of Oranjemund southwards to Cape Town and eastwards
south to Wallekraal inland of Hondeklip along the south coast to Riet River Mouth
Bay (Gess and Gess 1999). The principal in the Eastern Cape Province (15 sites) and
users ofthe shells were found tobe spiders an indigenous snail Tropidophora ligata, the
constructingsilk-bagnests and twonesting shells of which were encountered only on
categories of aculeate Hymenoptera: bur- the southeastern coast. Neither large num-
row excavators using sand-filled shells, bers of snail shells nor use of snail shells
two species of Quartinia; and nesters in for nesting was discovered in any other
pre-existing cavities using empty snail areas during the course of fieldwork
shells into which are brought in nesting conducted throughout the semi-arid to
materials, a eumenine wasp, Alastor ricae, arid areas of southern Africa-of particular
and megachilid bees - single species each interest, nowhere in the sandy areas of the
of Wainia (Caposmia) (as Wainia (Wainia) Namib Desert north of Liideritz, or from
and Hoplitis (Anthocopa) (Osmiini), and the Kuiseb River north to Terrace Bay nor
Afranthidium (Afranthidium) hamaticauda in the southern Kalahari, that is desertic
(as Afranthidium (Oranthidium) and in Gess areas outside the winter rainfall area. The
and Gess 2007 as Afranthidium (Afranthi- results of this more extensive study are
dium) ablusum) (Anthidiini). Recorded as- presented in the present paper together
sociates were undescribed Tricholabiodes with the earlier contributions giving an
(Mutillidae: Sphaeropthalminae: Dasylab- account of the present knowledge of
rini), undescribed Allocoelia (Chrysididae: nesting by aculeate wasps and bees in
Chrysidinae: Allocoeliini) and Apolysis snail shells in southern Africa.
(Bombyliidae: Ursiinae: Ursiini) from Names of authors of taxa are used in the
Quartinia nests, Eupelminae (Chalcidoidea: body of the paper only if they are not
Eupelmidae) reared from cocoons of H. available from the abstract and appendix.
(Anthocopa) and Trichodes aulicus Kl. (Cole-
METHODS
optera: Cleridae) from a shell used for
—
nesting by W. (Caposmia). Sampling sites. The shells of medium to
A sample of 13 Trigonephrus shells from large terrestrial snails, are abundantly
the GrootDerm in the ''Yellow Dunes, part available, empty and sand-filled, in the
of the northern Succulent Karoo, Northern desertic winter rainfall areas and the areas
Cape'', collected by members of the BIO- of sandy coastal dunes of southwestern
TA-Southern Africa Project, directed from Africa, where they offer abundant secure
Hamburg, Germany. From these unopened microhabitats in these areas of sparse low
shells deposited in the Natural History vegetation and unstable, often windswept
Museum, Berlin, Hoplitis sp., Alastor ricae sand (Figs. 2-5). Generally Asteraceae,
and a chrysidid wasp, Chrysis grootdermen- Aizoaceae: Mesembryanthema (forraerly
sis Koch emerged (Koch 2006). Mesembryanthemaceae) and monocotyle-
Since 1999 the authors have extended donous geophytes were represented and in
their sampling of Trigonephrus shells north- the northern desertic areas generally in
wards in the winter-rainfall desertic areas addition Geraniaceae (Sarcocaulon), Neur-
of southwestern Namibia to the northern adaceae (Grielum) and Zygophyllaceae
limit of the distribution of the snails (23 (Zygophyllum). Papilionoideae (Fabaceae)
additional sites) and southwards, princi- were present at some sites. All snail shell
pally along the coast to Cape Town (14 sampling sites, except the more eastern
additional sites). Where present they have ones on the coast lie within the winter
Journalof Hymenoptera Research
Fig. 1. DistributionofsnailshellsamplingsitesinsouthernAfrica.SolidspotsindicatesitesatwhichQiiartinia
(Vespidae: Masarinae) was found nesting in snail shells.
rainfall area. Sampling sites were selected Identity, size and distributions of snail
—
throughout this area (Fig. 1, map, and shells sampled. Snail shells sampled were
Appendix, list of all sites with their co- all of terrestrial snails (Mollusca: Gaster-
ordinates). opoda) (Figs. 6-11) - several species of
I, The Klingi artberge, a sparsely x'egetated desertic area in southwestern Namibia.
.
Volume \7, Number 1, 2008 89
m
—
~ .^' --'^^ ' > ,,
f
A''
M
%J<x.
^* "^^^r^^'^yr^M^^^^m
Fig. 3. Near Lutzville southwest ofVanrhynsdorp in the Western Cape, sandveld.
Fig. 4. Donkinsbaai, southwestcoast.WesternCape,wellvegetated dune slackbehind foredunes.
90 Journalof Hymenoptera Research
Fig. 5. VVit:?and, v\estern south coast, sparselyvegetated dunes.
Trigonephrus (Dorcasiidae), Tropidophora li- were also sampled. From Port Nolloth
gata (Pomatidae), and an invasive exotic south to Cape Town and east to Riet River
species, Theba pisana (Helicidae). in the Eastern Cape Theba pisana was
Trigonephrus shells are by far the largest abundant. Trigonephrus was not encoun-
with an altitude of 27-44 mm and a major tered east of Cape Town although one
mm
diameter of 24-40 (Gess and Gess species, Trigonephrus ambiguosus (Per.), has
1999). Tropidophora ligata shells are mark- been recorded as far east as Mossel Bay
edly smaller and therefore offer a cavity of (Connolly 1938, p. 238). Theba pisana was
considerably smaller volume but are sim- generally abundant to the eastern limit of
ilarly of greater altitude, 19.7 mm, than sampling, at Riet River Mouth in the
mm
major diameter, 16.8 (Connolly 1938). eastern Cape. Tropidophora ligata although
Theha pisana are also of considerably recorded from as far west as the Cape
smaller volume than Trigonephrus and in Peninsula (Connolly 1938, p. 543) was
addition differ from both Trigonephrus and found no further west than Port Elizabeth
Tropidophera in having a low spiral, the in the Eastern Cape, from where it was
altitude, 12.2 mm, being less than the sampled at sites eastwards to Riet River.
mm
major diameter, 17.0 (Connolly 1938). Theba pisana is principally coastal Medi-
In the winter rainfall area in southwest- terranean in origin but also occurs in
ern Namibia and immediately south of the Switzerland, Ireland and southwestern
Orange River all snail shells encountered Wales and England. It was apparently
were of Trigonephrus and so samples from introduced into Cape Town from Europe
sites in this area are limited to shells of circa 1881 (Connolly 1938, p. 269) from
snails of this genus. At sites in the vicinity where it has spread north along the west
of Port Nolloth shells of Theba pisana were coast, east (at least as far as East London,
of equal or even greater abundance and Mary Bursey pers. com.) along the south
Volume 17, Number 1, 2008 91
Figs 6-11. 6-7, Shells (MoUusca, Gasteropoda) ofTropidophora ligata (Pomatiidae), Trigonephrus (Dorcasiidae)
and Thebapisana (Helicidae) X 1 (scalebar = 2 cm). 8,ShellsoftwospeciesofTrigonephrustoshowvariationin
shell size. 9, Sand-filled shell of Trigonephrus showing entrance turret of Quartinia (Vespidae: Masarinae). 10,
Sand-filled shells ofTropidophora ligata. 11, Sand-filled shells of Theba pisana.
coast, and in the west has invaded vine- slime production is considered to make
yards (Ferreira and Venter 1996) and citrus infested plants unpalatable to stock and
orchards (Joubert and Du Toit 1998). It has pollinators. —
been the subject of considerable study in Sample size. A sample size of 100 shells
Israel, southern and western Australia and was chosen, however, sample size taken at
California, in which areas it is a serious some sites was smaller or larger than this
agriculturalpest (e.g. Arad and Avivi 1998, number. For example smaller samples of
Baker 1988, Deisler and Stange 2005). Trigonephrus shells resulted from scarcity
Apart from damaging crops its copious of shells at sites on the west coast in the
92 Journalof Hymenoptera Research
vicinity of Cape Town or from the short- purpose in daylight hours even in cold,
ness ofcollection time for some stops in the wet weather. In the calculation of percent-
Sperrgebiet (Diamond Area no. 1), south- age of shells used empty and sandfilled
western Namibia, in which the authors shells are combined and a shell used by
were required to be part of a multidisci- more than one occupant is counted only
plii^ary group with varied agendas. Larger once.
samples resulted either from the combina- In the desertic area from the Drachen-
tion of several 100 shell samples from a berg in the north to Alexander Bay/
single site or, in the case of Thebnpisana and Brandkaros in the south 29 sites were
Tropidophora ligata, at sites, where inci- sampled. All shells in this area were of
dence of use was very low, particularly Trigonephrus species. The percentage of
along the south coast and eastwards, the used snail shells ranged from 33% (disre-
decision to take larger samples. garding one exceptionally low percentage,
—
Sampling method. The area of sampling 8%, obtained from near Kaizer's Camp to a
sites was not measured but was approxi- remarkably high 92% (average 50%).
m
mately 50 by 50 m. Shells were collected In the coastal sandveld from Port Nol-
by walking too and fro across the square loth in the north to Wallekraal to the south
and progessing from one side to the other. 11 sites were sampled. At all sites samples
The shell samples were collected into of Trigonephrus shells were taken. The
sealed plastic bags. Later live insects which percentage of shells used ranged from
had escaped into the bags were captured 40% to 100% (average 69%). In the Port
and the shells were broken open and the Nolloth and Hondeklip Bay areas shells of
contents examined. Live immatures were r. pisana were present and at some sites
reared. Voucher specimens were deposited were verymuchmore abundant than those
in the Albany Museum,—Grahamstown. of Trigonephrus. In a sampling site five
Condition of material. As snail shells in kilometres east of Port Nolloth a sample of
arid areas become bleached but do not 125 shells of T. pisana was taken with ease
degrade it is not possible to establish how but only five shells of Trigonephrus were
long theyhavebeen available. Consequent- found in the same area. Only 25% of the
ly, apart from active nests - nests contain- shells of T. pisana hadbeenusedbutall five
ing live occupants - it is not possible to Trigonephrus shells had been used. Similar-
establish in what year the nests were ly in a sampling site at Hondeklip Bay a
constructed. The condition of nest material sample of 121 Theha pisana shells was taken
and dead occupants, however, varies con- with ease but only 11 shells of Trigonephrus
siderably, from some nests being in a very were found in the same area. Only 17% of
poor or fragmentary state, so that it is clear the T. pisana shells had been used in
thatin a sample thenests are from a spread contrast to 63% of the Trigonephrus shells.
of years. At sites in the sandveld and coastal dune
slacks to the south of Vanrhynsdorp south
RESULTS
to Cape Town Trigonephrus shells were
The results of the study, including relatively uncommon, increasingly so
published records, are presented in tabular southwards. The percentage of these shells
form in the Appendix. Temporary lodgers, used was consistently over 50% (56%-
those occupants which use the shells only 71%). Only two of the sites visited were
for sleeping and sheltering, have not been free from T. pisana. One was an inland
included. The present survey, however, sandveld site southeastofLutzville and the
confirmed the previous finding (Cess and other was in the coastal dunes at Donkins-
Cess 1999) that there is a very low baai. At these sites Trigonephrus shells were
incidence of the use of shells for this less common than at most of the northern
Volume V7, Number 1, 2008 93
sites but more common than at sites Much habitat destruction in many sand-
infested by T. pisana. At Lamberts Bay veld and coastal sites from Cape Town
and a site north of Blaauwberg only nine north along the west coast at least to
and seven Trigonephrus shells were found Alexander Bay and along the south coast
but T. pisana wasverycommon. Samples of east to East London resulted from the
106, 130 and 134 shells of the latter were introduction of the Australian Acacia Cy-
investigated. Of these only 18%, 24% and clops A. Cunn. ex Loudon (Fabaceae:
12% had been used. At Elands Bay and Mimosoideae) in the second half of the
Yzerfontein no Trigonephrus shells were nineteenth century for the stabilization of
found. Theha pisana shells were extremely sand flats and coastal dunes (Dennill et al.
abundant. Of the 147 shells from Elands 1999). In seriously infested areas this shrub
Bay investigated only 21% had been used has formed extensive monospecific stands
and of 200 and 256 shells from Yzerfontein entirely replacing the species diverse nat-
investigated only 9% and 17% had been ural vegetation.
—
used. Sand-filled shells. Sand-filled shells in
Along the south and east coasts Tropido- windswept areas offer within the spiral
phora ligata has been recorded from the protected sites for the excavation of bur-
Cape Peninsula to Maputo (formerly Lor- rows. This habitatis widelyutilized for this
enzo Marques) (Connolly 1938), however, purposebyQiiartinia (Vespidae: Masarinae)
in the present study shells of this snail in the desertic areas ofsouthwestern Nami-
were found no further west than Port bia and the western Northern Cape south-
Elizabeth but the invasive exotic T. pisana wards in the sandveld and supra-littoral
was found to be extremely common in dunes to CapeTown and eastwards atleast
coastal sand dunes along the south coast toWitsand (Fig. 5). Theonlyotheroccupant
east to Riet River Mouth, the most eastern inthiscategoryencounteredwasTachysphex
site sampled. Use of snail shells along this hermia (Crabronidae: Larrinae),however, as
section ofthe coast was very patchy. By far only a single instance was recorded, in the
the greatestpercentage use of T. pisana was Sperrgebiet, it seems that for this wasp this
at Witsand where there had been least was unusual behaviour. Occasionally a
disturbance by man but where building shell used as a pre-existing cavity for
development was in full swing. At this site nesting by a bee had later fiUed with sand
48% ofshellshadbeenused. Inthe eastthe and been used in addition as a sand-filled
percentage of these shells used was 5.34% shell for nesting by a Quartinia (Fig. 17).
down to 0%. In their initial study Gess and Gess
Although T. ligata shells were abundant (1999) recorded the use of sand-filled snail
at some sites in the east the only site with shells for nesting by two undescribed
an appreciable percentage, 19.9%, of used species of Quartinia. Since then Gess
shells was atKenton-on-Sea in a small area (2007) has described seven species collect-
of dune slacks set aside as abird sanctuary ed from nests in snail shells. Representa-
but now neglected and on the edge of tion of species from snail shells was:
housing development. At Salt Vlei, Port
Alfred from which the use of these shells • in the desertic winter rainfall area north
by a bee, V\lainia (Caposmia) elizabethae was of the Orange River to south of Liideritz
first recorded (as Hoplitis sp. in Gess and Bay, Q. obibensis and Q. refugicola either
Gess 1988), ongoingbuilding development singly or together;
initiated during the past decade has result- • in the desertic winter rainfall area south
ed in the destruction of the dune profile, of the Orange River to Hondeklip Bay,
thenaturalvegetationand associated snails Q. obibensis, Q. refugicola and Q. con-
and insects. chicola:
94 Journalof Hymenoptera Research
• in sandxeld in the vicinity of Hondeklip cells per nest in small, relatively low T.
Bay and Wallekraal, Q. nainaqua and Q. pisana shells.
namaqucnsis (otherwise known only All species nesting in snail shells, except
from the Kamiesberg); Q. australis for which no flower visiting
• in the sandveld area north of Vanrhyns- records were obtained, visit flowers of
dorp, Q. conchicola; "mesems'" (Aizoaceae: Mesembryan-
• in dune slacks along the coast from thema). Quartinia refugicola, however, was
Donkinsbaai, north of Lamberts Bay, collected more commonly from flowers of
south to 4 km north of Bloubergstrand Asteraceae (eight genera) and also from
in the Blaauwberg Conservation Area flowers of Zygophyllaceae (Zygophyllum),
north of Cape Town, Q. honaespei; Neuradaceae (Grielum) and Geraniaceae
• at Yzerfontein on the southwest coast in (Sarcocaulon), all characteristic of the desert
addition, surprisingly, Q. obibensis and areas where it occurs. Quartinia bonaespei in
Q. nainaqua; addition to ''mesems" was collected from
• in dune slacks at Witsand on the south flowers of Trachyandra (Asphodelaceae)
Coast east of Cape Town, Q. australis. which is a characteristic plant of the supra-
littoral dunes of the Western Cape. Full
In the area north ofPortNolloth to south of flower visiting records are given in Gess
Liideritz , in addition to Q. obibensis and Q. (2007). Mating takes place on flowers.
refiigicola, a third closely related species Q. As recorded in Gess and Gess (1999)
vexiUata Gess was collected from flowers at Tricholabiodes sp. (Mutillidae: Sphaer-
several sites but was not collected from opthalminae: Dasylabriini), an undescribed
snail shells. It has been suggested (Gess melanistic species of Allocoelia (Chrysidi-
2007) that this species most probably also dae: Chrysidinae: Allocoeliini) (Fig. 13)
nests in sand-filled snail shells. and a species of Apolysis (Bombyliidae:
All Quartinia nesting in snail shells Usiinae: Usiini) (also in Greathead 1999)
stabilize their nest turrets, burrows and (Fig. 12) were reared from nests of Quarti-
cell walls by the spinning together of sand nia in snail shells from sites in the desertic
grains with self-generated silk as first areas immediately north and south of the
described for Quartinia vagepimctata Orange River.
Schulthess (Gess and Gess 1992). Where Since 1999 further investigation of the
turrets were present they were of the short "Tricholabiodes" specimens has revealed
vertical or slightly curved form with that only one of the specimens, a female,
diameter equal to that of the burrow as from southwest of Brandkaros was in fact
described and illustrated in Gess and Gess Tricholabiodes. Three further females and a
(1999) (Fig. 9). No species constructing bag female each from southeast ofOranjemund
shaped turrets such as are constructed by and east of Alexander Bay are not of the
Q. vagepunctata (described and illustrated genus Tricholabiodes but of an undescribed
in Gess and Gess 1992) were found. genus (Brothers pers. com. April 2007).
Generally the shaft penetrates deep into Only one additional record of a mutillid,
the upper part of the spiral of the shell into probably of one of these genera, from
which the cells are closely packed (Fig. 13), Quartinia cells has been obtained. The
however, in some instances cell construc- new record is for the Drachenberg, the
tion had been commenced lower in the most northern site at which Trigonephrus
spiral. The number of cells that can be was encountered. It appears that the
constructed is limited by the volume of the frequency of occurrence of mutillids in
shell. The number of cells per nest in the Quartinia nests in snail shells is low, only
large, relatively high Trigonephrus shells seven records having been obtained from
can therefore be many times the number of 1,269 nests.
Volume 17, Number 1, 2008 95
considered that it was probably A. capicola
Hesse (Greathead 1999). However, subse-
quent sampling of snail shells has provid-
ed adult female and male specimens from
Quartinia nests from four additional sites
from the Orange River north to the
Klinghardtberge. The specimens have
made it possible to determine the species
as Apolysis hesseana (Greathead 2006). Apo-
lysis hesseana was described briefly from a
single denuded female specimen from the
Northern Cape (Evenhuis and Greathead
1999) but the new material enabled a full
description of the species to be given
(Greathead 2006). In nests from which
Apolysis has emerged the empty pupal
cuticle is found attached to the outside of
a Quartinia cell. Itappears thatfrequencyof
occurrence ofApolysis in Quartinia nests in
shells is low, only 14 records having been
obtained from 1,269 nests but more fre-
quent than by mutillids and Allocoelia.
Indeed overall in the areas sampled the
frequency of parasitism of Quartinia nests
in snail shells appears to be low, only 29
Figs 12-13. 12,CellsandfemaleofQuartiniaobibensis, records having been obtained from 1,269
andassociatedApolysishesseana,adultX 1.5(scalebar= nests. It is of interest that no incidents of
2cm). 13, Cells and female of Quartina refugicola, and parasitism were recorded from south of
associated Allocoelia n. sp., cocoon and imagine X 1.5
(scalebar = 2cm). Port Nolloth. —
Empty shells, empty snail shells were
Similarly the only new record ofAllocoe- found to be utilized as nesting cavities by
lia from Quartinia cells since the rearings bees throughout the study area. The only
from south of Rosh Pinah and east of Port other aculeate using the empty shells was
Nolloth is from the Drachenberg. It is Alastor ricae (Vespidae: Eumeninae) previ-
probable that it is present throughout the ouslyrecorded (as ''a eumenine'', Gess and
desertic areas to the south and north of the Gess 1999) from sites in the vicinity of
Orange River, however, it would appear Brandkaros to the south of the Orange
that frequency ofoccurrence ofAllocoelia in River and presumed to be the same
Quartinia nests in snail shells is low, only ''Eumenid-wasp" referred to by Hesse
eight records having been obtained from (1944) as nesting in empty snail shells in
1,269 nests. Allocoelia pupates outside the "the very arid and sandybelt ofthe Namib
Quartinia cell in which it developed so that desert''. In subsequent sampling of snail
in nests from which ithas emerged its past shells by the present authors its nests have
presence is readily visible in the form of its been found at 14 sites from north of the
empty cocoon attached to the outside of a Orange River to the Drachenberg and a
Quartinia cell. sample of snail shells collected by the
In 1999 the species identity of the BIOTA-Southern Africa Project at Groot-
Apolysis could not be established as only derm to the south of the Brandkaros sites
a pharate adult was available. It was yielded two specimens (Koch 2006). Even
