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Pupa menkeana Pfeiffer, 1853, Type Species of the Speciose Land Snail Genus Gulella Pfeiffer, 1856: Correction of Longstanding Misidentification and Designation of Neotype (Mollusca: Eupulmonata: Streptaxidae)

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Gulella menkeana (Pfeiffer, 1853) is the type species of the highly diverse genus Gulella Pfeiffer, 1856 and study of its morphology and DNA is critical to our ability to define the genus sensu stricto in relation to research on the systematics of this complex genus. However, we have established that current use of the name is inconsistent with both the original description and first figure of the species. The name-bearing type is lost, and purported paratypes in the Museum für Naturkunde der Humboldt-Universität, Berlin, on which Connolly's widely-followed redescription was based, are not authentic. They have no type status and are instead specimens of Gulella albersi (Pfeiffer, 1855). Recently collected specimens conforming to Pfeiffer's description and figure of G. menkeana have been identified and a neotype is designated. This material also corresponds with the current broad interpretation of G. adamsiana (Pfeiffer, 1859), which, with its several established synonyms, we include in the synonymy of G. menkeana. There is also considerable resemblance to G. wahlbergi (Krauss, 1848), a name based on composite material and for which the lectotype is lost. We also designate a neotype for this species so as to preserve current application of the name. The possible synonymy of G. menkeana and G. wahlbergi, an older name, is of nomenclatural concern, but is not critical to the issue of defining Gulella s.s.
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233
African Invertebrates Vol. 52 (2) Pages 233–242 Pietermaritzburg December, 2011
http://www.africaninvertebrates.org.za
Pupa menkeana Pfeiffer, 1853, type species of the speciose land
snail genus Gulella Pfeiffer, 1856: correction of longstanding
misidentication and designation of neotype
(Mollusca: Eupulmonata: Streptaxidae)
D. G. Herbert1 and B. Rowson2
1KwaZulu-Natal Museum, P. Bag 9070, Pietermaritzburg, 3200 South Africa and School of Biological &
Conservation Sciences, University of KwaZulu-Natal, Scottsville, 3206 South Africa; dherbert@nmsa.org.za
2Department of Biodiversity & Systematic Biology, National Museum of Wales, Cathays Park, Cardiff
CF10 3NP, UK; ben.rowson@museumwales.ac.uk
ABSTRACT
Gulella menkeana (Pfeiffer, 1853) is the type species of the highly diverse genus Gulella Pfeiffer, 1856
and study of its morphology and DNA is critical to our ability to dene the genus sensu stricto in relation
to research on the systematics of this complex genus. However, we have established that current use of the
name is inconsistent with both the original description and rst gure of the species. The name-bearing
type is lost, and purported paratypes in the Museum für Naturkunde der Humboldt-Universität, Berlin, on
which Connolly’s widely-followed redescription was based, are not authentic. They have no type status
and are instead specimens of Gulella albersi (Pfeiffer, 1855). Recently collected specimens conforming
to Pfeiffer’s description and gure of G. menkeana have been identied and a neotype is designated.
This material also corresponds with the current broad interpretation of G. adamsiana (Pfeiffer, 1859),
which, with its several established synonyms, we include in the synonymy of G. menkeana. There is also
considerable resemblance to G. wahlbergi (Krauss, 1848), a name based on composite material and for
which the lectotype is lost. We also designate a neotype for this species so as to preserve current application
of the name. The possible synonymy of G. menkeana and G. wahlbergi, an older name, is of nomenclatural
concern, but is not critical to the issue of dening Gulella s.s.
KEY WORDS: South Africa, Gulella menkeana, adamsiana, wahlbergi, type species, misidentication,
original gure, new synonyms, neotypes.
INTRODUCTION
The genus Gulella Pfeiffer, 1856 (Streptaxidae) is perhaps the most species-rich
genus of African land snails (Bruggen 1967; Richardson 1988; Schileyko 2000). How-
ever, as presently conceived, it includes a diverse array of species and much of the
supraspecic taxonomy of genus is based on very limited evidence, with little con-
sensus concerning the characterisation, distinctness and usage of the described sub-
generic entities. Furthermore, given recent systematic work (Rowson et al. 2010) and
the diversity of form exhibited in Gulella s.l., it seems probable that it is not a mo-
nophyletic radiation, instead comprising a polyphyletic assemblage of variously re-
lated lineages.
As an early step in a study examining phylogenetic relationships within the Strep-
taxidae and Gulella in particular, we propose to dene Gulella in terms of its type
spe cies, Pupa menkeana Pfeiffer, 1853, thus circumscribing Gulella s.s. in greater
de tail by providing morphological data on shell microsculpture, radula tooth form and
genital tract anatomy, as well as molecular sequence data. To do this unequivocally,
it is essential that the material studied be correctly identied. In the process of con-
rming identications it has become apparent to us that the current interpretation of
Gulella menkeana is at variance with the rst illustration of the species provided by
Pfeiffer (1859 in 1854–1860). Further investigation has revealed that this name has
234 AFRICAN INVERTEBRATES, VOL. 52 (2), 2011
been consistently applied incorrectly in the literature pertaining to South African land
snails. In this paper we detail the historical misidentication of G. menkeana and pro-
vide instead a more plausible interpretation of the species in terms of the ori ginal de-
scription and the rst illustration. To clarify its identity and thus that of Gulella itself,
we designate a neotype for the species. It is important that this matter be re solved be-
fore further research is undertaken on the species in relation to streptaxid systematics.
INSTITUTIONAL CODENS
NHMUK Natural History Museum, London, UK.
NMSA KwaZulu-Natal Museum, Pietermaritzburg, South Africa.
SMF Senckenberg Museum, Frankfurt, Germany.
SMNH Swedish Museum of Natural History, Stockholm, Sweden.
ZMHB Museum für Naturkunde der Humboldt-Universität, Berlin, Germany.
TAXONOMY
Genus Gulella Pfeiffer, 1856
Gulella: Pfeiffer 1856a: 173. Type species Pupa menkeana Pfeiffer, 1853 [S.D. Martens 1860: 298].
Gulella menkeana (Pfeiffer, 1853)
Pupa menkeana: Pfeiffer 1853: 552, No 161. Type loc.: Port Natal [= Durban, South Africa].
Ennea menkeana: Pfeiffer 1856b: 61; idem 1859 in 1854–1860: 113, pl. 32, gs 3–5.
Pfeiffer (1853) described Pupa menkeana from material in the collection of German
conchologist K.T. Menke (1791–1861) and gave Port Natal [= Durban, South Africa] as
the only locality. After Menke’s death, his collection was sold to a natural history dealer,
M.J. Landauer of Frankfurt, and dispersed via retail sale to private collectors (Zilch
1958). Thereafter, the location of the original material of G. menkeana has remained
unknown and Connolly (1939: 39) stated ‘the type appears to be lost’. However, he
iden tied two shells in the ‘Berlin Museum’ (ZMHB) as ‘paratypes’ and based his
description of the species on one of these, also providing a gure. In terms of locality
data, he repeated only the ‘Port Natal’ locality. In this same work (p. 39), Connolly
discussed Gulella albersi (Pfeiffer, 1855), stating that this was ‘merely a large edition
of menkeana, with which it is probably conspecic’. For G. albersi Connolly cited, in
addition to the type locality, several localities on the KwaZulu-Natal south coast (see
note below regarding type locality of this species). It is Connolly’s description and -
gure of the ZMHB paratype and his interpretation of G. menkeana that has informed
sub sequent identication and discussion of this species and its distribution (Aiken
1995; Herbert & Kilburn 2004), and the belief that G. albersi is a junior synonym.
In the process of assembling a formal synonymy and list of citations for Gulella
menkeana, and providing illustrations of the relevant type specimens, we have disco-
vered rstly, that the ZMHB paratypes do not correspond with the rst gure of the
species provided by Pfeiffer (1859 in 1854–1860: pl. 32, g. 4, here reproduced in
Fig. 1A), and secondly that there appears to be no justication for considering them
to be paratypes in the current sense of the Code. They were part of the collection of
J.C. Albers, who evidently obtained them from R.J. Shuttleworth. There is no label to
indicate that they were ever part of the Menke collection or that they represent type
material of any kind (Glaubrecht pers. comm. 24.iii.2011). Why Connolly should have
HERBERT & ROWSON: IDENTITY OF THE TYPE SPECIES OF GULELLA 235
considered them to be paratypes is not known and he was evidently not justied in so
doing.
When compared with Pfeiffer’s illustration (Fig. 1A) the upper labral tooth of the
Albers specimens (Figs 1B, 1C) is a far less robust structure. It is smaller than the
lower labral tooth and does not have a basal buttress on its upper side that lies almost
parallel to the parietal lamella. Conversely, the lower labral tooth in Pfeiffer’s gure is
smaller than the upper one and is clearly inset. In addition, the basal tooth is large and
trigonal in Pfeiffer’s gure, but narrow in the Albers specimens. The columella lamella
is poorly dened in Pfeiffer’s gure, but the Latin description, although not detailed in
terms of apertural tooth morphology, stated that the second tooth (what is now termed
the columella lamella – he worked anticlockwise, starting with the parietal lamella) is
excavata profunde ad columellam’ [deeply excavated at/toward the columella]. This
description does not match the columella lamella of the Albers specimens in which
this structure takes the form of a horizontal ridge-like tooth. In contrast, in the original
Fig. 1. (A) gure of Pupa menkeana (as Ennea) given by Pfeiffer (1859 in 1854–1860: pl. 32, g. 4); (B,
C) ZMHB specimens considered by Connolly (1939) to be paratypes of P. menkeana Pfeiffer, 1853,
length 13.5 mm and 13.6 mm (ZMHB 56871); (D) lectotype of Pupa albersi Pfeiffer, 1855, ‘Cape
Natal, Mus. Cuming’, length 15.3 mm (NHMUK 20110169); (E) Senckenberg Museum specimen
gured as Gulella menkeana by Zilch (1960, in 1959–1960: 571, g. 2000), ‘Natal’, length 10 mm
(SMF 83755, photograph S. Hof); (F, G) P. menkeana Pfeiffer, 1853, neotype, Burman Bush,
Durban, length 9.84 mm, diameter 5.05 mm (NMSA W7943/T2670).
236 AFRICAN INVERTEBRATES, VOL. 52 (2), 2011
description of G. albersi, Pfeiffer (1855) described the columella lamella as ‘compressa,
prominentiae umbilicali transverse imposita’ [attened, positioned transverse to the
umbilicus] which is an apt description of this structure in the Albers specimens. In
terms of size, these specimens (length 13.5 and 13.6 mm) are also closer to G. albersi
(length given as 15 mm) than they are to G. menkeana (length given as 11 mm). In
fact, these Albers specimens at ZMHB exhibit all the features of G. albersi and their
identication as G. menkeana is consistent with neither the original description nor the
rst illustration of that species. They are evidently referable to G. albersi (lectotype
[designated Connolly 1939: 39] illustrated in Fig. 1D), thus explaining Connolly’s view
that G. albersi was merely a large edition of G. menkeana. The description and gure
of G. menkeana show it to possess characters distinct from those of G. albersi and the
two names should no longer be considered synonyms. This conclusion is further
supported by the fact that Pfeiffer discussed and gured G. albersi in the same work
in which he gured G. menkeana for the rst time (Pfeiffer 1854 –1860), clearly in-
dicating that he considered them to be different species. Thus Connolly’s concept of
G. menkeana was based on misidentied specimens with no type status. Consequently,
material collected subsequent to his monograph (Connolly 1939) and identied in ac-
cor dance with his description of G. menkeana is misidentied and in fact represents
G. albersi (Aiken 1995; Herbert & Kilburn 2004; Rowson et al. 2010). The question
that then arises is — what is the real Gulella menkeana?
WHAT IS THE REAL GULELLA MENKEANA?
In the absence of authentic type material of Pupa menkeana, we have only Pfeiffer’s
brief original description (Pfeiffer 1853), and his subsequent illustration (reproduced in
Fig. 1A) to guide us in determining the true identity of this taxon. Other treatments and
gures of the species, prior to Connolly’s monograph (e.g. Sowerby 1878; Tryon 1885;
Möllendorff & Kobelt 1904 in 1903–1905), are of little value since these largely repeat
or abbreviate Pfeiffer’s earlier description and copy his gure. More recent illustrations
of specimens in European museums, most probably collected in the late 1800s or early
1900s (Zilch 1960 in 1959–1960: 571, g. 2000; Schileyko 2000: g. 1067A) show a
species with a large, buttressed upper labral tooth, suggesting that the early European
concept of the species was in accordance with the original gure and at variance with
Connolly’s later interpretation (Connolly 1939). However, the specimens concerned
were not types and the illustrations are thus not denitive. In reality, a photograph of
the specimen illustrated by Zilch (1960) reveals it to be a specimen of Gulella wahl-
bergi (Krauss, 1848) (Fig. 1E, courtesy of R. Janssen) and earlier labels written by
Werner Blume identify it as such (Janssen pers. comm. 13.iv.2011).
Apertural dentition is critical in the identication of Gulella species (e.g. Connolly
1939; Verdcourt 1962; Herbert & Kilburn 2004) and, as already emphasised, the form
of the upper labral tooth and its juxtaposition almost parallel to the parietal lamella,
as well as the smaller size and inset position of the lower labral tooth, are perhaps the
most signicant features evident in Pfeiffer’s gure of G. menkeana. Such a pattern of
labral dentition is shown by several other Gulella species, including other similarly-
sized, axially ribbed species found in the Durban area, namely G. adamsiana (Pfeiffer,
1859) and G. wahlbergi.
HERBERT & ROWSON: IDENTITY OF THE TYPE SPECIES OF GULELLA 237
The current view of G. adamsiana is that it is a relatively widely distributed taxon
(north-eastern Eastern Cape to northern KwaZulu-Natal, from the coast to altitudes of
ca 1300 m) and it correspondingly exhibits considerable variation in size, shell pro-
por tions and strength of apertural dentition (Bruggen 1980; Herbert & Kilburn 2004).
Some of this variation, particularly shell size, may be linked to differences in habitat,
specimens from drier thornveld habitats at inland localities being smaller than those
from more mesic coastal forests. However, the format of the aper tu ral dentition re-
mains essentially the same, and although the strength and shape of the individual aper-
tural teeth may vary, there appears to be no clear pattern in this va riation. The shells
illustrated in Fig. 2, including type specimens of G. adamsiana and its various sy nonyms
are illustrative of this variation. We concur with Burnup (in Connolly 1932) that G.
socratica (Melvill & Ponsonby, 1893) is based on a deformed spe cimen of G. adam-
siana. The species is evidently prone to abnormalities (Warren 1933; Bruggen 1980).
Specimens referable to this concept of G. adamsiana, collected in Durban (the type
locality for Pupa menkeana), closely resemble Pfeiffer’s gure of Pupa menkeana and
are concordant also with the original description. We believe that these can legitimately
be considered to represent the species named Pupa menkeana by Pfeiffer (1853). Since
this description predates that of Pupa adamsiana Pfeiffer, 1859, the latter must be con-
sidered a junior synonym. So as to stabilise this nomenclature we designate as neo type
for Pupa menkeana a specimen from this Durban population (see below).
Krauss’ G. wahlbergi also resembles Pfeiffer’s gure of G. menkeana and Bruggen
(1980) has noted the considerable similarity between the former and G. adam siana.
However, he observed that G. wahlbergi differs (inter alia) in having a relatively nar-
row ba sal denticle, in the form of an in-running ridge as opposed to a trigonal or sub-
quadrate, trans versely-set peg (see also Connolly 1939; Herbert & Kilburn 2004).
Pfeiffer’s gure of G. menkeana clearly shows the basal denticle as a trigonal structure
like that of many G. adamsiana specimens and unlike that of G. wahlbergi. We follow
Bruggen (1980) in considering Helix fanulus Pfeiffer, 1856, from ‘Port Natal’ which
Connolly (1939) associated with G. adamsiana, to be an unidentiable juvenile Gulella
and thus a nomen dubium.
An updated synonymy for G. menkeana incorporating these nomenclatural changes
is given below. In so redening the species, we are aware that adherence to the original
concept of G. menkeana might be considered to conict with prevailing usage of the
name. However, the species has been rarely mentioned in the literature, beyond mere
mention of the name as the type species of Gulella. We are aware of only three instances
where the species, as conceived by Connolly (1939), has been cited subsequently in
print (Aiken 1995; Herbert & Kilburn 2004; Rowson et al. 2010). Conversely, the ori-
ginal concept of the species has not completely fallen out of use and was employed by
Schileyko (2000). Since both concepts of the species have been employed in relatively
recent times, it is logically correct to expunge the one based on an error and to employ
the name in a manner consistent with the original description and gure.
Gulella menkeana Pfeiffer, 1853
Pupa menkeana: Pfeiffer 1853: 552, No. 161. Type loc.: Port Natal [= Durban, South Africa]. Neotype
de signated herein.
Ennea menkeana: Pfeiffer 1856b: 61; 1859 in 1854–1860: 113, pl. 32, gs 3–5; Tryon 1885: 97, pl. 18, g.
79; Melvill & Ponsonby 1898b: 168; Sturany 1898: 15 [555]; Connolly 1912: 79.
238 AFRICAN INVERTEBRATES, VOL. 52 (2), 2011
Ennea adamsiana: Pfeiffer 1859 in 1854 –1860: 114, pl. 32, gs 9–11; 1859: 339; Melvill & Ponsonby
1898b: 166. Type loc.: Port Natal. Lectotype designated by Connolly (1939: 84). Syn. n.
Gulella menkeana: Martens 1860: 298; Zilch 1960 in 1959 –1960: 571, g. 2000; 1961: 95; Richardson
1988: 101; Schileyko 2000: 816, g. 1067A.
Ennea (Gulella) menkeana: Pfeiffer 1878 in 1878–1881: 19; Möllendorff & Kobelt 1904 in 1903–1905:
191, pl. 24, g. 6; Kobelt 1909: 54; 1910: 161.
Pupa menkeana: Sowerby 1878, pl. 19, g. 176 [confused with Carychium menkeanum Pfeiffer, 1821
= Azeca goodalli (Férussac, 1821) and erroneous locality given].
Enneastrum menkeanum: Bourguignat 1889: 127.
Ennea socratica: Melvill & Ponsonby 1893: 109, pl. 3, g. 14. Type loc.: Pietermaritzburg. Syn. n.
Ennea impervia: Melvill & Ponsonby 1896: 315, pl. 16, g. 1; 1898b: 168. Type loc.: Natal. Syn. n.
Ennea aurisleporis: Melvill & Ponsonby 1898a: 25, pl. 8, g. 3; 1898b: 167. Type loc.: Natal. Syn. n.
Gulella adamsiana: Connolly 1939: 84, text-g. 5; Bruggen 1980: 7, gs 1, 3 [see for additional citations];
Richardson 1988: 50; Aiken 1995: 18; Herbert & Kilburn 2004: 195.
Gulella adamsiana var. impervia: Connolly 1939: 85.
Gulella aurisleporis: Connolly 1939: 86; Richardson 1988: 51; Aiken 1995.
Gulella impervia: Richardson 1988: 51.
Gulella socratica: Richardson 1988: 51.
? Helix fanulus: Pfeiffer 1856c: 33. Type loc.: Port Natal [=Durban, South Africa]. Nomen dubium
non Gulella menkeana: Connolly 1939: 38, pl. 1, g. 15; Aiken 1995: 7; Herbert & Kilburn 2004: 169;
Rowson et al. 2010: 10 [= Gulella albersi Pfeiffer, 1855].
DESIGNATION OF NEOTYPE FOR PUPA MENKEANA PFEIFFER, 1853
Since Pupa menkeana is the type species of the genus Gulella Pfeiffer, 1856, its
identity is of critical importance in dening the genus and it is vital that the name be
applied correctly. We believe, in accordance with Article 75.3.4 of the Code (ICZN
1999), that the original type material of Pupa menkeana was lost when Menke’s col-
lec tion was sold and dispersed to private collectors after his death (Zilch 1958). As dis-
cussed above, the purported paratypes in the ZMHB (Connolly 1939) are neither types
nor are they referable to G. menkeana (they are in reality specimens of G. albersi).
Having now identied material that matches Pfeiffer’s description and gure of P.
men keana more closely than any other material known to occur in the province of
KwaZulu-Natal, South Africa, we consider it is necessary to designate a neotype for
the taxon in order to remove any further doubt concerning the species represented by
this name. The specimen (Figs 1F, 1G) is selected from a population occurring at the
type locality (given only as ‘Port Natal’ = Durban). In addition to the neotype, we
have collected from this same population a growth series of shells, as well as live-
col lected specimens preserved for anatomical study and tissue samples for molecular
se quencing (NMSA W7878, W7896). This material will enable us to dene the species
and thus the genus Gulella s.s. in terms of features of the adult and embryonic shell,
ra dula teeth, reproductive tract morphology and molecular sequence data (Rowson &
Herbert, in prep).
We consider that the neotype is also identiable as the larger coastal form of what
has been known as the variable Gulella adamsiana (Figs 2A, 2B) and as stated above
consider G. adamsiana and G. menkeana to be conspecic and therefore synonyms.
The distinguishing features of this species and the extent of its intraspecic variability
have been discussed in detail under the name G. adamsiana by Bruggen (1980).
Neotype: SOUTH AFRICA: KwaZulu-Natal: Durban, Burman Bush, beside road near scout camp, 29.81490°S:
31.01740°E, 75 m, in accumulations of leaf-litter at roadside, Station 11-05, 29.iii.2011, D. Herbert & L.
Davis (NMSA W7943/T2670). Length 9.84 mm, diameter 5.05 mm.
HERBERT & ROWSON: IDENTITY OF THE TYPE SPECIES OF GULELLA 239
Fig. 2. (A, B) lectotype and paralectotype of Ennea adamsiana Pfeiffer, 1859 in 1854–1860, ‘Port Na tal,
M.C. [Mus. Cuming]’, length 8.45 and 8.1 mm (NHMUK 20110168); (C, D) syn types of Ennea
im per via Melvill & Ponsonby, 1896, ‘Natal’, length 8.8 and 8.75 mm (NHMUK 1903.3.11.85–86);
(E) holotype of Ennea aurisleporis Melvill & Ponsonby, 1898, ‘Natal’, length 6.65 mm (NHMUK
1903.3.11.69); (F) holotype of Ennea socratica Melvill & Ponsonby, 1893, ‘Pietermaritzburg’,
length 8.45 mm (NHMUK 1903.3.11.78); (G) Gulella menkeana, small inland form, Cumberland,
Pietermaritzburg, length 7.2 mm (NMSA V9709); (H) G. menkeana, large form from coastal E.
Cape, Port St Johns, length 9.24 mm (NMSA W561); (I) Pupa wahlbergi Krauss, 1848, probable
paratype in SMNH, length 10.1 mm (SMNH-Type-2112); (J, K) P. wahlbergi Krauss, 1848, neotype,
Durban Bluff, length 9.84 mm, diameter 4.92 mm (NMSA W7942/T2669); (L) holotype of Ennea
formosa Melvill & Ponsonby, 1898, Pietermaritzburg, length 7.75 mm (NHMUK 1903.3.11.79).
240 AFRICAN INVERTEBRATES, VOL. 52 (2), 2011
CAVEAT
We acknowledge that our use of the name Gulella menkeana as an earlier name
for the species currently known as G. adamsiana may ultimately need to be revised.
The prevailing broad interpretation of G. adamsiana includes small, narrow specimens
from drier inland localities (Fig. 2G), as well as larger, broader specimens with strong
apertural dentition from the coast (Figs 2C, 2D form impervia) and a somewhat dis junct
population of similarly large specimens from the central coastal area of the Trans kei
region, Eastern Cape (Fig. 2H). In due course, phylogeographic analysis of mo lecular
data may reveal this to be a composite taxon. However, this is immaterial to the issue
at hand, the crux of which is to verify the identity of G. menkeana. Should the species
eventually be shown to be composite, this will not change the fact we have established
the true identity of G. menkeana.
Ultimately, if the broad interpretation of a morphologically variable G. menkeana
proves to be robust, it may also include the form currently known as G. wahlbergi.
This taxon differs from G. menkeana only in relatively small details that could be
subsumed within the variability of one species. If such is the case, since it is an earlier
name (1848), it would take precedence over G. menkeana (1853). Again, however, this
would not detract from the fact that we have xed the identity of G. menkeana for the
purposes of dening Gulella s.s.
Connolly (1939) indicated that the type material of Pupa wahlbergi Krauss, 1848
in Stutt gart (two specimens, both now lost) was composite and selected as ‘the type’
[= lectotype] an axially costulate specimen with a supercial tooth on the columella
which matched Krauss’ gure. A lot containing three probable paralectotypes of P.
wahlbergi in the SMNH is also composite (Herbert & Warén 1999). Only one, a some-
what damaged specimen, is costulate and has a supercial tooth on the columella
(Fig. 2I). Although damaged, this specimen conforms with Krauss’ original and most
subsequent illustrations of the taxon (Küster 1855 in 1841–1855: 158, pl. 19, gs 6–9
(plate dated 1854); Sowerby 1878: pl. 20, g. 187; Möllendorff & Kobelt 1904 in 1903–
1905: 190, pl. 24, gs 3, 4; Burnup 1925: pl. 8, g. 35) and with current application of
the name (Connolly 1939; Bruggen 1980; Herbert & Kilburn 2004). Tryon’s illustration
(Tryon 1885: 96, pl. 19, g. 99), as pointed out by Burnup (1925), erroneously shows
three labral teeth, and seems to have been inuenced by Pfeiffer’s inclusion of the
basal tooth as a third labral tooth (Pfeiffer 1848). The remaining two specimens are
smooth except for axial pliculae radiating onto the base from the umbilicus and lack
a supercial columella tooth. They resemble Gulella kosiensis (Melvill & Ponsonby,
1908) but are considerably larger (length 9.2 mm) than any other specimens referable
to that species (length up to 7.0 mm) and their identity is puzzling. Since P. wahlbergi
was based on material of more than one species (evident also in Krauss’ description),
we consider it necessary designate a neotype for the taxon such that application of the
name will preserve prevailing usage. As the single SMNH specimen reecting this
usage is in poor condition (Fig. 2I) we prefer (as permitted by Art. 75 of the Code) to
se lect a more recently collected, undamaged, topotypic specimen as the neotype (Figs
2J, 2K): Durban Bluff, length 9.84 mm, diameter 4.92 mm (NMSA W7942/T2669).
G. formosa (Melvill & Ponsonby, 1898) from the KwaZulu-Natal Midlands (holo-
type Fig. 2L), though generally less strongly sculptured, may also fall within this va-
riable concept of G. menkeana and represent a mist-belt ecomorph, though we refrain
HERBERT & ROWSON: IDENTITY OF THE TYPE SPECIES OF GULELLA 241
from proposing synonymy at this stage. The locality ‘Durban’ given in the original
description is dubious. That of ‘Pietermaritzburg’ cited on the labels in the type lot
(NHMUK 1903.3.11.79) is more probable.
NOTE ON THE TYPE LOCALITY OF GULELLA ALBERSI (PFEIFFER, 1855)
For his new taxon Pupa albersi, Pfeiffer (1855) gave as the locality ‘Port Natal
(Stan ger)’ [later misspelt as ‘Strangier’ (Pfeiffer, 1859: 339)]. Subsequently, Connolly
(1939) cited several additional localities on the KwaZulu-Natal south coast (Scottburgh,
Port Shepstone, Port Edward). In reality, there are to date no conrmed records for this
species from further north than Scottburgh (30.288°S). It is not known from the Durban
area and it seems probable that the Port Natal locality was simply an imprecise one
referring to the KwaZulu-Natal coast. Stanger is a more precise locality, but lies 120 km
to the north of the known distribution of the species. Given that the malacofauna of this
region is relatively well known and that the south coast of KwaZulu-Natal is home to
other locally endemic land snails that do not range as far north as Durban (Herbert &
Kilburn 2004), we believe that the original locality data must be considered imprecise
in the case of ‘Port Natal’ and erroneous in the case of ‘Stanger’. Errors such as this are
not unusual for material in the Cuming collection. We take this opportunity to emend
the type locality to Port Shepstone, where the species is particularly common.
ACKNOWLEDGEMENTS
For the loan of types, and assistance in examination and photography of specimens we thank J. Ablett
(NHMUK), M. Glaubrecht and Ch. Zorn (ZMHB), R. Janssen (SMF) and A. Warén (SMNH).
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