Gulella menkeana Pfeiffer, 1853

Herbert, D. G. & Rowson, B., 2011, Pupa menkeana Pfeiffer, 1853, type species of the speciose land snail genus Gulella Pfeiffer, 1856: correction of longstanding PLVLGHQWL ¿ FDWLRQ DQG GHVLJQDWLRQ RI QHRW \ SH (Mollusca: Eupulmonata: Streptaxidae), African Invertebrates 52 (2), pp. 233-233 : 237-241

Gulella menkeana Pfeiffer, 1853

Pupa menkeana: Pfeiffer 1853: 552 , No. 161. Type loc.: Port Natal [= Durban, South Africa]. Neotype designated herein.

Ennea menkeana: Pfeiffer 1856 b: 6 l; l8ƽ9 LQ l8ƽ4±l86Ɵ: ll³, SO. ³2, ¿JV ³±ƽ; 7U\RQ l88ƽ: 97, SO. l8, ¿J. 79; Melvill & Ponsonby 1898 b: 168; Sturany 1898: 15 [555]; Connolly 1912: 79.

Ennea adamsiana : 3IHLIIHU l8ƽ9 LQ l8ƽ4±l86Ɵ: ll4, SO. ³2, ¿JV 9±ll; l8ƽ9: ³³9; 0HOYLOO & 3RQVRQE\ 1898 b: 166. Type loc.: Port Natal. Lectotype designated by Connolly (1939: 84). Syn. n.

Gulella menkeana : 0DUWHQV l86Ɵ: 298; =LOFK l96Ɵ LQ l9ƽ9±l96Ɵ: ƽ7l, ¿J. 2ƟƟƟ; l96l: 9ƽ; 5LFKDUGVRQ l988: lƟl; 6FKLOH\NR 2ƟƟƟ: 8l6, ¿J. lƟ67$.

Ennea (Gulella) menkeana: Pfeiffer 1878 in 1878–1881: 19; Möllendorff & Kobelt 1904 in 1903–1905: l9l, SO. 24, ¿J. 6;.REHOW l9Ɵ9: ƽ4; l9lƟ: l 6l.

Pupa menkeana : 6RZHUE\ l878, SO. l9, ¿J. l76>FRQIXVHG ZLWK Carychium menkeanum Pfeiffer, 1821 = Azeca goodalli (Férussac, 1821) and erroneous locality given].

Enneastrum menkeanum: Bourguignat 1889: 127 .

Ennea socratica : 0HOYLOO & 3RQVRQE\ l89³: lƟ9, SO. ³, ¿J. l4. 7\SH ORF.: 3LHWHUPDULW]EXUJ. Syn. n.

Ennea impervia : 0HOYLOO & 3RQVRQE\ l896: ³lƽ, SO. l6, ¿J. l; l898 b: 168. Type loc.: Natal. Syn. n.

Ennea aurisleporis: Melvill & Ponsonby 1898 a: 2 ƽ, SO. 8, ¿J. ³; l898 b: 167. Type loc.: Natal. Syn. n.

Gulella adamsiana : &RQQROO\ l9³9: 84, WH[W‒¿J. ƽ; %UXJJHQ l98Ɵ: 7, ¿JV l, ³>VHH IRU DGGLWLRQDO FLWDWLRQV @; 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 1856 c: 33 . Type loc.: Port Natal [=Durban, South Africa]. Nomen dubium non Gulella menkeana : &RQQROO\ l9³9: ³8, SO. l, ¿J. lƽ; $LNHQ l99ƽ: 7; +HUEHUW &.LOEXUQ 2ƟƟ4: l69; 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 LGHQWLW\ LV RI FULWLFDO LPSRUWDQFH LQ GH¿QLQJ WKH JHQXV DQG LW LV YLWDO WKDW WKH QDPH EH 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 collection was sold and dispersed to private collectors after his death ( Zilch 1958). As discussed 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 ). +DYLQJ QRZ LGHQWL¿HG PDWHULDO WKDW PDWFKHV 3IHLIIHU¶V GHVFULSWLRQ DQG ¿JXUH RI P. menkeana 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 livecollected specimens preserved for anatomical study and tissue samples for molecular VHTXHQFLQJ (106$:7878,:7896). 7KLV PDWHULDO ZLOO HQDEOH XV WR GH¿QH WKH VSHFLHV and thus the genus Gulella s.s. in terms of features of the adult and embryonic shell, radula teeth, reproductive tract morphology and molecular sequence data (Rowson & Herbert, in prep).

: H FRQVLGHU WKDW WKH QHRW\SH LV DOVR LGHQWL¿DEOH DV WKH ODUJHU FRDVWDO IRUP RI ZKDW has been known as the variable Gulella adamsiana ( Figs 2A, 2B ) and as stated above consider G. adamsiana and G. menkeana WR EH FRQVSHFL ¿F DQG WKHUHIRUH V\QRQ\PV. 7KH GLVWLQJXLVKLQJ IHDWXUHV RI WKLV VSHFLHV DQG WKH H[WHQW RI LWV LQWUDVSHFL¿F YDULDELOLW\ 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.

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 disjunct population of similarly large specimens from the central coastal area of the Transkei region, Eastern Cape ( Fig. 2H ). In due course, phylogeographic analysis of molecular 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 ZRXOG QRW GHWUDFW IURP WKH IDFW WKDW ZH KDYH ¿[HG WKH LGHQWLW\ RI G. menkeana for the SXUSRVHV RI GH¿QLQJ Gulella s.s.

Connolly (1939) indicated that the type material of Pupa wahlbergi Krauss, 1848 in Stuttgart (two specimens, both now lost) was composite and selected as ‘the type’>=OHFWRW\SH@ DQ D[LDOO\ FRVWXODWH VSHFLPHQ ZLWK D VXSHU¿FLDO WRRWK RQ WKH FROXPHOOD ZKLFK PDWFKHG.UDXVV¶ ¿JXUH. $ ORW FRQWDLQLQJ WKUHH SUREDEOH SDUDOHFWRW\SHV RI P. wahlbergi in the SMNH is also composite ( Herbert & Warén 1999). Only one, a some- ZKDW GDPDJHG VSHFLPHQ, LV FRVWXODWH DQG KDV D VXSHU¿FLDO WRRWK RQ WKH FROXPHOOD ( Fig. 2I ). Although damaged, this specimen conforms with Krauss’ original and most VXEVHTXHQW LOOXVWUDWLRQV RI WKH WD[RQ (.üVWHU l8ƽƽ LQ l84l±l8ƽƽ: lƽ8, SO. l9, ¿JV 6±9 (SODWH GDWHG l8ƽ4); 6RZHUE\ l878: SO. 2Ɵ, ¿J. l87; 0|OOHQGRUII &.REHOW l9Ɵ4 LQ l9Ɵ³± l9Ɵƽ: l9Ɵ, SO. 24, ¿JV ³, 4; %XUQXS l92ƽ: SO. 8, ¿J. ³ƽ) DQG ZLWK FXUUHQW DSSOLFDWLRQ RI the name (Connolly 1939; Bruggen 1980; Herbert & Kilburn 2004). Tryon’s illustration (7U\RQ l88ƽ: 96, SO. l9, ¿J. 99), DV SRLQWHG RXW E\ %XUQXS (l92ƽ), HUURQHRXVO\ VKRZV WKUHH ODEUDO WHHWK, DQG VHHPV WR KDYH EHHQ LQÀXHQFHG E\ 3IHLIIHU¶V LQFOXVLRQ RI WKH 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 D VXSHU¿FLDO FROXPHOOD WRRWK. 7KH\ UHVHPEOH Gulella kosiensis (Melvill & Ponsonby, 1908) but are considerably larger (length 9.2mm) 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 QDPH ZLOO SUHVHUYH SUHYDLOLQJ XVDJH. $V WKH VLQJOH 601+ VSHFLPHQ UHÀHFWLQJ WKLV usage is in poor condition ( Fig. 2I ) we prefer (as permitted by Art. 75 of the Code) to select 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 (holotype Fig. 2L ), though generally less strongly sculptured, may also fall within this variable concept of G. menkeana and represent a mist-belt ecomorph, though we refrain 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 (Stanger)’ [later misspelt as ‘Strangier’ (Pfeiffer, 1859: 339)]. Subsequently, Connolly (1939) cited several additional localities on the KwaZulu-Natal south coast (Scottburgh, 3RUW 6KHSVWRQH, 3RUW (GZDUG).,Q UHDOLW\, WKHUH DUH WR GDWH QR FRQ¿UPHG UHFRUGV IRU WKLV 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 120km 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.

Figures

Fig. 2. (A, B) lectotype and paralectotype of Ennea adamsiana Pfeiffer, 1859 in 1854–1860, ‘Port Natal, M.C. [Mus. Cuming]’, length 8.45 and 8.1 mm (NHMUK 20110168); (C, D) syntypes of Ennea impervia 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).