European Journal of Taxonomy 37: 1-46

http://dx.doi.org/10.5852/ejt.2013.37

ISSN 2118-9773

WWW. europeanj oumaloftaxonomy. eu

2013 • Ben Rowson & Peter Tattersfield

This work is licensed under a Creative Commons Attribution 3.0 License.

Research article

um:lsid:zoobank.org:pub:ADF6394E-77B5-4309-87FE-4097FDE0A3FD

Revision of Dadagulella gen. nov., the “Cwfe/Za radius group”

(Gastropoda: Streptaxidae) of the eastern Afrotropics,

including six new species and three new subspecies

Ben ROWSON' & Peter TATTERSFIELD"

Biodiversity & Systematic Biology, National Museum of Wales, Cathay s Park, Cardiff, UK CFIO 3NP.

Email: [email protected]

^ um:lsid:zoobank.org:author:05D46259-DA45-45A4-9127-5AF361D566A7

" um:lsid:zoobank.org:author:8C55642E-B18A-461B-9BF9-31CB6DB7A070

Abstract. The genus Dadagulella gen. nov. is described to include 16 species of small, dentate, ovate-

acuminate Afrotropical snails. An identification key is provided and biogeography, anatomy and

systematics are discussed. The type species is the Kenyan D. radius (Preston, 1910) comb, nov., whose

name has informally been used for part of the group in the past. Substantial intraspecific variation occurs

in three species: D. radius itself, D. browni (van Bruggen, 1969) comb. nov. mdD. minuscula (Morelet,

1877) comb. nov. (= Ennea fischeriana Morelet, 1881) (non Gulella minuscula Emberton & Pearce,

2000) . We recognise subspecies within each of these: D.radius radius (Preston, 1910) comb, nov.,

D. r calva (Connolly, 1922) comb, et stat. nov., D. browni browni (van Bruggen, 1969) comb, nov.,

D. b. mafiensis subsp. nov., D. b. semulikiensis subsp. nov., D. minuscula minuscula (Morelet, 1877)

comb, nov., D. m. mahorana subsp. nov. Six new Tanzanian species are described: D. cresswelli sp. nov.,

D. delta sp. nov., D. ecclesiola sp. nov., D. frontier arum sp. nov., D. minareta sp. nov., andZJ. pembensis

sp. nov. The genus includes seven other previously described species: D. cuspidata (Verdcourt, 1962)

comb, nov.; D. rondoensis (Verdcourt, 1994) comb, nov.; D. conoidea (Verdcourt, 1996) comb, nov.; D.

selene (van Bruggen & Van Goethem, 1999) comb, nov.; fJ. meredithae (van Bruggen, 2000) comb, nov.;

D. nictitans (Rowson & Fange, 2007) comb, nov.; andD. delgada (Muratov, 2010) comb. nov.

Key words. Mollusca, Africa, Eastern Arc Mountains, Albertine Rift, biogeography.

Rowson B. & Tattersfield P. 2013. Revision of Dadagulella gen. nov., the ''Gulella radius group” (Gastropoda:

Streptaxidae) of the eastern Afrotropics, including six new species and three new subspecies. European Journal of

Taxonomy 3)1 \ 1-46. http://dx.doi.org/10.5852/eit.2013.37

Introduction

The Streptaxidae J. Gray, 1860, or “hunter snails” (Herbert & Kilbum 2004) are the most speciose

terrestrial mollusc family in the Afrotropics. One group of small, ribbed, ovate-acuminate streptaxids,

usually with a pointed apex and with characteristic apertural dentition, constitutes a distinctive part

of the eastern Afrotropical fauna. They are litter-dwellers in lowland and montane forest and other

well-vegetated habitats and can be locally common (e.g. van Bruggen & Appleton 1977; Rowson et al.

2010b). The species are among those currently included in Gulella F. Pfeiffer, 1856, an extremely large

European Journal of Taxonomy 37 : 1-46 ( 2013 )

genus much in need of revision. Gulella includes several subgenera, some monotypic, erected for the

conchologically more characteristic species. However, no such name is available for the group of taxa in

question. The group includes those species first referred to by van Bruggen (2000) as the ''Gulella radius

group”, the name being taken from the Kenyan Ennea radius Preston, 1910. The "Gulella radius group”

has never been revised, but has been touched upon by Verdcourt (1962, 1985, 1996), van Bruggen &

Meredith (1984) (who called it the "Gulella browni group”), Tattersfield (1998b), van Bruggen (2000),

Rowson (2007a), Rowson & Lange (2007), Rowson et al. (2010a, b) and Muratov (2010). These authors

appear to disagree on the composition of the group. For example, van Bruggen (1969, 2000) included

the Kenyan G. pretiosa (Preston, 1911), which other authors did not. Muratov (2010) did not mention G.

selene van Bruggen & Van Goethem, 1999, G. meredithae van Bruggen, 2000 or G. nictitans Rowson &

Lange, 2007, three species which were each considered to belong to the "Gulella radius group” by their

original authors. Preston (1910) noted the similarity between E. radius and Pupa minuscula Morelet,

1877 of the Comoros, a comparison not drawn by later authors. Verdcourt, meanwhile, did not associate

the above named East African species with one another in his checklists (1983, 2006). In Richardson’s

(1988) catalogue, the relevant species are treated in separate genera. There is thus a need to discuss

the evidence for the existence of such a group and to define it more precisely. We here delimit it on

conchological grounds and describe its key features, including available anatomical and molecular

data. All species previously assigned to the "Gulella radius group”, with the exception of G. pretiosa

(excluded because of sequence and anatomical reasons, see below), are included, as are several others.

A second difficulty is the apparent conchological variability of some species that have been attributed

to this group. For example, G. cuspidata Verdcourt, 1962 was distinctive enough to key out first in

Verdcourt’s (1962) keys to the then known East African Gulella species. However, the other relevant

species in the key, G. radius (Preston, 1910) and G. calva Connolly, 1922 key out in not two but

three other, separate places despite Verdcourt’s (1962) suggestion that one of them is “scarcely more

than a variety” of the other. Verdcourt later (1985) speculated whether the group consisted of only a

single variable species G. radius, yet maintained G. radius, G. calva and G. cuspidata as distinct in

his checklists (1983, 2006). He had almost certainly not seen material of most of the species we here

recognise as new. We consider most of these to be readily separable from G. radius using the key and

other information we provide. In other cases the separation is more difficult and some taxa could be

shown to be more closely related to one another as more material becomes available, although we have

studied material from many additional populations as well as the relevant types. This appears sufficient

to assess intrapopulation variation and to detect examples of species occurring in sympatry without

morphological intermediates, at least in Tanzania (two species occur at Pugu, two at Kwamgumi, and

three at Kimboza). We also emphasize that, although we intend our taxa to reffect monophyletic lineages,

this is a revisionary study and not a phylogenetic one. Further phylogenetic work is necessary to test

hypotheses on the age and relationships of the taxa.

Material and Methods

Type and other material was studied at or on loan from the following museums, or has been deposited

with them: National Museum of Wales, Cardiff, UK (NMW); The Natural History Museum, Eondon,

UK (NHMUK; was BMNH); Netherlands Centre for Biodiversity Naturalis, Eeiden, the Netherlands

(RMNH); Royal Museum for Central Africa, Tervuren, Belgium (MRAC); Royal Belgian Institute of

Natural Sciences, Brussels, Belgium (RBINS); Senckenberg Forschungsinstitut und Naturmuseum,

Frankfurt, Germany (SMF); National Museums of Kenya, Nairobi, Kenya (NMK); National Museum of

Tanzania, Dar es Salaam, Tanzania (NMT); and the KwaZulu-Natal Museum, Pietermaritzburg, South

Africa (NMSA). Intact adult shells were measured with Vernier callipers and photographed in apertural

view with further measurements made using the software ImageJ (Rasband 1997-2011). The number of

whorls was counted in this view to the nearest 0.5 whorl. The number of ribs per mm on the penultimate

ROWS ON B. & TATTERSFIELD R, Revision of Dadagulella gen. nov. (Streptaxidae)

whorl was used as a measure of seulpture beeause the front of the body whorl was often ealloused or

eroded smooth. A spire angle was measured as the angle enelosed by a pair of lines beginning at the ends

of the upper suture of the penultimate whorl and meeting at the shell apex.

A note on terminology is required. Our preferred term for the overall shell shape of the majority of

speeies, where the maximum width is reaehed at approximately the middle of the shell (e.g. all of Figs

6-26) is ‘ovate-aeuminate’ (after van Bruggen & Van Goethem 1999; van Bruggen 2000; Muratov 2010,

ete.). In nearly all speeies of the group the spire is substantially tapered towards the apex. The spire has

itself usually been referred to as ‘aeuminate’ (e.g. Preston 1910; Verdeourt 1962; van Bruggen & Van

Goethem 1997; Muratov 2010). The sides of the upper whorls of the spire appear eoneave in a number

of speeies (e.g. Figs 29-31) for whieh our preferred term is ‘eoeloeonoid’. Where the upper whorls of

the spire appear eonvex we use the term ‘eyrtoeonoid’. The apex of the shell, i.e. that part of the spire

formed by the embryonie whorls, is pointed in most speeies, sharply in some. In others it is rounded at

a maeroseopie seale (any point is of eourse aetually rounded at the most mieroseopie of seales). This

variation appears to be at least partly independent of the spire angle, e.g. Figs 33-34 have a similar spire

angle but very different apiees.

For dimensions, see Table 1.

Abbreviations

ad. = adult shell

juv. = juvenile shell

BR = B. Rowson

PT = P. Tattersfield

MBS= M.B. Seddon

CFN = C.F. Ngereza

Results

Order Stylommatophora Sehmidt, 1855

Superfamily Streptaxoidea J. Gray, 1860

Family Streptaxidae J. Gray, 1860

Genus Dadagulella gen. nov.

um:lsid:zoobank.org:aet:lC3959C3-50E5-4DFF-A5AC-67BB9F05A042

Figs 1-84; Table 1

Type species

Ennea radius Preston, 1910.

Included species

Sixteen. In addition, D. radius (Preston, 1910) eomb. nov. andD. minuscula (Morelet,I877) eomb. nov.

are treated as having two subspeeies and D. browni (van Bruggen, 1969) eomb. nov. as having three

(see below).

Diagnosis

Shell small (2.3-5.5 mm high), ovate to ovate-aeuminate with an aeuminate (sometimes eoeloeonoid

or eyrtoeonoid) spire; strongly and often sinuously radially ribbed, never smooth. Peristome refleeted.

Aperture dentate, with at least one parietal and one palatal tooth. Juvenile aperture (known from

7-8 speeies) dentate (exeept possibly in D. nictitans eomb. nov.), aperture not downtumed. Radula

Table 1. Measurements and dentition seores for adult shells of Dadagulella gen. nov. speeies and subspeeies. Ranges are given with the mean (for

measurements) or mode (for tooth eounts) in braekets when n (number of shells measured) was greater than 2. Seores and measurements mentioned in

the Table were taken either from all available adult material or from literature.

European Journal of Taxonomy 37 : 1-46 ( 2013 )

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ROWS ON B. & TATTERSFIELD R, Revision of Dadagulella gen. nov. (Streptaxidae)

(known from three speeies) bearing multieuspid teeth, the outer eusps mueh smaller than the inner.

Other anatomy (known from six speeies): salivary gland bilobed or Y-shaped. Vas deferens thiekened

or with divertieulum prior to insertion on penis, penis laeking sheath, with a museular apex in whieh is

embedded a spatulate seoop with a mieroseopieally serrated tip, usually assoeiated with one or two large

apieal hooks, and a few (<40) smaller hooks lower down. For dimensions, see Table 1.

Etymology

Prefix from Swahili noun ''dadd" meaning sister, with referenee to the apparent relationships and East

Afriean eentre of diversity of the group.

Description

Shell (Figs 1-64). Small to medium-sized relative to speeies of Gulella (2.3-5.5 mm high x 1.4-2.8 mm

wide), of 4.5-8 whorls. Ovate to ovate-aeuminate (maximum width being approximately at middle of

shell, usually at penultimate whorl) or subeonieal (maximum width being in bottom third of the shell,

at body whorl). Spire eharaeteristieally narrowly to broadly aeuminate, oeeasionally eoeloeonoid or

eyrtoeonoid (spire angle 43-77°), angle varying a little within most speeies. Apex (i.e. top of spire)

pointed in most speeies, but rounded in others. Sutures usually deep, shells never eompletely smooth¬

sided. Umbilieus elosed or narrowly open, this sometimes varying within a speeies. Embryonie whorls

usually smoothly granulate, with fine radial striae in two speeies, or irregularly punetate in another. The

last part of the embryonie shell is seulptured with very fine radial striae in most (possibly all) speeies but

these are worn away in all but the freshest individuals. Eater whorls never smooth, with radial ribs (5-27

per mm on penultimate whorl) running from suture to suture, often strong and/or sinuous, lamella-like

in some speeies. Peristome eomplete, or ineomplete parietally, always refleeted to some extent. Outer

palatal surfaee of aperture with a depression, sometimes furrow-like, eorresponding to the palatal tooth,

and sometimes another eorresponding to the basal tooth. Dentition 3-fold (rarely, and debatably, 2-fold)

to 8-fold. Dentition eonsisting of at least one parietal tooth and one palatal tooth, the latter often slab¬

like and/or bifld, sometimes forming a parieto-palatal sinus. Usually also with one deep-set eolumellar

baffle, folded or sub-bifld or sub-trifld in some speeies. Often with additional parietal, palatal, basal

and/or eolumellar teeth or dentieles. Dentition is variable in some speeies but the form, as well as the

number, of the teeth is often eonsistent enough to distinguish speeies. Juvenile shells (known from 7-8

speeies) always with 3-fold to 4-fold apertural dentition (Figs 43-50) exeept in one speeies, D. nictitans

eomb. nov., where the only known juvenile has no teeth. In several speeies, some or all of the teeth in

juvenile shells appear to be resorbed at intervals.

Cephalopodium. Pale eream or yellow, usually with aprieot-eoloured to orange tentaele retraetors and

somet im es with brown speekles on the mantle.

Pallial complex. Sigmurethrous, with long zone of eontaet between long, oblong kidney and hindgut,

pulmonary eavity not strongly vaseularised.

Salivary glands (Figs 69-72). United, soft, not tumid, elongate, bilobed to Y-shaped; eaeh duet leaving

at the apex of the lobe and evenly thiek throughout.

Buccal mass. Very small and elongate, radula eorrespondingly small, tightly enrolled and diffleult to

loeate or prepare.

Radula (Figs 65-68). (known from three speeies) With a unieuspid eentral tooth and 9-15 laterals

in eaeh half-row, diminishing gradually in size laterally. Most or aft laterals distinetively bieuspid or

multieuspid, with outer eusps smaller or mueh smaller than inner eusps. Teeth are somewhat delieate (or

flexible; Aiken 1981) and are short and flake-like at the ventral end of the radular ribbon.

European Journal of Taxonomy 37 : 1-46 ( 2013 )

Genitalia. (Figs 73-83) (known from six species) Hermaphroditic duct diverticulum (talon) short, large,

sausage-shaped, not convoluted. Bursa copulatrix attending albumen gland. Acini of prostate indistinct

to distinct. Vagina attenuate. Oviduct often containing a single large egg. Vas deferens either thickened

prior to insertion on penis, or appearing as such but actually with a diverticulum lying parallel to it. Vas

deferens entering penis subapically. Penial retractor muscle branching off columellar muscle, attaching

partly to vas deferens, thus sometimes bifid or nearly so. Penis elongate, tubular. Penial sheath absent,

but lower part sometimes surrounded by a thin sheath-like layer contiguous with walls of lower penis.

Interior of penis with weak radial pilasters and small rhombic pads, sometimes with a longitudinal

pilaster or short rounded lobe. Apical part of penis with a spatulate or broad “scoop”. One end with a

microscopically serrated tip, the other end deeply embedded in muscular apex of penis. Scoop usually

associated with one or two large, broad hooks, lying just behind or underneath it. Scoop and hook(s)

could conceivably act together as a grip or pincer. Elsewhere in penis, a longitudinal row or group of few

(<40) short, simple hooks mounted on rhombic pads. “Spermatophore” (see Discussion) detected in two

species: comma-shaped, lying with reservoir in vagina with longitudinally ribbed tail extending into mid

penis. End of tail probably originally weakly attached to wall of penis [as in one specimen of D. radius

radius (Preston, 1910) comb, nov.] but subsequently detached. Partially digested tail fragments present

in bursa of one specimen of D. pembensis sp. nov.

Known distribution

Eastern Kenya, eastern Tanzania, western Uganda, eastern DR Congo, Malawi, eastern Zambia,

Mozambique, north-eastern South Africa, Comoros archipelago (Fig. 84).

Gender

Feminine.

Comparison and remarks

See General Discussion.

Dadagulella radius radius (Preston, 1910) comb. nov.

Figs 1-5, 9-20, 43-45, 52-53, 65, 69, 73, 77-78, 83-84; Table 1

Ennea radius Preston, 1910: 529, pi. Vll, fig. 8.

Gulella radius - Verdcourt 1962: 13, 17, ? not 22 (see D. radius calva below). — van Bruggen 1969:

71. —Verdcourt 1983: 234; 1985: 119, fig. 17; 1996: 136. — Tattersfield 1998a: 83; 1998b: 37. —

Verdcourt 2000: 415. — van Bruggen 2000: 233. — Eange & Mwinzi 2003: 64-66. — Verdcourt 2006:

48. — Rowson 2007a: 441-442, fig. 56 (not fig. 54). — Rowson & Eange 2007: 31. — Muratov 2010:

277. —Rowson et al. 2010b: 28-29, in part. —Ndalila2011: 24.

Ennea radius - Richardson 1988: 26.

Type material examined

KENYA: lectotype (here designated) MRAC.117592: 1 ad., “Shimbi Hills, British East Africa”, standing

as 'Jadius'\ labelled “ex Putzeys-Musee, 1935”, “type” and apparently the shell figured in Preston

(1910: pi. Vll, fig. 8). Verdcourt (1985) wrote that a second shell from Shimbi in the NHMUK type

collection could not be located at that time. We too failed to trace this specimen. Van Bruggen (1969)

referred to two other NHMUK shells from Gazi as “paratypes”, presumably because they were labelled

as such. However, Gazi (4.42°S, 39.50°E) is around 30 km from the type locality of Shimbi (Shimba)

Hills (4.25°S, 39.38®E), and Preston (1910) distinguished between the two localities elsewhere in his

paper. Thus it cannot be assumed that Gazi material has type status.

ROWS ON B. & TATTERSFIELD R, Revision of Dadagulella gen. nov. (Streptaxidae)

Other material examined

KENYA: NMW. 1955.158.25050: 3 ads, 7 juvs, Gazi, standing as >adius^\ RMNH.MOE.273272: 1 ad.,

Gazi, standing as ''radius''. RMNH.MOE.273826: 1 ad., near Mombasa (approx. 4.04°S, 39.66°E),

amongst river debris, Sep. 1987. NMW.Z. 1990.067.00001: 1 ad., Arabuko-Sokoke Forest (3.32°S,

39.87°E), 20 km SW of Gedi, Brachystegia Benth. woodland on sandy soil with some leaf litter, leg. MBS,

12 Aug. 1989. NMW.Z.2012.042.00002: 2 ad., DianiBeaeh (4.27°S, 39.59°E), south ofMombasa, under

deep mixed broadleaf litter, 18 m alt., leg. P. E. Cresswell, 24 Apr. 1997. NHMUK. 1911.10.12.146-147:

2 ads, Gazi, standing as "radius".

TANZANIA: NMW.Z.2009.013.00226: 2 ads, Jozani Forest Reserve (6.23°S, 39.42°E), Unguja L,

Zanzibar, leg. BR, B.H. Warren, & CFN, 5 Feb. 2009. NMW.Z.2004.014.00023: 10 ads, 1 juv., several

Figs 1-5. Eeetotype of Ennea radius Preston, 1910, type speeies of Dadagulella gen. nov. 1. Apieal

view. 2. Apertural view. 3. East two whorls, from left side. 4. East two whorls, from right side.

5. Umbilieal view.

European Journal of Taxonomy 37 : 1-46 ( 2013 )

in poor condition, Jozani Forest Reserve, Unguja L, Zanzibar, leg. PT, MBS, & CFN, 11-12 Mar. 2000.

NMW.Z.2003.001.00006: 2 ads, Mkungwe Forest Reserve (6.90°S, 37.91°E), Ulugum Mts, Morogoro

Distriet, submontane forest, approx. 900 m alt., leg. BR & CFN, 7 Feb. 2003. NMW.Z.2003.001.00007:

1 ad., 1 juv., Kimboza Forest Reserve (7.01°S, 37.78°E), Ulugum Mts, Morogoro Distriet, lowland forest

on limestone, approx. 300 m alt., leg. BR, PT, MBS, & CFN, 5 Feb. 2003. NMW.Z.2007.012.00001: 1 ad.,

Pugu Hills Nature Reserve (6.89°S, 39.09°E), SE of Dar es Salaam, leg. CFN, Jan. 2007 (sequeneed by

Rowson etal. 2010a as ''Gulella radius"'). NMT.2000/0130: 6 ads, 6 juvs Mbudya 1. (6.66°S, 39.25°E),

Dar es Salaam, leg. CFN & NMT team, 14 May 1998. NMW.Z. 1995.016.00005: 3 ads, Kiono Forest

(6.15°S,38.59°E),NWofDares Salaam (site 111), leg. PT,5Mar. 1995. NMW.Z. 1995.016.00006: 1 ad.,

Pugu, S of Dar es Salaam (site 11), leg. PT, 6 Mar. 1995. NMW.Z. 1995.016.00007: 2 ads, Pugu Forest,

S of Dar es Salaam (site 1), leg. PT, 6 Mar. 1995. NMW.Z. 1995.016.00008: 1 ad., Amboni eaves and

Mkulumuzi Gorge (5.06°S, 39.05°E), near Tanga (site 1), leg. PT, 3 Mar. 1995. NMW.Z. 1995.016.00010:

3 ads, 2 juvs, Amboni, near Tanga (site 1), leg. PT, 3 Mar. 1995. NMW.Z. 1995.016.00009: 2 ads, Amboni,

near Tanga (site 111), leg. PT, 3 Mar. 1995. NMW.Z. 1995.016.00011: 6 ads, Amboni, near Tanga (site 11),

leg. PT, 3 Mar. 1995. NMW.Z.2003.074.00005: 1 ad., Kwamgumi Forest Reserve (4.94°S, 38.75°E),

East Usambara Mts, Muheza Distriet, lowland forest at approx. 350 m alt., leg. Frontier Tanzania, 4

Dee. 1996. NMW.Z. 2003.074.00006: 1 ad. in poor eondition, Mtai Forest Reserve (4.87°S, 38.77°E),

East Usambara Mts, Muheza Distriet (plot 73), riverine forest at 200 m alt., leg. Frontier Tanzania, 23

Jan. 1996.

Description

Shell (Figs 1-5, 9-20, 43-50, 52-53). Very variable in size, shape and dentition, small to large (2.80-

4.60 mm high x 1.60-2.44 mm wide), of 6.0-7.0 whorls. Ovate-aeuminate, spire narrowly to broadly

aeuminate (spire angle 55-70°). Apex pointed. Embryonie whorls smoothly granulate. Eater whorls

with relatively eoarse and few ribs (7-16 per mm on penultimate whorl). Sutures relatively deep.

Umbilieus elosed or narrowly open. Peristome eomplete or (more often) ineomplete parietally. Outer

palatal surfaee of aperture with a depression eorresponding to the palatal tooth and sometimes another

one eorresponding to the basal tooth. Dentition 5-fold to 7-fold, eonsisting of at least: one lamella-like

parietal tooth; one slab-like palatal tooth, often bifid; one basal dentiele; one shallow eolumellar dentiele

(very weak in some eases) and one deep-set eolumellar baffle, always visible. Parieto-palatal sinus, if

present, broad, seldom narrow or parallel-sided. Neither the eolumellar baffle nor basal dentiele were

mentioned in Preston’s (1910) original deseription, although both are elearly visible on the leetotype

whieh is apparently the originally figured speeimen. We assume this to have been a lapsus. One or two

additional eolumellar dentieles may be present (e.g. Figs 12, 14, 15). Juvenile shells (Figs 43-45) with

3-fold dentition: one parietal lamella; one bifid basal tooth; and one eolumellar tooth or thiekening.

Some earlier teeth are retained in some juveniles and even adults (e.g. some speeimens from Jozani).

Cephalopodium. Pale eream or yellow, with aprieot to orange tentaele retraetors and often with brown

speekles on the mantle.

Salivary glands (Fig. 69). United, soft, not tumid, elongate, bilobed to nearly Y-shaped; eaeh duet

leaving at the whitened apex of the lobe and evenly thiek throughout.

Radula (Fig. 65). With a unieuspid eentral tooth and around 13 laterals in eaeh half-row, diminishing

gradually in size laterally. All laterals bieuspid or trieuspid, with outer eusps mueh smaller than inner

eusps.

Genitalia (Figs 73, 77, 83). Vas deferens appearing thiekened prior to insertion on penis but aetually with

a short, broad parallel divertieulum. Penial sheath absent but with a thin sheath-like layer eontiguous

with walls of lower penis. Interior of penis with weak radial pilasters and small rhombie pads, sometimes

ROWS ON B. & TATTERSFIELD R, Revision of Dadagulella gen. nov. (Streptaxidae)

with a longitudinal pilaster or short, rounded and weakly ehitinized lobe. Apieal, museular part of penis

with a single large hook, assoeiated with a “seoop” with mieroseopieally serrated tip. Elsewhere in

penis, a single longitudinal row of short, simple hooks mounted on rhombie pads. “Spermatophore”

(see Diseussion) present in penis and vagina of one Amboni speeimen; “spermatophore” tail, apparently

attaehed to penis wall, present in penis of one Pugu speeimen.

Range and habitat

Eastern Kenya and eastern Tanzania, ineluding Unguja (Zanzibar) island. Apparently replaeed by other

Dadagulella gen. nov. speeies on Pemba island and in some lowland areas and montane forests, and by

D. browni eomb. nov. sd. in Tanzania from around 7.8°S southwards. Dadagulella r radius eomb. nov. is

sympatrie with other Dadagulella gen. nov. speeies at Kwamgumi, Pugu and Kimboza [it has also been

reeorded from Kimboza by Verdeourt (2006: 48)]. The reeords are mainly from forest and other well-

vegetated habitats; Verdeourt (2000) suggested the habitat was “woodland/foresf’. Eange & Mwinzi

(2003) found D. radius eomb. nov. aeross several forest types at Arabuko-Sokoke, but Ndalila (2011)

found only three speeimens in the Shimba Hills, all in serub and grassland rather than forest.

Remarks

This appears to be the most widespread and variable speeies of Dadagulella gen. nov. In this we eoneur

with van Bruggen (1969) and Verdeourt (1985) that D. radius eomb. nov. is a speeies variable enough

to inelude shells as different as the leetotype (Figs 1-5, 11) and those from Diani Beaeh (Fig. 14). We

retain one sueh extreme form, subsequently deseribed by Connolly (1922) as Gulella calva Connolly,

1922, as a subspeeies of D. radius eomb. nov. (see below) but we refrain from deseribing additional

subspeeies sinee these forms are not obviously geographieally isolated (e.g. at Amboni, Fig. 15) and

oeeur throughout the geographieal range of the speeies. The variation between them often appears

eontinuous rather than diserete (e.g. eompare Figs 17-20, eaeh from a similar latitude and arranged

in order from W to E). Furthermore, some variability is often present at a single loeality, e.g. at Gazi

(eompare Figs 12 and 13). Despite this, D. radius eomb. nov. sd. differs from other similar, lowland

speeies as follows. It differs from D. browni eomb. nov. sd. in never having either an additional parietal

dentiele or two basal dentieles, and in normally having a broader (or no) parieto-palatal sinus. It differs

from D. delgada (Muratov, 2010) eomb. nov. in laeking the flaring, lamella-like ribs and nearly always

having a more broadly aeuminate spire. It differs from D. ecclesiola sp. nov. in not having the basal

dentiele hidden behind the palatal tooth, in having at least one shallow eolumellar tooth, and in having a

broader (or no) parieto-palatal sinus. The distinetion between D. radius eomb. nov. sd. andD. minuscula

eomb. nov. of the Comoros, with whieh it was eompared by Preston (1910), is diseussed under the latter

speeies.

Dadagulella radius calva (Connolly, 1922) eomb. et stat. nov.

Figs 6-8, 51, 84; Table 1

Gulella calva Connolly, 1922: 495, pi. XIV, fig. 35.

Gulella calva-NQx&comi 1962: 17; 1983: 234. —Riehardson 1988: 62. —Verdeourt 2000: 215; 2006: 49.

? ''Gulella radius (Preston) var. (K, Mrima Hill Forest)” - Verdeourt 1962: 22.

Type material examined

KENYA: leetotype (here designated) NHMUK. 1937.12.30.486: 1 ad., Tam Desert (i.e. a semi-arid

area of southeastern Kenya now partly in the Tsavo East National Park, approx. 3.40°S, 39.00°E), leg.

Pereival, labelled “type”, and apparently the shell figured in Connolly (1922: pi. XIV, fig. 35).

European Journal of Taxonomy 37 : 1-46 ( 2013 )

Figs 6-26. Adult shells of Dadagule I la radius (Preston, 1910) eomb. nov., D. browni (vanBmggen, 1969)

eomb. nov. and their respeetive subspeeies. 6-20. D. radius eomb. nov. sd. 6-8. D. radius calva (Connolly,

1922): 6. Leetotype, Tam Desert. 7. Tam Desert near Malindi. 8. Near Mombasa. 9-20. D. radius radius'.

11. Leetotype, Shimba Hills. 12. Gazi. 13. also Gazi. 14. Diani Beaeh. 15. Am bon i. 16. Kimboza.

17. Mkungwe. 18. Pugu. 19. Mbudya 1. 20. Jozani. 21-26. D. browni eomb. nov. s.l.'. 21. D. browni

mafiensis subsp. nov., holotype, Mlula. 22. D. browni semulikiensis subsp. nov., holotype, Semuliki NP.

23-26. D. browni browni'. 23. Mwanihana. 24. Mzelezi. 25. Pomene Bay. 26. Holotype, Lake Sibayi

(after van Bmggen 1969).

ROWS ON B. & TATTERSFIELD R, Revision of Dadagulella gen. nov. (Streptaxidae)

Other material examined

KENYA: NMW. 1955.158.25052: 5 ads, “ea. lat. 3°5’, long. 39°27’”, standing as “ca/va Co.” and later

labelled “ef radius’" by B. Verdeourt. Assuming that the latitude refers to a point south rather than north

of the equator, the eoordinates eorrespond to a point in the Tam Desert , ±75 km inland of Malindi.

RMNH.MOE.288087: 1 ad., near Mombasa (approx. 4.04°S , 39.66°E), amongst river debris, Sep. 1987.

Description

Shell (Figs 6-8, 51). Earge (4.00-4.60 mm high x 2.10-2.50 mm wide), of 6.5-8.0 whorls. Ovate-

aeuminate, but more eolumnar than other Dadagulella gen. nov., spire narrowly to broadly aeuminate

(spire angle 49-62°). Apex pointed. Embryonie whorls smoothly granulate. Eater whorls with

eharaeteristieally eoarse, few and widely spaeed ribs (5-10 per mm on penultimate whorl). Sutures

deep. Umbilieus elosed or nearly so. Peristome ineomplete parietally. Outer palatal surfaee of aperture

with a depression eorresponding to the palatal tooth. Dentition eharaeteristieally simple, 4-fold to 5-fold,

eonsisting of one lamella-like parietal tooth; one slab-like palatal tooth without parieto-palatal sinus;

one basal dentiele; and one deep-set eolumellar baffle, always visible, sometimes with a very slight,

shallower eolumellar tooth. Juvenile shells and anatomy unknown.

Range and habitat

Eowlands of extreme southeastern Kenya, ineluding Tam Desert. This subspeeies has also been reeorded

from Malindi (Verdeourt 1962: 17) and Mrima Hill (4.48°S, 39.25°E) (Verdeourt 2006). Verdeourt

(2000) suggested the habitat was “bushland to forest”, in eontrast to “woodland/foresf ’ for D. r radius

eomb. nov. Shells of both D. r radius eomb. nov. and D. r calva eomb. et stat. nov. have been found in

river debris near Mombasa, but may have been washed in from different loealities.

Remarks

The leetotype and similar material of this subspeeies (Figs 6-8, 51) differ from D. r radius eomb. nov.

in the more widely spaeed ribs, more eolumnar shape and simpler dentition, although some speeimens

seem to show intermediate eharaeters. Verdeourt evidently had diffleulty separating them. Although

Verdeourt (1962: 17) suggested G. calva was “seareely more than a variety of G. radius"", it keyed out

in a separate part of his key, with radius appearing in two other plaees (1962: 13, 22). The seeond of

these (p.22) was listed as ''Gulella radius (Preston) var. (K [Kenya], Mrima Hill Forest)”. Verdeourt’s

measurements, deseription, and range of apertural tooth formulas allow attribution of this taxon to D. r

calva eomb. et stat. nov.. In his later eheeklists, Verdeourt (1983: 234; 2006: 48-49) maintained the two

as separate speeies, and gave Mrima Hill as a loeality for calva but not radius. He also kept them apart

in his list of eoastal molluses (2000). Thus it appears likely that he later deeided that (1962: 22) ''radius

var.” belonged to what we treat as D. r calva eomb. et stat. nov. and not to what we treat as D. r radius

eomb. nov. It also appears that he deeided not to separate the two taxa. We too found this a diffleult

deeision to take, owing to the morphologieal, habitat and distributional differenees, whieh may of eourse

be interrelated. To aeknowledge this diffleulty, and the potential diffleulties in assigning future material

to either, we rank calva as a subspeeies of radius.

Dadagulella browni browni (van Bruggen, 1969) eomb. nov.

Figs 23-26, 47-48, 71, 74, 82, 84; Table 1

Gulella browni van Bruggen, 1969: 69-71, figs 25-26.

Gulella - van Bruggen & Appleton 1977: 31-32. —Aiken 1981: 321-323, fig. 5. —Riehardson

1988: 60. — Verdeourt 1990: 349. — van Bruggen & Van Goethem 1999: 40. — van Bruggen 2000:

233-234. —Herbert & Kilbum 2004: 212. —Rowson & Eange 2007: 31. —Muratov 2010: 277.

European Journal of Taxonomy 37 : 1-46 ( 2013 )

“a closely related speeies from Zululand” - Verdeourt 1985: 119.

Gulella ef radius - Tattersfield et al. 2006: 52, 54, 58.

Type material examined

SOUTH AFRICA: holotype RMNH.MOL.54894: 1 ad.. Lake Sibayi (approx. 27.34°S, 32.71°E),

KwaZulu-Natal, dune forest on the east shore, leg. D.S. Brown, 12 Jun. 1966 (examined digital

photograph only).

Other material examined

TANZANIA: NMW.Z.2003.001.00008: 6 ads, 5 juvs, Mt. Mwanihana Forest Reserve (7.82°S,

36.83°E), Udzungwa Mts National Park, Kilombero Distriet, lowland forest at approx. 600-900 m

alt., leg. BR, PT, MBS & CFN, 29 Jan. 2003. NMW.Z.2003.001.00009: 4 ads, data as previous but

700 m alt. NMW.Z. 1997.007.00003: 5 ads, 2 juvs, Mzelezi Forest Reserve (8.79°S, 36.72°E), Mahenge

Mts, Ulanga Distriet, forest on limestone, approx. 645 m alt., leg. PT, MBS & CFN, 6 Feb. 1997.

NMW.Z. 1997.007.00004: 1 ad., data as previous. NMW.Z. 1997.007.00005: 2 ads, 6 juvs, data as

previous.

MOZAMBIQUE: NMW.Z.2012.040.00001: 2 ads, Pomene Bay, Inhambane Provinee, near lighthouse

S of old hotel (22.9417°S, 35.5913°E), degraded dune forest, in leaf-litter, leg. D. Herbert, Oet. 2002 (ex

NMSA.E5797). NMSA.E5797: 2 ads, data as previous.

SOUTH AFRICA: NMW.Z. 2012.040.00002: 3 ads, Kosi Bay mouth, KwaZulu-Natal (26.89°S,

32.88°E), dune forest, in sandy leaf-litter, leg. D. Herbert & E. Davis, “11-41”, 13 Oet. 2011 (exNMSA.

W8517). NHMUK.20110173: 2 ads, Eake Sibaya area of Tongaland (approx. 27.35°S, 32.67°E),

Zululand, S. Afriea, leg. C.C. Appleton, Dee. 1972-Nov. 1973.

Description

Shell (Figs 23-26, 47-48). Small (2.60-3.20 mm high x 1.60-1.90 mm wide), of 5.5-6.0 whorls. Ovate-

aeuminate, spire broadly aeuminate (spire angle 66-76°). Apex pointed. Embryonie whorls smoothly

granulate. Eater whorls with relatively eoarse ribs (11-15 per mm on penultimate whorl). Sutures

relatively shallow. Umbilieus elosed or narrowly open. Peristome ineomplete parietally. Outer palatal

surfaee of aperture with a depression eorresponding to the palatal tooth and another mueh weaker one

eorresponding to the basal tooth. Dentition 7-fold to 8-fold, eonsisting of one lamella-like parietal

tooth and (almost always) one additional parietal dentiele; one slab-like palatal tooth, forming a narrow

parieto-palatal sinus; two basal dentieles; a deep-set eolumellar baffle, sometimes folded, and one or

two shallower eolumellar dentieles. Dentition of 8-fold individuals alternatively reeognisable as 7-fold,

depending on the interpretation of the eolumellar and parietal dentieles (van Bruggen 1969). Juvenile

shells of D. browni eomb. nov. were not known to van Bruggen (1969, 2000). Herbert & Kilbum (2004)

deseribe them simply as dentate, but with dentition different from that of the adult. Juveniles from

southern Tanzania (Figs 47-48) have dentition similar to, but weaker than, juveniles of D. r radius

eomb. nov. (Figs 43-45).

Cephalopodium. Pale eream, with pink-orange tentaele retraetors and a few purple-brown speekles in

the mantle.

Salivary glands (Fig. 71). United, soft, not tumid, elongate, bilobed; eaeh duet leaving at the whitened

apex of the lobe and evenly thiek throughout.

Radula. Deseribed and figured by Aiken (1981) as having a unieuspid, ‘heart-shaped’ eentral tooth and

nine laterals in eaeh half-row, of whieh at least the inner six are trieuspid, diminishing gradually in size

laterally. The outer eusps of most teeth are smaller than the inner eusp. Aeeording to Aiken (1981) the

teeth are fiexible when pressed by a eover slip and there are 47 rows.

ROWS ON B. & TATTERSFIELD R, Revision of Dadagulella gen. nov. (Streptaxidae)

Genitalia (Figs 74, 82). Upper parts of spermoviduet not reeovered. Vas deferens appearing thiekened

prior to insertion on penis but aetually with a very short, broad parallel divertieulum. Penial sheath

absent. Interior of penis with weak radial pilasters. Apieal, museular part of penis with a single large

hook, assoeiated with a spatulate “seoop” with mieroseopieally serrated tip. Elsewhere in penis, a single

longitudinal row of short, simple hooks.

Range and habitat

Coast of north-eastern South Afriea: dune forest, eoastal forest and bush; also eentral Mozambique: high

forest in Chiluvo [Xiluvo] hills (19.24°S, 34.06°E) at 700 ft (approx. 213 m) elevation (van Bruggen

1969; van Bruggen & Appleton 1977; Herbert & Kilbum 2004). Also in lowland forest, up to 900 m, in

southern Tanzania, as far north as 7.8°S.

Remarks

We agree with van Bruggen (1969) that the eomplex dentition of D. browni eomb. nov. s.l. separates

it from D. radius eomb. nov. sd., whieh never seems to have 8-fold dentition, in partieular the extra

parietal and basal dentieles. The shells of D. browni eomb. nov. sd. are also mueh smaller, less elongate

(i.e. have a lower height/width ratio), and have a mueh narrower and more horizontal parieto-palatal

sinus than is usual for D. radius eomb. nov. sd. Although some speeimens of D. r. radius eomb. nov.

approaeh D. browni eomb. nov. s.l. in one or more of these respeets (e.g. Figs 10, 14, 17) all ean be

readily assigned to one or the other and the type speeimens are morphologieally quite different (Figs 1-5,

11, 52 vs. Fig. 26). Material intermediate between D. browni eomb. nov. and D. radius eomb. nov.

might be expeeted in the intervening area of southern Tanzania or northern Mozambique. However, all

the relevant southern Tanzanian material here (e.g. from Mafia and Mahenge) is eloser to D. browni

eomb. nov. s.l. in dentition and other aspeets. Areeent survey in northern Mozambique (Muratov 2010)

eneountered only D. delgada eomb. nov.

Dadagulella browni mafiensis subsp. nov.

Figs 21, 54, 84; Table 1

Etymology

From Mafia Island, with the final ‘a’ elided for euphony.

Type material examined

NHMUK.20110174: 1 ad., Mlula, Mafia I. (approx. 7.87°S, 39.76°E), evergreen eoastal thieket on

elay loam eoral rag, leg. Frontier Tanzania, Oet.-Nov. 1990, labelled ''Gulella radius aggreg.” by B.

Verdeourt.

Other material examined

None.

Description

Shell (Figs 21, 54). Small (3.30 mm high x 1.80 mm wide), of 6.0 whorls. Ovate-aeuminate, spire

broadly aeuminate (spire angle 65°). Apex eonieal. Embryonie whorls smoothly granulate. Eater whorls

with relatively eoarse ribs (12 per mm on penultimate whorl). Sutures relatively shallow. Umbilieus

narrowly open. Peristome ineomplete parietally. Outer palatal surfaee of aperture with a depression

eorresponding to the palatal tooth. Dentition 5-fold, eonsisting of one lamella-like parietal tooth and

one additional parietal dentiele; one slab-like palatal tooth, forming a narrow parieto-palatal sinus; one

basal dentiele; and a deep-set, folded eolumellar baffle, but no shallower eolumellar dentieles. Shells

and anatomy of juveniles unknown.

European Journal of Taxonomy 37 : 1-46 ( 2013 )

Range and habitat

Evergreen eoastal thieket at the type loeality.

Remarks

This taxon is in some respeets intermediate between the type speeimens of D. r radius eomb. nov. and

D. b. browni eomb. nov., and thus distinet from either, yet difheult to plaee. It is larger and not as squat

as other D. browni eomb. nov. s.l, and laeks the additional basal dentiele and shallower eolumellar

dentition. Conversely, it has a parietal dentiele not seen in D. radius eomb. nov. s.l, is smaller and

squatter than most D. radius eomb. nov. sA., and has a longer and narrower parieto-palatal sinus than

any D. radius eomb. nov. sA. Mafia Island, from whieh no other Dadagulella gen. nov. are yet known,

lies at almost the same latitude (7.8°S) as the apparent northernmost limit of D. b. browni eomb. nov.

in mainland Tanzania. Given the morphology of this speeimen, its latitude and its isolation as an island

population, we treat it as a subspeeies of D. browni eomb. nov.

Dadagulella browni semulikiensis subsp. nov

Figs 22, 55, 84; Table 1

Etymology

From Semuliki.

Type material examined

UGANDA: holotype NMW.Z. 1997.009.00004: 1 ad., Semuliki National Park (00.82°N, 30.16°E),

Bwamba County, Bundibugyo Distriet, lowland Guineo-Congolian rainforest (site HR), approx. 760 m

alt., leg. PT & J.A. Allen, 14 Jul. 1996. Paratype NMW.Z. 1997.009.00005: 1 ad., data as holotype.

Other material examined

UGANDA: NMW.Z. 1997.009.00006: 1 juv., data as holotype.

Description

Shell (Figs 22, 55). Small (3.20-3.30 mm high x 1.90 mm wide), of 6.0-6. 5 whorls. Ovate-aeuminate,

spire broadly aeuminate (spire angle around 67°). Apex pointed. Embryonie whorls smoothly granulate.

Eater whorls with relatively eoarse ribs (around 12 per mm on penultimate whorl). Sutures relatively

shallow. Umbilieus elosed. Peristome eomplete. Outer palatal surfaee of aperture with a depression

eorresponding to the palatal tooth, and another mueh weaker one eorresponding to the basal tooth.

Dentition 8-fold, eonsisting of one lamella-like parietal tooth and one additional parietal dentiele;

one slab-like palatal tooth, forming a long, narrow parieto-palatal sinus; two basal dentieles; a deep-

set eolumellar baffle and two shallower eolumellar dentieles. Juvenile shell with 3-fold dentition: one

parietal lamella; one bifid basal tooth; and one baso-eolumellar tooth. Anatomy unknown.

Range and habitat

Eowland Guineo-Congolian forest at the type loeality in the valley of the Albertine Rift.

Remarks

This subspeeies is very similar to D. b. browni eomb. nov. and, if found together, the two might not

at first glanee be distinguished. However, D. b. semulikiensis subsp. nov. is approximately 10% taller

and wider, has shallower sutures and a more eomplete peristome. Its parieto-palatal sinus is longer and

narrower than that of D. b. browni eomb. nov., and does not widen appreeiably towards its inner end.

The type loealities of D. b. browni eomb. nov. and D. b. semulikiensis subsp. nov. are over 3000 km

ROWS ON B. & TATTERSFIELD R, Revision of Dadagulella gen. nov. (Streptaxidae)

apart, in very different biogeographie and elimatie regions. Although several snail speeies are known

to range between South and East Afriea, most are either speeies of the eoastal strip, or are widely

distributed throughout the entire area. Dadagulella gen. nov. has not been reeorded from mueh of the

intervening area of East Afriea and is represented in the Albertine Rift otherwise only by D. selene

(van Bruggen & Van Goethem, 1999) eomb. nov. It remains possible that the Semuliki population has

deseended from D. b. browni eomb. nov., introdueed by man from Tanzania or further to the south, and

that the morphologieal differenees are mainly eeophenotypie. However, in the light of these differenees

and the great geographie separation, we treat it as a subspeeies of D. browni eomb. nov. as we do with

D. b. mafiensis subsp. nov.

Dadagulella delgada (Muratov, 2010) eomb. nov.

Figs 27, 84; Table 1

Gulella delgada Muratov, 2010: 274, 276-277, figs 37, 39-45.

Type material examined

None.

Type locality

“Mozambique: Cabo Delgado: 1.1 km WNW of lighthouse, 19 km NE of Palma, 10.68883°S,

40.62806°E, alt. 11 m, 24 Nov. 2009, TV. Muratov.” (Muratov 2010).

Other material examined

None.

Description

Shell (Fig. 27). Medium-sized (3.80-4.00 mm high x 1.80-1.80 mm wide), of 6.5-7.5 whorls. Elongate

ovate-aeuminate, spire narrowly aeuminate (spire angle 43-49°). Apex sharply pointed (with axis slightly

deviated in one syntype). Embryonie whorls smooth. Eater whorls with very few, widely-spaeed, fiaring,

subtriangular, lamella-like ribs (about 7 per mm on penultimate whorl). Sutures very deep. Umbilieus

narrowly open. Peristome ineomplete parietally. Outer palatal surfaee of aperture with a depression

eorresponding to the palatal tooth. Dentition 5-fold, eonsisting of one lamella-like parietal tooth; one

slab-like palatal tooth, forming a parieto-palatal sinus; one basal dentiele; a deep-set eolumellar baffle

and one shallower eolumellar dentiele. Juvenile shells with 3-fold dentition: one parietal lamella; one

basal tooth and one eolumellar thiekening. Muratov (2010) found no internal dentition in the upper

whorls. Anatomy unknown.

Range and habitat

At the type loeality, vegetation on eoral rag (Muratov 2010: 284).

Remarks

The few, fiaring, lamella-like ribs and elongate, narrowly aeuminate spire of D. delgada eomb. nov.

allow it to be separated from other speeies, ineluding some D. radius eomb. nov. sA. whieh it resembles

in size and dentition.

European Journal of Taxonomy 37 : 1-46 ( 2013 )

Dadagulella ecclesiola sp. nov.

urn:lsid:zoobank.org:act:CCE56537-B647-4144-AB07-FE322A139087

Figs 28, 49, 56, 84; Table 1

Etymology

From Latin 'ecclesiola\ feminine, diminutive of‘ehureh’ (or ‘the ehureh’); used arbitrarily to distinguish

the speeies from D. minareta sp. nov. with whieh it oeeurs.

Type material examined

TANZANIA: holotype NMW.Z.2003.001.00015: 1 ad., Kimboza Forest Reserve (7.01°S, 37.78°E),

Uluguru Mts, Morogoro Distriet, lowland forest on limestone, approx. 300 m alt., leg. BR, PT, MBS, &

CFN, 5 Feb. 2003. Paratypes NMW.Z.2003.001.00016: 8 ads, data as holotype. Paratype MRAC.

MT.803794: 1 ad., data as holotype. Paratype NHMIJK.20120259: 1 ad., data as holotype. Paratype

NMK: 1 ad., data as holotype. Paratype NMT: 1 ad., data as holotype. Paratype NMSA.L8692/T3061:

1 ad., data as holotype. Paratype RMNH.MOL.288089: 1 ad., data as holotype.

Other material examined

TANZANIA: NMW.Z.2003.001.00017: 1 ad., data as holotype, sequeneed by Rowson et al. (2010a)

as ''Gulella ef browni Uluguru”. NMW.Z.2003.001.00018: 3 ads in poor eondition, 3 juvs, data as

holotype.

Description

Shell (Figs 28, 49, 56). Small to medium-sized (3.20-3.70 mm high x 1.80-1.90 mm wide), of 5.5-7.0

whorls. Ovate-aeuminate, spire narrowly aeeuminate (spire angle 53-65°). Apex pointed. Embryonie

whorls smoothly granulate. Later whorls with relatively eoarse, often sinuous ribs (8-14 per mm on

penultimate whorl). Sutures deep. Umbilieus elosed. Peristome eomplete. Outer palatal surfaee of

aperture with a depression, often furrow-like, eorresponding to the palatal tooth. Dentition 4-fold,

eonsisting of one strongly oblique parietal tooth, V-shaped when shell turned to the left; one large slab¬

like palatal tooth, forming a narrow, horizontal parieto-palatal sinus; and a deep-set eolumellar baffle.

A basal dentiele is also present, presumably in all speeimens, but is partly or eompletely hidden by the

palatal tooth whieh oeeludes mueh of the aperture. The dentiele is visible when the shell is turned to the

right (Fig. 56). Juvenile shells not known with eertainty: an individual from Kimboza (Fig. 49), with

dentition like that of a juvenile D. r radius eomb. nov., might belong to this speeies. Anatomy unknown.

Range and habitat

In forest at the type loeality in the eastern Tanzanian lowlands.

Remarks

This speeies has mueh simpler dentition than D. minareta sp. nov. (whieh also oeeurs at Kimboza), and

is usually smaller, with straight rather than sinuous ribs. Its dentition is more like that of D. r radius

eomb. nov. (whieh again also oeeurs at Kimboza; Fig. 16) andD. minuscula minuscula (Morelet, 1877)

eomb. nov. but D. ecclesiola sp. nov. laeks the shallow eolumellar tooth, has a hidden or partly hidden

basal dentiele, and has a mueh narrower parieto-palatal sinus. It further differs from D. minuscula

minuscula eomb. nov. in having stronger ribs.

ROWS ON B. & TATTERSFIELD R, Revision of Dadagulella gen. nov. (Streptaxidae)

Figs 27-35. Adult shells of Dadagulella gen. nov. speeies. 27. D. delgada (Muratov, 2010) eomb. nov.,

syntype, Cabo Delgado (after Muratov 2010). 28. D. ecclesiola sp. nov., holotype, Kimboza. 29. D.

cresswelli sp. nov., holotype, Ngorongoro. 30. D. minareta sp. nov., holotype, Kimboza. 31. D. cuspidata

(Verdeourt, 1962) eomb. nov., holotype, Shume. 32. D. pembensis sp. nov., holotype, Ngezi. 33. D.

nictitans (Rowson & Eange, 2007) eomb. nov., holotype, Maeha. 34. D. frontier arum sp. nov., holotype,

Mtai. 35. D. delta sp. nov., holotype, Mwanihana.

European Journal of Taxonomy 37 : 1-46 ( 2013 )

Dadagulella cresswelli sp. nov.

um:lsid:zoobank.org:act:B3362D6F-2118-4DlD-8B25-Q98E25BQC3B2

Figs 29, 57, 84; Table 1

Etymology

After Pete Cresswell, who eolleeted the speeimen.

Type material examined

TANZANIA: holotype NMW.Z.2012.042.00001: 1 ad., Ngorongoro Crater, Arusha Region, erater rim

on southeastern side, heavy rainforest leaf litter, leg. P.L. Cresswell, 2 Jun.1996.

Other material examined

None.

Description

Shell (Figs 29, 57). Medium-sized (3.70 mm high x 1.80 mm wide), of 7.0 whorls. Ovate-aeuminate,

spire eoeloeonoid (spire angle 52°). Apex sharply pointed. Embryonie whorls smoothly granulate. Eater

whorls with relatively fine ribs (13 per mm on penultimate whorl). Sutures shallow. Umbilieus narrowly

open. Peristome eomplete. Outer palatal surfaee of aperture with a very deep, long, furrow-like depression

eorresponding to the palatal tooth. Dentition 5-fold, eonsisting of one V-shaped parietal tooth; one

bifid slab-like palatal tooth, forming a elear parieto-palatal sinus, with the upper eusp projeeting into

the sinus; a deep-set eolumellar baffie and two shallower eolumellar dentieles. Shells and anatomy of

juveniles unknown.

Range and habitat

Forest at the type loeality in northern Tanzania. The vegetation is presumably of a montane type, sinee

the erater fioor is above 1700 m while the rim rises to over 2400 m or higher.

Remarks

This speeies is distinetive in its deep, long furrow on the outer palatal surfaee in eombination with the

eoeloeonoid spire and dentition. D. minerata sp. nov. shares these features, but differs in having weaker

ribs and less eomplex dentition. It is the only Dadagulella gen. nov. speeies thus far eolleeted in the

voleanie (as opposed to bloek-faulted) highlands of Tanzania or Kenya.

Dadagulella minareta sp. nov.

um:lsid:zoobank.org:aet:6AFlE4AF-DlD2-4C5A-9FCA-AC5B15A07204

Figs 30, 50, 58, 84; Table 1

Etymology

From English ‘minaret’, the tower of a mosque; Eatinized by the addition of the feminine ending ‘-a’;

used as a noun in apposition.

Type material examined

TANZANIA: holotype NMW.Z.2003.001.00010: 1 ad., Kimboza Forest Reserve (7.01°S, 37.78°E),

Uluguru Mts, Morogoro Distriet, lowland forest on limestone, approx. 300 m alt., leg. BR, PT,

MBS & CFN, 5 Feb. 2003. Paratype NMW.Z.2003.001.00011: 1 ad., data as holotype. Paratypes

NMW.Z.2003.001.00012: 3 ads, data as holotype. Paratype NMW.Z.2003.001.00013: 1 ad., data as

holotype. Paratype MRAC.MT.803795: 1 ad., data as holotype. Paratype NHMUK.20120260: 1 ad..

ROWS ON B. & TATTERSFIELD R, Revision of Dadagulella gen. nov. (Streptaxidae)

data as holotype. Paratype NMK: 1 ad., data as holotype. Paratypes NMT: 2 ads, data as holotype.

Paratype NMSA.E863/T3062: 1 ad., data as holotype. Paratype RMNH.288088: 1 ad., data as holotype.

Other material examined

TANZANIA: NMW.Z.2003.001.00014: 1 juv., data as holotype. NMW.Z.2004.016.00008: 2 ads, Kanga

Forest Reserve (6.01°S, 37.72°E), Ngum Mts, Mvomero Distriet, lowland forest at approx. 450 m alt.,

leg. BR, PT & CFN, 26 Jun. 2004. NMW.Z. 1995.016.00012: 4 ads, 1 juv. PuguForest (6.89°S, 39.09°E),

S of Dar es Salaam (site I), leg. PT, 6 Mar. 1995. The juv. in the Pugu lot may not be eonspeeifie.

Description

Shell (Figs 30, 50, 58). Medium-sized (3.40-4.00 mm high x 1.80-2.00 mm wide), of 6.0-7.0 whorls.

Ovate-aeuminate, spire aeuminate to eoeloeonoid (spire angle 48-63°). Apex sharply pointed. Embryonie

whorls with fine regular radial striae; smoothly granulate where worn. Eater whorls with relatively

eoarse, usually sinuous ribs (10-13 per mm on penultimate whorl). Sutures deep. Umbilieus narrowly

open. Peristome eomplete or ineomplete parietally. Outer palatal surfaee of aperture with a furrow-like

depression eorresponding to the palatal tooth. Dentition 5-fold to 7-fold, eonsisting of one V-shaped

parietal tooth; one bifid, slab-like, deeply in-running palatal tooth, not usually forming a parieto-palatal

sinus; and two eolumellar teeth. The upper of these is large and eharaeteristieally squarish, running in

to form a eolumellar baffle whieh is eonneeted to it; the lower eolumellar tooth is a small in-running

dentiele. Additional teeth inelude an extra parietal tooth and/or one or two shallow eolumellar dentieles.

Juvenile shell (Fig. 50) with 3-fold to 4-fold dentition: one very short parietal lamella; two basal teeth

(or one bifid tooth); and one eolumellar tooth. Earlier basal and eolumellar teeth are retained.

Cephalopodium. Pale yellow, with aprieot tentaele retraetors.

Salivary glands. United, soft, not tumid, elongate, Y-shaped; eaeh duet leaving at the apex of the lobe

and evenly thiek throughout.

Radula. Not sueeessfiilly prepared.

Genitalia. Vas deferens thiekened prior to insertion on penis but apparently without divertieulum. Penial

sheath absent but with a thin sheath-like layer eontiguous with wall of lower penis. Interior of penis

with weak radial pilasters and small rhombie pads. Apieal part of penis with a broad “seoop” with

mieroseopieally serrated tip, but without large hook. Elsewhere in penis a single longitudinal row of

short, simple hooks mounted on rhombie pads.

Range and habitat

In forest at the type loeality and in Pugu Hills Nature Reserve, both in the eastern Tanzanian lowlands,

and in lowland forest on Mt. Kanga in the Ngum Mts.

Remarks

The dentition, most notieeably the squared-off shallow eolumellar tooth that mns in to eonneet with the

baffle, readily distinguishes this speeies from D. ecclesiola sp. nov. (whieh also oeeurs at Kimboza) and

D. r radius eomb. nov. (whieh also oeeurs at both Kimboza and Pugu). It also distinguishes it from D.

cuspidata (Verdeourt, 1962) eomb. nov., with whieh it shares the strong upper eolumellar tooth and fine

regular radial striae on the embryonie whorls. It differs from D. cresswelli sp. nov., with whieh it shares

the fiirrow-like depression on the outer palatal surfaee, in dentition and in having stronger ribs.

European Journal of Taxonomy 37 : 1-46 ( 2013 )

Dadagulella cuspidata (Verdcourt, 1962) comb. nov.

Figs 31, 59, 84; Table 1

Gulella cuspidata Verdcourt, 1962: 3, 27-28, pi. 3, fig. 2.

"'Gulella sp. Verdcourt 1958: 94, 100, fig. 10.

Gulella cuspidata - van Bruggen 1969: 71. — Verdcourt 1983: 232. — Richardson 1988: 72. —

Verdcourt 1996: 136. — Tattersfield 1998b: 37. — Tattersfield et al. 1998: 135. — van Bruggen 2000:

233. — Verdcourt 2006: 46. — Rowson & Lange 2007: 31. — Muratov 2010: 277.

Type material examined

TANZANIA: holotype MRAC.792352: 1 ad.. Worlds View, Shume, West Usambara Mts (4.70°S,

38.20°E), in rather dry evergreen forest, leg. B. & L. Verdcourt, Dec. 1956.

Other material examined

None.

Description

Shell (Figs 31,59). Large (4.30-4.80 mm high x 1.80 x 2.50 mm wide), of 7.75 whorls. Ovate-acuminate,

spire coeloconoid (spire angle 58°). Apex sharply pointed. Embryonic whorls appear smooth but with

fine regular radial striae. Eater whorls with relatively few, coarse ribs (about 8 per mm on penultimate

whorl). Sutures of intermediate depth. Umbilicus narrowly open. Peristome incomplete parietally.

Outer palatal surface of aperture with a furrow-like depression corresponding to the upper palatal tooth.

Dentition 6-fold (alternatively recognisable as 5-fold), consisting of one lamella-like parietal tooth;

two palatal teeth, the upper much larger and forming a parieto-palatal sinus; one basal, in-running tooth;

and a strong, squarish columellar tooth, running in to form a columellar baffle which is connected to it.

Juvenile shells and anatomy unknown.

Range and habitat

Elevation not stated but probably between 1500-2000 m at the type locality in northeastern Tanzania.

Remarks

This species differs from D. minareta sp. nov. in having two palatal teeth, in having a basal tooth, in the

shape of the parietal tooth, and in size.

Dadagulellapembensis sp. nov.

um:lsid:zoobank.org:act:4442210F-A097-400D-B34F-96A4B7677CE7

Figs 32, 46, 60, 66-68, 70, 75, 79-80, 84; Table 1

'“'Gulella” radius (Preston, 1910)” - Rowson etal. 2010b: 9-10, 28-29, figs 60-61.

Etymology

From Pemba Island, with the final ‘a’ elided for euphony.

Type material examined

TANZANIA: holotype NMW.Z.2009.013.00227: 1 ad., Ngezi Forest Reserve (4.94°S, 39.69°E), Pemba

L, Zanzibar, dry forest on dark, sandy soil on coral rag on Tondooni peninsula within reserve, leg. BR, B. H.

Warren & CFN, 7 Feb. 2009; fig. 60 in Rowson a/. (2010b). ParatypesNMW.Z.2009.013.00228: 2 ads,

data as holotype. Paratypes MRAC.MT. 803796-7:2 ads, data as holotype. Paratypes NHMUK.20120261:

ROWS ON B. & TATTERSFIELD R, Revision of Dadagulella gen. nov. (Streptaxidae)

2 ads, data as holotype. Paratypes NMK: 2 ads, data as holotype. Paratypes NMT: 2 ads, data as holotype.

Paratypes NMSA.E8694/T3063: 2 ads, data as holotype. Paratypes RMNH.MOE.288090: 2 ads, data

as holotype.

Other material examined

TANZANIA: NMW.Z.2009.013.00229: 15 juvs, data as holotype; one figured as fig. 61 in Rowson

et al. (2010b). NMW.Z.2009.013.00230: 3 ads, near Chwaka (5.39°S, 39.77°E), Pemba L, Zanzibar,

elove and fruit tree woodland on dark, sandy soil, leg. BR, B.H. Warren & CFN, 15 Feb. 2009.

NMW.Z.2009.013.00231: 3 ads, Msitu Mkuu Forest Reserve (5.00°S, 39.83°E), Pemba L, Zanzibar,

moist forest on dark (not sandy) soil on eoral rag in high forest in north of reserve, leg. BR, B.H. Warren &

CFN, 10 Feb. 2009. NMW.Z.2009.013.00232: 3 ads, Ngezi Forest Reserve, Pemba L, Zanzibar, dry

forest and thieket on dark, sandy soil on eoral rag on eoast of Tondooni peninsula within reserve, leg.

BR, B.H. Warren & CFN, 8 Feb. 2009. NMW.Z.2009.013: Many additional ads and juvs from the type

loeality and other sites on Pemba L, Feb. 2009; see table 2 of Rowson et al. (2010b).

Description

Shell (Figs 32, 46, 60). Farge (4.80-5.50 mm high x 2.00-2.50 mm wide), of 6.5-7.5 whorls. Ovate-

aeuminate, spire narrowly to broadly aeuminate (spire angle 51-65°). Apex sharply pointed. Embryonie

whorls smoothly granulate. Eater whorls with relatively eoarse ribs (12-15 per mm on penultimate whorl).

Sutures shallow, shell appearing relatively straight-sided. Umbilieus elosed or nearly so. Peristome

eomplete, or ineomplete parietally. Outer palatal surfaee of aperture with a depression eorresponding

to the palatal tooth and often another eorresponding to the basal tooth. Dentition 6-fold, eonsisting of

one lamella-like parietal tooth; one slab-like palatal tooth, forming a parieto-palatal sinus; one basal

dentiele; a deep-set eolumellar baffle, sometimes folded, sub-bifid or sub-trifid; and two shallower,

broad eolumellar dentieles. Juvenile shells (Fig. 46) with 3-fold dentition, similar to some forms of D.

r radius eomb. nov.: one parietal lamella; one basal tooth; and one eolumellar thiekening. Earlier basal

teeth are retained in juveniles.

Cephalopodium. Pale yellow, with aprieot to orange tentaele retraetors.

Salivary glands (Fig. 70). United, soft, not tumid, elongate, Y-shaped; eaeh duet leaving at the apex of

the lobe and evenly thiek throughout.

Radula (Figs 66-68). With a unieuspid eentral tooth and around 15 laterals in eaeh half-row, gradually

diminishing in size laterally. All laterals bieuspid, trieuspid, or quadrieuspid, with outer eusps mueh

smaller than inner eusps. Teeth delieate, short and fiake-like at the ventral end of the radular ribbon.

Genitalia (Figs 75, 79, 80). Vas deferens appearing thiekened prior to insertion on penis, but aetually

with a short, broad parallel divertieulum. Penial sheath absent but with a thin sheath-like layer

eontiguous with wall of lower penis. Interior of penis with weak radial pilasters and small rhombie

pads. Apieal, museular part of penis with a single large hook, assoeiated with a spatulate “seoop” with

mieroseopieally serrated tip. Elsewhere in penis, one or two longitudinal rows of short, simple hooks

mounted on rhombie pads. “Spermatophore” (see Diseussion) present in penis of two Ngezi speeimens;

partially digested remains in bursa of another.

Range and habitat

Pemba Island, Tanzania, where widespread in forest and other vegetated habitats.

European Journal of Taxonomy 37 : 1-46 ( 2013 )

Remarks

This species was discussed by Rowson et al. (2010b), who suggested a thorough revision of the group

to establish whether the Pemba populations were part of a variable D. radius comb. nov. or a subspecies

or species endemic to Pemba. This uncertainty was incorporated into their conclusions on endemism

on the island (Rowson et al. 2010b: 28-29, 32). We find here that there is little or no overlap with D.

radius comb. nov. s.l. or other species, so conclude that this is an island endemic, which we here name

D. pembensis sp. nov. Like specimens of D. r radius comb. nov. on the opposite mainland at Amboni

(e.g. Figs 15, 53), D. pembensis sp. nov. is large and has a complex pattern of 6-fold dentition. However

it is distinguishable from them by its even larger size, shallower sutures, and having on average an extra

0.5 whorls. It differs from D. minuscula mahorana subsp. nov. which has similar dentition, in its larger

size, shallower sutures, and stronger and more widely spaced ribs.

Dadagulella nictitans (Rowson & Lange, 2007) comb. nov.

Figs 33, 84; Table 1

Gulella nictitans Rowson & Lange, 2007: 27-31, figs 4-7, 14-19.

Type material examined

KENYA: holotype NMK: 1 ad., Taita Hills, Macha Forest Reserve (3.45°S, 38.37°E), leaf litter in indigenous

forest (site IB), 1550 m alt, leg. C.N. Eange, 27 Nov. 1998. Paratypes NMW.Z.2007.024.00007-00009: 3 ads,

data as holotype. Paratype NMW.Z.2007.024.00010: 1 juv, data as holotype.

Other material examined

None.

Description

Shell (Fig. 33). Medium-sized (3.60-3.90 mm high x 2.00-2.20 mm wide), of 5.5 whorls. Ovate-

acuminate, although spire (spire angle 64-69°) less acuminate than in other Dadagulella gen. nov. Apex

rounded. Embryonic whorls smoothly granulate. Eater whorls with relatively coarse ribs (13-15 per

mm on penultimate whorl). Sutures of intermediate depth. Umbilicus closed or nearly so. Peristome

complete. Outer palatal surface of aperture with a long, furrow-like depression corresponding to the

palatal tooth. Dentition 5-fold or 6-fold, consisting of one V-shaped parietal tooth; one slab-like palatal

tooth, forming a conspicuous and narrow parieto-palatal sinus; and two to three shallow columellar

teeth, the lower the largest. A deep set-columellar bafihe is partly or completely hidden by the constricted

aperture, while a basal denticle is completely hidden behind the palatal tooth. However, the bafhe and

basal denticle are probably present in all specimens. One broken juvenile shell is known; it appears to

lack teeth.

Cephalopodium. Pale cream, with pale tentacle retractors.

Salivary glands. United, soft, not tumid, elongate, Y-shaped; each duct leaving at the apex of the lobe

and evenly thick throughout.

Radula. Not successfully prepared.

Genitalia. Figured in Rowson & Eange (2007).Vas deferens thickened prior to insertion on penis but

apparently without diverticulum. Penial sheath absent. Interior of penis with weak radial pilasters, a

single longitudinal pilaster, and small rhombic pads. Apical, muscular part of penis with two large

hooks, associated with a spatulate “scoop” with microscopically serrated tip. Elsewhere in penis a single

longitudinal row of short, simple hooks mounted on rhombic pads.

ROWS ON B. & TATTERSFIELD R, Revision of Dadagulella gen. nov. (Streptaxidae)

Range and habitat

Montane forest (1400-1900 m) at the type loeality and three other Forest Reserves in the Taita Hills,

southeastern Kenya (Rowson & Eange 2007).

Remarks

This speeies is ineluded in Dadagulella gen. nov. on the basis of the apieal penial seoop and hooks, and

the Y-shaped salivary gland. Its shell features are eonsistent with inelusion in Dadagulella gen. nov.

although the apex is more rounded than in other speeies. A slightly or strongly rounded apex oeeurs in

D. selene eomb. nov. andD. meredithae eomb. nov. The parieto-palatal sinus, hidden basal dentiele, and

eolumellar dentition of D. nictitans eomb. nov. allow it to be separated from other speeies.

Dadagulella frontierarum sp. nov.

um:lsid:zoobank.org:aet:9D028B17-9FCB-47E8-96Bl-5901B8E3EA4E

Figs 34, 61, 84; Table 1

Etymology

After Frontier Tanzania, the organisation whieh eolleeted the speeimens; given the ending '-arum" for a

feminine noun in the genitive plural.

Type material examined

TANZANIA: holotype NMW.Z.2003.074.00001: 1 ad., Mtai Forest Reserve (4.87°S, 38.77°E), East

Usambara Mts, Muheza Distriet (plot 69/3), submontane forest at 970 m alt., leg. Frontier Tanzania, 8 Mar.

1997. Paratypes NMW.Z.2003.074.00002: 2 ads, data as holotype. Paratype NMW.Z.2003.074.00003:

1 ad., data as holotype. Paratype NMW.Z.2003.074.00004: 1 ad., data as holotype but 7 Mar. 1997.

Paratypes NMT: 2 ads, data as holotype.

Other material examined

None.

Description

Shell (Figs 34, 61). Medium-sized (3.15-3.40 mm high x 1.80-2.05 mm wide), of 4.5-5.0 whorls. Ovate-

aeuminate, spire narrowly aeuminate (spire angle 68-77°). Apex sharply pointed. Embryonie whorls

smoothly granulate. Eater whorls with few, widely-spaeed, eoarse ribs (5-7 per mm on penultimate

whorl). Sutures very deep. Umbilieus narrowly open. Peristome eomplete, almost detaehed. Outer

palatal surfaee of aperture with a depression eorresponding to the lower palatal teeth. Dentition 6-fold,

eonsisting of one eomplex, V-shaped and flaring parietal tooth; three palatal teeth, the lower two larger

and set low down on the palatal surfaee, not forming a parieto-palatal sinus; one basal, in-running

dentiele; and one shallow but strong, in-running eolumellar tooth. Shells and anatomy of juveniles

unknown.

Range and habitat

Submontane forest (970 m elevation) in the East Usambara Mountains, northeastern Tanzania.

Remarks

The eomplex dentition and detaehed peristome of D. frontierarum sp. nov. are unlike that of any other

Dadagulella gen. nov. speeies. It is also eharaeteristie for its few whorls, few ribs and sharply pointed

apex. Biogeographieally a elose relationship with other speeies in and around the East Usambaras would

seem likely but there is no strong resemblanee.

European Journal of Taxonomy 37 : 1-46 ( 2013 )

Dadagulella delta sp. nov.

um:lsid:zoobank.org:act:BF3B1698-C8EF-4B28-89A8-DlAA85773A42

Figs 35, 72, 76, 81, 84; Table 1

""Gulella sp. 12” - Tattersfield et al. 2006: 52-53.

Etymology

From Greek 'delta', after the letter we originally used as an informal morphospeeies name for the

speeies; used as a noun in apposition.

Type material examined

TANZANIA: holotypeNMW.Z.2003.001.00019: 1 ad., Mt. Mwanihana Forest Reserve (7.82°S, 36.83°E),

Udzungwa Mts National Park, Kilombero Distriet, forest at 1050 m alt., leg. BR, PT, MBS & CFN, 29 Jan.

2003. Paratype NMW.Z.2003.OOF00020: 1 ad., data as holotype but 1200 m alt, 30 Jan. 2003.

Other material examined

None.

Description

Shell (Fig 32). Earge (4.20-4.30 mm high x 2.10-2.40 mm wide), of 6.5 whorls. Ovate-aeuminate, spire

broadly aeuminate (spire angle 56-66°). Apex pointed. Embryonie whorls smoothly granulate. Eater

whorls with eoarse, sinuous ribs (12-13 per mm on penultimate whorl). Sutures of intermediate depth.

Umbilieus narrowly open. Peristome ineomplete parietally. Outer palatal surfaee of aperture with a

depression eorresponding to the palatal tooth. Dentition weak, 3-fold (although eould be interpreted as

2-fold or 4-fold), eonsisting of: one lamella-like parietal tooth; one weak palatal tooth, not forming a

parieto-palatal sinus; a eolumellar baffle so deeply set as to be almost invisible in apertural view; and a

very weak shallow eolumellar swelling. Juvenile shells unknown.

Cephalopodium. Pale eream, with pale tentaele retraetors.

Salivary glands (Fig. 72). United, soft, not tumid, elongate, bilobed; eaeh duet leaving at the whitened

apex of the lobe and evenly thiek throughout.

Radula. Not sueeessfully prepared.

Genitalia (Figs 76, 81). Vas deferens appearing thiekened prior to insertion on penis but aetually with

an elongate, parallel divertieulum. Penial sheath absent. Interior of penis with weak radial pilasters and

small rhombie pads. Apieal part of penis with a single large hook, assoeiated with a spatulate “seoop”

with mieroseopieally serrated tip. Elsewhere in penis, an elongate eluster of short, simple hooks mounted

on rhombie pads.

Range and habitat

Montane forest (1000-1200 m) in the Udzungwa Mountains, eentral Tanzania.

Remarks

This speeies is distinetive in its simple and weak dentition, weaker than in any other Dadagulella

gen. nov. speeies exeept in D. rondoensis (Verdeourt, 1994) eomb. nov. and D. conoidea (Verdeourt,

1996) eomb. nov. whieh have a more eonieal shape. It ean be distinguished from Gulella udzungwensis

ROWS ON B. & TATTERSFIELD R, Revision of Dadagulella gen. nov. (Streptaxidae)

van Bmggen, 2003, whieh also oeeurs on Mt. Mwanihana (van Bmggen 2003), by having a more

aeuminate spire and more pointed apex, deeper sutures, sinuous ribs and no basal tooth.

Dadagulella selene (van Bmggen & Van Goethem, 1999) eomb. nov.

Figs 36, 84; Table 1

Gulella selene van Bmggen & Van Goethem, 1999: 39-40, figs 11-12.

Gulella selene - van Bmggen 2000: 233.

Type material examined

DEMOCRATIC REPUBEIC OF CONGO: paratype RBINS.MT.1645: 1 ad., Kivu Provinee, Vimnga

National Park, northern seetor, Kalivina River, tributary of the TalyaNord River, bamboo forest, Hagenia

Moeneh, 1000-2720 m (approx. 0.0°S, 29.5°E), leg. P. Vansehuytbroeek & H. Synave, 28 Aug. 1953

(examined digital photograph only).

Type locality

“D.R.Congo, Kivu Provinee, Vimnga National Park, northern seetor, at Musabaki (=Musavaki) River,

2720 m” (van Bmggen & Van Goethem 1999) (approx. 0.0°S, 29.5°E).

Other material examined

None.

Description

Shell (Fig. 36). Medium-sized (3.60-3.70 mm high x 2.10-2.20 mm wide), of 6.0-6.25 whorls. Ovate-

aeuminate, although spire (spire angle 70°) less aeuminate than in other Dadagulella gen. nov. Apex

rounded. Embryonie whorls “eompletely smooth, although there is a hint of spiral engraving” (van

Figs 36-40. Adult shells of Dadagulella gen. nov. speeies. 36. D. selene (van Bmggen & Van Goethem,

1999) eomb. nov., holotype, Vimnga NP (after van Bmggen & Van Goethem 1999). 37. D. meredithae

(van Bmggen, 2000), holotype, Nyika NP. 38. D. minuscula minuscula (Morelet, 1877), leetotype,

Nzwani. 39. D. minuscula minuscula, paraleetotype, Nzwani. 40. D. minuscula mahorana subsp. nov.,

holotype, Mayotte.

European Journal of Taxonomy 37 : 1-46 ( 2013 )

Bmggen & Van Goethem 1999). Later whorls with numerous, fine, very weak ribs (about 16 per mm on

penultimate whorl). Sutures of intermediate depth. Umbilieus elosed or nearly so. Peristome eomplete.

Outer palatal surfaee of aperture with a depression eorresponding to the palatal tooth. Dentition 5-fold,

eonsisting of one lamella-like parietal tooth; one bifid slab-like palatal tooth, not forming a parieto-

palatal sinus; a baso-eolumellar tooth or dentiele; a deep-set eolumellar baffle and a shallower eolumellar

tooth. Dentition alternatively reeognisable as 6-fold, if the bifid palatal tooth is interpreted as two teeth

(van Bmggen 1969). Juvenile shells and anatomy unknown.

Range and habitat

Montane forest (1000-2720 m) in the Albertine Rift. The type loeality and the other in the Vimnga

National Park are in bamboo forest (van Bmggen & Van Goethem 1999). Wronski & Hausdorf (2010)

also reeord it from montane forest at around 2300 m in Eehuya Forest Reserve, southwestern Uganda.

Remarks

Van Bmggen & Van Goethem (1999) eonsidered this speeies to belong to the group of speeies among

whieh van Bmggen (1969) plaeed D. browni eomb. nov. They noted that within this group D. selene

eomb. nov. was the most weakly seulptured and had distinetive dentition. These eomments are still

applieable among the larger number of speeies we attribute to Dadagulella gen. nov. However, like

D. browni semulikiensis subsp. nov., D. selene eomb. nov. is notable for oeeurring far to the northwest

of other Dadagulella gen. nov speeies. It deserves further study from the perspeetive of Conogulella

Pilsbry, 1919 (see Diseussion).

Dadagulella meredithae (van Bmggen, 2000) eomb. nov.

Figs 37, 84; Table 1

Gulella meredithae van Bmggen, 2000: 226-232, figs 1-7.

“... an as yet unidentified speeies from Malawi” - van Bmggen & Meredith 1984: 165.

Gulella meredithae - Rowson & Fange 2007: 31.

Type material examined

MAFAWl: holotype RMNH.59399: 1 ad., Nyika National Park (approx. 10.6°S, 33.8°E), Rumphi

Distriet, Juniper Forest, approx. 2100 m alt., leg. H.M. Meredith, 16 Sep. 1983.

Other material examined

None.

Description

Shell (Fig. 37). Small (2.30-3.10 mm high X 1.40-1.60 mm wide), of 5.5-“< 6” whorls. Ovate-aeuminate,

although spire (spire angle 58-77°) less aeuminate than in other Dadagulella gen. nov speeies. Apex

rounded (in holotype) to weakly pointed. Embryonie whorls smoothly granulate. Eater whorls with

very fine, very numerous ribs (about 27 per mm on penultimate whorl). Sutures of intermediate depth.

Umbilieus elosed or nearly so. Peristome ineomplete parietally. Outer palatal surfaee of aperture

with a depression eorresponding to the palatal tooth. Dentition 3-fold to 4-fold, eonsisting of at least:

one lamella-like parietal tooth; one slab-like palatal tooth, not forming a parieto-palatal sinus; and a

mammillate eolumellar baffle. Additional teeth limited to a shallow, weak eolumellar swelling. Juvenile

shells with 2-fold to 3-fold dentition: one parietal tooth; one eolumellar tooth; and usually one basal

tooth (termed labral by van Bmggen 2000). Van Bmggen (2000) showed that earlier sets of dentition are

ROWS ON B. & TATTERSFIELD R, Revision of Dadagulella gen. nov. (Streptaxidae)

visible through the shells of some transparent juveniles, even in the preeeding whorls, suggesting slow

or no resorbtion. Anatomy unknown.

Range and habitat

Montane forest (above 1500 m to around 2450 m) in northern and eentral Malawi, and adjaeent part

of Zambia (Chowo Forest). Van Bruggen (2000) suspeeted it to range into parts of Tanzania adjoining

northern Malawi.

Remarks

This speeies is distinetive in its small size, very fine, numerous ribs, and dentition. It differs from D.

radius eomb. nov. sA. and D. browni eomb. nov. sJ. in the laek of a basal tooth or dentiele. The apex is

rounded in the holotype but more eonieal in paratypes figured by van Bruggen (2000).

Dadagulella minuscula minuscula (Morelet, 1877) eomb. nov.

Figs 38-39, 62-63, 84; Table 1

Pupa minuscula MoTQlQt, 1877: 340.

Pupafischeriana Morelet, 1877: pi. XII, fig. 5 (objeetive synonym).

Enneafischeriana - Morelet 1881: 230-231. —Riehardson 1988: 103 (objeetive synonym).

Ennea (Gulella) fischeriana - Try on 1885: 100, pi. 20, fig. 39.

Ennea (Uniplicaria)fischeriana -Kobelt 1905: 166, pi. 22, fig. 10; 1910: 158.

Pupa minuscula -Kohelt 1905: 166; 1910: 158.—van Bruggen 1975: 166.

Ennea minuscula-Preston 1910: 529.

Gulella minuscula Morelet {non Gulella minuscula Emberton & Pearee, 2000) - Fiseher-Piette &

Vukadinovie 1974: 55. — Riehardson 1988: 103. — Rowson 2007a: 441-442. — Rowson & Fange

2007: 31. —Muratov2010: 277.

Type material examined

COMOROS: leetotype (here designated) andparaleetotypeNHMUK. 1893.2.4.87-88: 2 ads, “Anjouanl.”

(NzwaniT, approx. 12.22°S, 44.45°E), labelled “types”. The smaller of the two speeimens is apparently the

shell figured in Morelet (1877: pi. XII, fig. 5) under the name ''fischeriana’'. It was evidently formerly on

a white eard mount. This shell is here designated leetotype, while the larger is eonsidered a paraleetotype

(although Morelet’s deseription eould in faet have been made from a single shell).

Type locality

“Dans File d’Anjouan (Johanna)” (Morelet 1877; from title of paper).

Description

Shell (Figs 38-39, 62-63). Variable in size, shape and dentition, small to large (3.00-4.10 mm high x

1.90-2.00 mm wide), of 6.5-7.0 whorls. Ovate-aeuminate, spire broadly aeuminate (spire angle 48-63°).

Apex pointed. Embryonie whorls smoothly granulate. Eater whorls with relatively fine, numerous ribs

(10-13 per mm on penultimate whorl). Sutures relatively deep. Umbilieus elosed or nearly so. Peristome

eomplete, or ineomplete parietally. Outer palatal surfaee of aperture with a depression eorresponding

to the palatal tooth. Dentition 4-fold (leetotype. Figs 38, 62) to 5-fold (paraleetotype. Figs 39, 63),

eonsisting of one lamella-like parietal tooth; one slab-like palatal tooth, often bifid, but with parieto-

palatal sinus barely present; one basal dentiele; and one deep-set eolumellar baffie, always visible. An

additional shallow eolumellar dentiele is present in the paraleetotype. Shells and anatomy of juveniles

unknown.

European Journal of Taxonomy 37 : 1-46 ( 2013 )

Range and habitat

Nzwani (Anjouan) island in the Comoros archipelago; habitat unknown.

Remarks

Morelet (1877) described this species from Anjouan (type locality implied by the paper’s title) under

the name ''Pupa minusculd" but figured it under the name "Pupa fischeriand" (pi. Xll, fig. 5). In the

paper he compared it to three species from “He Bourbon” (Reunion), but none is very like it, suggesting

Morelet was unaware of other Dadagulella gen. nov. species and probably meaning that D. m. minuscula

comb. nov. was the first species of the group to be collected. Two of the Reunion species are streptaxids of

the Mascarene genus Gonospira Swainson, 1840 while the third belongs to the Pupillidae Turton, 1831

(see Griffiths & Florens 2006). Although Kobelt (1905, 1910) classified D. m. minuscula comb. nov.

(under fischeriana) in the subgenus Ennea (Uniplicaria) L. Pfeiffer, 1856, it is very unlike Pupa cerea

Dunker, 1848, the type species of Uniplicaria. Preston (1910) made a more relevant comparison in his

description of Ennea radius, noting that his species had coarser ribs, a columellar denticle, and a bifid

palatal tooth that were absent in D. m. minuscula comb. nov. Given these remarks and the fact that

Preston cited only Anjouan as a locality for D. m. minuscula comb, nov., it thus appears he examined

only the lectotype of the latter or material that was very like it. We agree that these features distinguish the

lectotype ofD. m. minuscula comb. nov. from that of A r radius comb. nov. However, the paralectotype

(if it is such; see also below) of D. m. minuscula comb, nov., while having similarly fine ribs, has a

stronger shallow columellar denticle that resembles that of D. r radius comb. nov. Although we have

seen no mainland specimen that exactly matches either of them, both specimens of D. m. minuscula

comb. nov. fall within the range of variation seen in D. r radius comb. nov. in other respects. It is

therefore possible that the two taxa are synonymous, in which case Morelet’s name would take priority

over Preston’s. It is even possible that D. minuscula comb. nov. sA. consists of populations of D. radius

comb. nov. sA. derived from specimens introduced from the African mainland (the alternative, that D.

radius comb. nov. sA. was introduced into East Africa from the Comoros appears unlikely given the

variation seen in East African populations and the many similar species there). That said, the isolated,

volcanic Comoros archipelago comprises a region of endemism in itself and D. minuscula comb. nov.

sA. could easily be a Comoros endemic whose overlapping variation with D. radius comb. nov. sA. is

due to homoplasy. Resolving the relationships between these two taxa, and D. m. mahorana subsp. nov.

below, is likely to require anatomical or molecular data. Until such information becomes available, and

while we focus on other issues within Dadagulella gen. nov., we maintain the two species as separate.

Van Bruggen (1986) took a similar approach with two other streptaxids, one described by Preston from

the Shimba Hills, the other by Morelet from Mayotte.

There is a nomenclatural issue concerning the name minuscula Morelet, 1877. Morelet (1881) tried to

replace the name used for the (1877) description. Pupa minuscula, with the name used for the (1877)

figure, Ennea fischeriana. Morelet (1881) wrote that the name P. minuscula had been published in error.

Subsequently both Try on (1885: 100) and Kobelt (1905: 166; 1910: 158) accepted fischeriana as the

species name. However, when revising the fauna of the Comoros, Fischer-Piette & Vukadinovic (1974)

disagreed. They treated E. fischeriana is an objective synonym of P. minuscula since the latter was not

preoccupied and was validly introduced. This point of view was followed by Richardson (1988). As a

consequence, the lectotype of D. m. minuscula comb. nov. in NHMUK also becomes the lectotype of

E. fischeriana and the latter name remains unavailable for a taxon founded on another specimen. This

includes the paralectotype, which is further discussed below.

Finally, we note that, although Emberton & Pearce (2000) described a junior homonym of minuscula

Morelet, 1877 in Gulella (G. minuscula Emberton & Pearce, 2000) no replacement name is required

since minuscula Morelet is here transferred from Gulella to Dadagulella gen. nov.

ROWS ON B. & TATTERSFIELD R, Revision of Dadagulella gen. nov. (Streptaxidae)

Dadagulella minuscula mahorana subsp. nov.

Figs 40, 64, 84; Table 1

? Ennea {Uniplicaria)fischeriana “grossere Form auf Mayotte” -Kobelt 1905: 166.

Etymology

From English noun or adjeetive ‘Mahoran’, denoting a person or thing from Mayotte; Fatinized by the

addition of the feminine ending ‘-a’ and used as a noun in apposition.

Type material examined

MAYOTTE: holotype NMW. 1955.158.25051: 1 ad., “Mayotte T”, standing as ''fischeriana''. Paratypes

NMW. 1955.158.25065: 2 ad., data as holotype.

Type locality

Mayotte (approx. 12.85°S, 45.15°E), Comoros arehipelago.

Other material examined

MAYOTTE: NHMUK. 1882.5.27.6-7: 2 ad., “Mayotte” (approx. 12.85°S, 45.15°E), standing as

"fischeriana". NHMIJK.20110171: 2 ad., “Mayotte”, standing as "fischeriana". RMNH.MOF.273928:

1 ad., “Mayotte”, standing as "fischeriana".

Description

Shell (Figs 40, 64). Variable in size, shape and dentition, large (4.30-4.60 mm high x 1.90-2.20 mm

wide), of 6.5-7.5 whorls. Ovate-aeuminate, spire narrowly aeuminate to eoeloeonoid (spire angle 50-

57°). Apex pointed. Embryonie whorls smoothly granulate. Subsequent whorls with finer, mueh more

numerous ribs than the leetotype of D. r radius eomb. nov. (15-23 per mm on penultimate whorl).

Sutures relatively deep. Umbilieus elosed or nearly so. Peristome eomplete, or ineomplete parietally.

Outer palatal surfaee of aperture with a depression eorresponding to the palatal tooth. Dentition 6-fold,

eonsisting of one lamella-like parietal tooth, sometimes slightly bifid; one slab-like palatal tooth, often

bifid, but with parieto-palatal sinus barely present; one basal dentiele; two shallow eolumellar dentieles;

and one deep-set eolumellar baffle, folded and sub-trifid, always visible. Shells and anatomy of juveniles

unknown.

Range and habitat

Mayotte; habitat unknown.

Remarks

This Mayotte subspeeies is here separated from its eounterpart on Nzwani for the first time. Speeimens of

Dadagulella gen. nov. speeies from Mayotte were first diseussed by Morelet (1881) under his proposed

replaeement name for Pupa minuscula (see above). He noted speeimens from Mayotte were larger

and had more eomplex dentition than those from Anjouan. Tryon (1885) and Kobelt (1905) followed

Morelet in treating D. fischeriana (i.e. D. minuscula eomb. nov. s.l.) as a variable speeies that was

present on the “Comoros Islands” (Tryon 1885) or on Anjouan, with a “grossere Form auf Mayotte”

(Kobelt 1905, although his dimensions of 9 mm x 2 mm must be erroneous). Both authors appear to

have eopied Morelet’s (1877) figure of the leetotype of D. m. minuscula eomb. nov., unless they saw

material very like it, and appear to have repeated (translated) Morelet’s (1881) eomments on dentition.

Tryon (1885) says “sometimes the parietal lamina is aeeompanied by a more profound very small tooth”

but this may be a misinterpretation of Morelet’s (1881) “le pli parietal pent etre aeeompagne d’une tres

petite dentieule plus profonde”, with “plus profonde” meaning ‘deeper’, rather than ‘profound’, i.e.

European Journal of Taxonomy 37 : 1-46 ( 2013 )

‘more obvious’ or ‘more important’. Morelet’s “dentieule” is likely to refer to the bifid shape of the

parietal tooth, rather than to an additional parietal dentiele (like that of D. browni eomb. nov. sJ.) whieh

is not present in any of the speeimens of D. minuscula eomb. nov. s.l. we examined. It is not elear to

whieh of the other teeth Morelet’s (1881) “einquieme petite dent, dans la gorge de la eoquille” refers,

beeause all Mayotte speeimens we examined have 6-fold dentition. It eould even be the fifth tooth, i.e.

the shallow eolumellar dentiele, of the paraleetotype of D. m. minuscula eomb. nov., sinee Morelet

(1881) is not explieit about the shell he is referring to. This raises another unfortunate possibility: that

the paraleetotype of D. m. minuscula eomb. nov., apparently from Anjouan, was in faet a speeimen

without type status that was added later, and may even have eome from Mayotte (or elsewhere), whieh

would aeeord with Morelet’s (1881) amendments to his deseription. However, we have no other reason

to doubt the existing labels.

We here introduee the name D. m. mahorana subsp. nov. for the larger, eoeloeonoid speeimens from

Mayotte, the name fischeriana Morelet being unavailable (see above). They are indeed larger than the

two speeimens of D. m. minuscula eomb. nov. (the size ranges do not overlap), are more elongate, have

a more eoeloeonoid spire, finer and more numerous ribs, and 6-fold dentition ineluding two shallow

eolumellar dentieles. These same features also distinguish them from D. radius eomb. nov. s.l, whieh

they overlap in size but only barely in the higher number of ribs per millimetre. Although as with D.

radius eomb. nov. s.l, it is diffieult to distinguish intra- and interspeeifie variability, it is elear that the

differenees between the holotype of D. m. mahorana subsp. nov. and the leetotype of D. m. minuscula

eomb. nov., eaeh eolleeted on different islands, are substantial and suffieient for most modem authors to

eonsider them distinet speeies. Alternatively, and given the residual uneertainty about the paraleetotype

of D. m. minuscula eomb. nov., they may form part of a eomplex of subspeeies. Again, further data is

needed to address this.

Dadagulella rondoensis (Verdeourt, 1994) eomb. nov.

Figs 41, 84; Table 1

Gulella rondoensis YQrdcomt, 1994: 137-139, fig. 1.

Gulella rondoensis - Verdeourt 2006: 48.

Type material examined

TANZANIA: holotype SMF.310150: 1 ad.. Rondo Forest Reserve, Rondo Plateau, Lindi Distriet

(10.12°S, 39.22°E), evergreen forest with Milicia Sim, Albizia Durazz., Dialium L., ete. in small gully

and amphitheatre around a well on esearpment, 650 m alt., leg. Bidgood, Abdallah & Vollesen, 10 Feb.

1991 (examined digital photograph only).

Other material examined

None.

Description

Shell (Fig. 41). Large (4.10 mm high x 2.20 mm wide), of 6.25 whorls. Subeonieal (maximum width

being in bottom third of the shell, at body whorl). Spire narrowly aeuminate, almost eyrtoeonoid

(eonvex) rather than eoeloeonoid (spire angle 52°). Apex sharply pointed. Embryonie whorls punetate

or malleate, rather than merely granulate. Later whorls with relatively fine ribs (about 14 per mm on

penultimate whorl). Sutures of intermediate depth. Umbilieus narrowly open. Peristome ineomplete

parietally. Dentition weak, 3-fold (although eould be interpreted as 2-fold), eonsisting of one lamella¬

like parietal tooth, with a swelling above it that reealls the one in D. conoidea; one weak palatal tooth,

not forming a parieto-palatal sinus; and one very weak, shallow eolumellar swelling. Further minute

ROWS ON B. & TATTERSFIELD R, Revision of Dadagulella gen. nov. (Streptaxidae)

swellings just pereeptible in the holotype (Fig. 41) were not noted by Verdeourt (1994) who interpreted

the dentition as 2-fold. Shells and anatomy of juveniles unknown.

Range and habitat

Forest at the type loeality, southeastern Tanzania.

Remarks

This speeies differs from D. conoidea eomb. nov. in its weaker dentition, smaller size, and in having

punetate apieal whorls. Along with D. conoidea eomb. nov. and D. delta sp. nov. it has weaker dentition

than other Dadagulella gen. nov. However, bothZ). rondoensis eomb. nov. and/), conoidea eomb. nov.

are distinetively more eonieal, i.e. less ovate than other Dadagulella gen. nov. These two are attributed to

the genus somewhat speeulatively, on the basis of their aeuminate spire and pointed apex. These features

mean they do not obviously fit into Gulella or any of its named subgenera, or indeed other plausible

streptaxid genera. No anatomieal or juvenile shell data are available for either speeies, both being

known from single speeimens. The punetuate apex of D. rondoensis eomb. nov., unique in Dadagulella

Figs 41-42. Adult shells of Dadagulella gen. nov. speeies. 41. D. rondoensis (Verdeourt, 1994)

eomb. nov., holotype. Rondo (photo by R. Janssen). 42. D. conoidea eomb. nov., holotype, Kwamgumi

(after Verdeourt 1996).

European Journal of Taxonomy 37 : 1-46 ( 2013 )

gen. nov., may not be signifieant in this respeet (see Rowson 2007b for a diseussion on the value of

apieal seulpture in distinguishing Afriean streptaxid genera). The apex of D. conoidea eomb. nov. is

less obviously punetuate, although it may have been worn smooth (Verdeourt 1996). In the deseription

of D. rondoensis eomb. nov., Verdeourt (1994) diseussed a resemblanee only to Gulella galactochila

(Crosse, 1885), a mueh larger and more broadly aeuminate Tanzanian speeies that we eonsider to laek

the eharaeteristie features of Dadagulella gen. nov. Although G. galactochila has not been disseeted, the

anatomy of another speeies very similar to it (G. udzungwensis van Bruggen, 2003) laeks the anatomieal

features of Dadagulella gen. nov. (Rowson unpublished). In his diseussion of D. conoidea eomb. nov.,

Verdeourt (1996) made no referenee to either D. rondoensis eomb. nov. or G. galactochila, but only

to two speeies that we here treat in Dadagulella gen. nov. {D. r radius eomb. nov. and D. cuspidata

eomb. nov.). We eontend firstly that D. rondoensis eomb. nov. and D. conoidea eomb. nov. are more

similar to one another than either is to G. galactochila, and seeondly that the resemblanee between D.

conoidea eomb. nov., D. r radius eomb. nov. andO. cuspidata eomb. nov. extends also to D. rondoensis

eomb. nov.. Our attribution of them to Dadagulella gen. nov. refieets this point of view.

Dadagulella conoidea (Verdeourt, 1996) eomb. nov.

Figs 42, 84; Table 1

Gulella conoidea YQrdcomt, 1996: 135-137, fig. 1.

Gulella conoidea - Tattersfield 1998b: 37. — Verdeourt 2006: 48.

Type material examined

TANZANIA: Holotype RMNH.MOL.57150: 1 ad.. East Usambara foothills, Kwamgumi/Segoma

Reserve, about 4.57°-4.58°S, 38.43°-38.45°E, near Muzi R/Sigi R. junetion, leg. Frontier Tanzania

(examined digital photograph only).

Other material examined

None.

Description

Shell (Fig. 42). Earge (5.5 mm high x 2.80 mm wide), of 8.0 whorls. Subeonieal (maximum width being

in bottom third of the shell, at body whorl). Spire narrowly aeuminate, almost eyrtoeonoid (eonvex)

rather than eoeloeonoid (spire angle 48°). Apex sharply pointed. Embryonie whorls “probably smooth

but worn” (Verdeourt 1996). Eater whorls with relatively weak ribs (about 11 per mm on penultimate

whorl). Sutures relatively shallow. Umbilieus narrowly open. Peristome ineomplete parietally, or nearly

so. Dentition 3-fold (although eould be interpreted as 4-fold), eonsisting of one lamella-like parietal

tooth, with a swelling above it that reealls that in D. rondoensis eomb. nov.; one strongly bifid palatal

tooth (or pair of teeth), not forming a parieto-palatal sinus; and a shallow eolumellar tooth or dentiele.

Shells and anatomy of juveniles unknown.

Range and habitat

Altitude and habitat not stated but probably in lowland forest at approximately 200 m at the type loeality,

northeastern Tanzania. Another speeies {D. radius eomb. nov.) also oeeurs at Kwamgumi.

Remarks

See D. rondoensis eomb. nov. above.

ROWS ON B. & TATTERSFIELD R, Revision of Dadagulella gen. nov. (Streptaxidae)

Key to species and subspecies (adult shells)

E Dentition simple, 2-fold to 4-fold (Figs 35, 38, 41,42, 51, 56, 62), never ineluding a parietal dentiele

to the left of parietal tooth . 2

- Dentition eomplex, 5-fold to 8-fold (Figs 52-55, 57-61, 63, 64) (if in doubt key from here). 8

2. Basal dentiele absent. 3

- Basal dentiele present (may be partly or wholly hidden behind palatal tooth, e.g. Fig. 56, so if

alternative fails return here). 6

3. Shell small (up to 3.1 mm high) with very fine, very numerous ribs (about 27 per mm on penultimate

whorl) (Fig. 37). D. meredithae (van Bruggen, 2000) eomb. nov.

- Shell large (4.1-5.5 mm high) and with 13 or fewer ribs per mm on penultimate whorl (Figs 35, 41-

42) . 4

4. Shell ovate-aeuminate (maximum width approximately at middle of shell); spire broadly aeuminate

(spire angle 56° or more); ribs sinuous (Fig. 35). D. delta sp. nov.

- Shell subeonieal (maximum width in bottom third of shell); spire narrowly aeuminate (spire angle

52° or less); ribs not sinuous (Figs 41-42). 5

5. One palatal tooth, eolumellar dentition weak or absent; 4.10 mm high (Fig. 41) .

. D. rondoensis (Verdeourt, 1994) eomb. nov.

- Two palatal teeth; one strong shallow eolumellar tooth; 5.5 mm high (Fig. 42).

. D. conoidea (Verdeourt, 1996) eomb. nov.

6. Deep-set eolumellar bafihe elearly visible through aperture (if not so elearly visible, then apex

pointed and not rounded) (e.g. Figs 51-57, 62-64) . 7

- Columellar bafide barely visible through aperture; apex rounded (Taita Hills, Kenya) (Fig. 33) .

. D. nictitans (Rowson & Fange, 2007) eomb. nov.

7. No eolumellar dentition other than deep-set bafde; basal dentiele present, either eompletely hidden

by a slab-like palatal tooth or visible only when shell is turned (Kimboza, Uluguru Mts, Tanzania)

(Figs 28, 56). D. ecclesiola sp. nov.

- Usually with shallow eolumellar dentition as well as deep-set bafide; basal dentiele elearly visible in

normal apertural view. 19

8. Parietal tooth strongly V-shaped, fiaring, direeted outwards; three palatal teeth; shell of 5.0 or fewer

whorls (Figs 34, 61). D. frontierarum sp. nov.

- Shell without this eombination of eharaeters. 9

9. Upper shallow eolumellar tooth strong, squarish, projeeting into aperture as far or further than any

deeper eolumellar bafide (Figs 30-31, 36, 58-59) . 10

- Shallow eolumellar dentition projeeting less far into aperture than any deeper eolumellar bafde (Figs

51-57,60-64) . 12

10. Apex rounded; spire broadly aeuminate, spire angle 70°; ribs weak, almost disappearing in the

middle of the whorl (Virunga NP, DR Congo) (Fig. 36) .

. D. selene (van Bruggen & Van Goethem, 1999) eomb. nov.

- Apex sharply pointed; spire aeuminate or eoeloeonoid, spire angle 65° or less; ribs prominent,

running from suture to suture; apex finely radially striate in fresh speeimens (Figs 30-31) . 11

European Journal of Taxonomy 37 : 1-46 ( 2013 )

11. Shell more than 4.0 mm high; parietal tooth simple, lamella-like; two palatal teeth (W. Usambara

Mts, Tanzania) (Figs 31, 59). D. cuspidata (Verdeourt, 1962) eomb. nov.

- Shell up to 4.0 mm high; parietal tooth V-shaped; one palatal tooth (elsewhere in Tanzania) (Figs 30,

58) . D. minareta sp. nov.

12. Outer palatal surfaee of aperture with very deep, long, furrow-like depression; basal dentiele absent

or not visible in apertural view (Figs 29, 33, 57). 13

- Outer palatal surfaee of aperture with shallow, not furrow-like depression; basal dentiele elearly

visible in apertural view. 14

13. Apex sharply pointed; spire eoeloeonoid (Ngorongoro, Tanzania) (Figs 29, 57).. D. cresswelli sp. nov.

- Apex rounded (Taita Hills, Kenya) (Fig. 33). D. nictitans (Rowson & Lange, 2007) eomb. nov.

14. Seulpture of widely-spaeed, flaring, subtriangular, lamella-like ribs; spire narrowly aeuminate (spire

angle <50°) (Cabo Delgado, Mozambique) (Fig. 27). D. delgada (Muratov, 2010) eomb. nov.

- Shell without this eombination of eharaeters. 15

15. Shell 4.8 mm-5.5 mm high (Pemba L, Tanzania) (Figs 32, 60) . D. pembensis sp. nov.

- Shell up to 4.6 mm high. 16

16. Shell up to 3.3 mm high; parieto-palatal sinus narrow, appearing parallel-sided in apertural view;

nearly always with an additional, small parietal dentiele to the left of parietal lamella; nearly always

with two basal dentieles (Figs 21-26, 54-55)... 17 (D. browni (van Bruggen, 1969) eomb. nov. sJ.)

- Shell up to 4.6 mm high; parieto-palatal sinus wide, not parallel-sided in apertural view; always without

a small parietal dentiele to the left of parietal lamella, even if shell less than 3.3 mm high. 19

17. Only one basal dentiele present; no eolumellar dentition other than a deep-set baffle (Mafla L,

Tanzania) (Figs 21, 54) . D. browni mafiensis subsp. nov.

- Two basal dentieles almost always present; one or two eolumellar dentieles present in addition to a

deep-set baffle (Figs 22-26, 55) . 18

18. Shell 3.15-3.25 mm high; sutures relatively shallow; peristome eomplete; parieto-palatal sinus

longer and narrower, not widening appreeiably towards its inner end (Semuliki NP, Uganda) (Figs

22, 55; eompare with Figs 23-26). D. browni semulikiensis subsp. nov.

- Shell (2.55-3.20 mm high; sutures relatively deep; peristome ineomplete parietally; parieto-palatal

sinus shorter and broader, widening towards its inner end (South Afriea, Mozambique, S. Tanzania)

(Figs 23-26; eompare with Figs 22, 55). D. browni browni (van Bruggen, 1969) eomb. nov.

19. Seulpture of numerous, flne ribs (15 or more per mm on penultimate whorl); spire eoeloeonoid,

elongate (Mayotte) (Figs 40, 64) . D. minuscula mahorana subsp. nov.

- Shell without this eombination of eharaeters (e.g. Figs 1-20, 38-39). 20

20. Shell 4.0 mm or more high, seulpture of few, eoarse, widely-spaeed ribs (5-10 per mm on penultimate

whorl); dentition 4-fold to 5-fold (Tam Desert and elsewhere in SE Kenya) (Figs 6-8, 51).

. D. radius calva (Connolly, 1922) eomb. et stat. nov.

- Shell without this eombination of eharaeters. 21

21. Dentition 5-fold or more, although when shallow eolumellar dentition is weak, might be eonsidered

4-fold (lowlands of SE. Kenya andNE. Tanzania) (Figs 1-5, 9-20, 52-53) .

. D. radius radius (Preston, 1910) eomb. nov.

- Dentition 4-fold, laeking a shallow eolumellar tooth (as in the leetotype) or 5-fold, with a shallow

eolumellar tooth (as in the paraleetotype) (Nzwani [Anjouan] L, Comoros) (Figs 38-39, 62-63).

. D. minuscula minuscula (Morelet, 1877) eomb. nov.

ROWS ON B. & TATTERSFIELD R, Revision of Dadagulella gen. nov. (Streptaxidae)

Discussion

Shell

The shells of Dadagulella gen. nov. differ from those of Gulella sA. in the aeuminate (sometimes

eoeloeonoid or eyrtoeonoid) spire; the majority also has a mueh more pointed apex. They are never

smooth, unlike many Gulella. Patterns of adult apertural dentition that are superfieially similar to those

of Dadagulella gen. nov. speeies oeeur in some Gulella, espeeially where the dentition is simple, but

not in eombination with the above mentioned features. A possible exeeption is the South Alfiean G.

contingens Bumup, 1925 whieh has an aeuminate spire but rounded apex (TV. Muratov 2012, pers.

eomm.). It may prove to belong to Dadagulella gen. nov. The shells of juvenile Dadagulella gen. nov.

are almost always dentate, while those of juvenile Gulella are very rarely so. Two speeies of Gulella s.l.

Figs 43-50. Juvenile shells of Dadagulella gen. nov. speeies, with dentition emphasized in basal view

ineluding earlier dentition visible through the shell. 43-45. D. radius radius, Mbudya I. 46. D. pembensis

sp. nov., Ngezi. 47-48. D. browni browni, Mwanihana. 49. ? D. ecclesiola sp. nov., Kimboza. 50. D.

minareta sp. nov., Kimboza.

European Journal of Taxonomy 37 : 1-46 ( 2013 )

with dentate juveniles, the Tanzanian G. subringens (Crosse, 1886) and the South Afriean G. phyllisae

Bumup, 1925 have been shown not to belong to Dadagulella gen. nov. by sequeneing (Rowson et al.

2010a; Rowson & Herbert, unpublished). The shells of Dadagulella gen. nov. do not eorrespond to

those of any other existing streptaxid genus or subgenus. They differ from those of the Central-West

Afriean Conogulella Pilsbry, 1919 in having a more aeuminate spire and more pointed apex, in laeking

strong spiral striae on the apieal whorls, and in dentition. Moreover, juvenile Conogulella laek apertural

dentition (A.J. de Winter 2012, pers. eomm.). Dadagulella selene eomb. nov., a speeies with a more

rounded apex and apieal whorls with “a hint of spiral engraving”, deserves further investigation from

this perspeetive although its authors (van Bruggen & Van Goethem 1999) eonsidered it part of the

same group as D. browni eomb. nov.. Another Central-West Afriean genus, Costigulella Pilsbry, 1919

has dentate juveniles (de Winter 2008) but is otherwise unlike Dadagulella gen. nov. A further speeies

from the eentral Congo basin, Ptychotrema (Nsendwea) nobrei Dupuis & Putzeys, 1923 (sole speeies

51 52 53 54 55

Figs 51-64. Apertures of adult shells of Dadagulella gen. nov. speeies (not to seale). 51. D. radius calva,

leetotype, Tam Desert. 52. D. radius radius, leetotype, Shimba Hills. 53. D. radius radius, Amboni.

54. D. browni mafiensis subsp. nov., holotype, Mlula. 55. D. browni semulikiensis subsp. nov., holotype,

Semuliki NP. 56. D. ecclesiola sp. nov., holotype, Kimboza (two views of shell showing basal tooth only

visible when shell angled). 57. D. cresswelli sp. nov., holotype, Ngorongoro. 58. D. minareta sp. nov.,

holotype, Kimboza. 59. D. cuspidata eomb. nov., holotype, Shume (note shell angled slightly differently

to that in Fig. 31). 60. D. pembensis sp. nov., holotype, Ngezi. 61. D. frontier arum sp. nov., holotype,

Mtai. 62. D. minuscula minuscula, leetotype, Nzwani. 63. D. minuscula minuscula, paraleetotype,

Nzwani. 64. D. minuscula mahorana subsp. nov., holotype, Mayotte.

ROWS ON B. & TATTERSFIELD R, Revision of Dadagulella gen. nov. (Streptaxidae)

of Nsendwea Dupuis & Putzeys, 1923) superfieially resembles some Dadagulella gen. nov. in size and

in its broadly eonieal spire. However, we agree with Adam et al. (1995) and Sehileyko (2000) that its

other shell features, ineluding the inrunning palatal fold, support its elassifieation among the subgenera

of Ptychotrema E. Pfeiffer, 1853. Other Ptychotrema ean likewise be distinguished from Dadagulella

gen. nov. by these folds. Finally, juvenile Dadagulella gen. nov. are distinet from the East Afriean

Juventigulella Tattersfield, 1998, with whieh they oeeur at several Tanzanian sites (Tattersfield 1998b)

in laeking the downtumed aperture of adult Juventigulella.

Salivary glands

The shape of Dadagulella"s salivary glands differs from that of the type speeies of Gulella in whieh the

united gland is not appreeiably divided into two lobes (Rowson & Herbert, unpublished).

Radula

Dadagulella gen. nov. is highly unusual among Streptaxidae in having multieuspid radular teeth. Aiken

(1981) believed the eontrast between the radula of D. browni eomb. nov. and of all other South Afriean

Gulella he studied was indieative of a differenee in diet. The only prior reports of multieuspid streptaxid

radulae appear to be Connolly (1930) and Verdeourt (1953) (see also Verdeourt 1990). Connolly (1930)

Figs 65-68. Radulae of Dadagulella gen. nov. speeies. 65. D. radius radius, Amboni. 66. D. pembensis

sp. nov., Ngezi, radula in situ on odontophore, ventral view. 67. D. pembensis sp. nov., Ngezi. 68. D.

pembensis sp. nov., Ngezi, teeth from ventral end of radular ribbon. Abbreviations: e = eentral tooth;

ii, iii, and iv = examples of bi-, tri- and quadrieuspid teeth respeetively. All sealebars =10 pm.

European Journal of Taxonomy 37 : 1-46 ( 2013 )

Figs 69-76. Salivary glands and genitalia of Dadagulella gen. nov. speeies. 69. D. radius radius, Amboni.

70. D. pembensis sp. nov., Ngezi. 71. D. browni browni, Kosi Bay. 72. D. delta sp. nov., Mwanihana.

73. D. radius radius, Amboni. 74. D. browni browni, Kosi Bay. 75. D. pembensis sp. nov., Ngezi.

76. D. delta sp. nov., Mwanihana. Abbreviations: ad = anterior duet of salivary gland; ag = albumen

gland; at = atrium; be = bursa eopulatrix; bd = bursa duet; dv = divertieulum of vas deferens; e = egg; fo

= free oviduet; hdd/t = hermaphroditie duet divertieulum / talon; ma = museular apex; p = penis; pd =

posterior duet of salivary gland; pr = penial retraetor musele; v = vagina; vd = vas deferens.

ROWS ON B. & TATTERSFIELD R, Revision of Dadagulella gen. nov. (Streptaxidae)

deseribed the radula of the Ugandan Conogulella conospira subsp. polynematica (Pilsbry, 1919) as

eonsisting of evenly bieuspid teeth (formula 47-1-47); this is also seen in other Conogulella speeies,

the eusps being nearly equal in size (A.J. de Winter 2012, pers. eomm.). Verdeourt (1953) deseribed the

radula of the Tanzanian Gulella usambarica (Craven, 1880) as eonsisting almost entirely of multieuspid

teeth (formula 26-1-26); again, the eusps are nearly equal in size. Sequenee data (Rowson et al. 2010a)

show that Dadagulella gen. nov. is only distantly related to G. usambarica. Thus a relationship between

Dadagulella gen. nov. and Conogulella based on the multieuspid radula eannot be ruled out, but remains

unproven sinee multieuspid teeth oeeur in distantly related lineages.

Genitalia

Dadagulella gen. nov. differs from the type speeies of Gulella (Rowson & Herbert, unpublished) in

having a swelling of or divertieulum on the vas deferens, a museular apex, an apieal penial “seoop” and

one or more large apieal penial hooks, and a smaller number of penial hooks in total. The genitalia differ

Figs 77-83. Penes and spermatophores of Dadagulella gen. nov. speeies. 77. D. radius radius, Amboni,

penis with spermatophore in situ. 78. Same, hook and seoop in situ when hook lifted baek to expose

seoop. 79. D. pembensis sp. nov., Ngezi, penis. 80, same, spermatophore from penis of another individual.

81. D. delta, Mwanihana, penis. 82. D. browni browni, Kosi Bay, penis (diagrammatie). 83. D. radius

crassa, Amboni, seoop magnified to show serrated edge. Abbreviations: el = ehitinized lobe; Ih = large

apieal hook; ma = museular apex; pr = penial retraetor musele; se = apieal seoop; st = spermatophore

tail; sr = spermatophore reservoir; vd = vas deferens.

European Journal of Taxonomy 37 : 1-46 ( 2013 )

from those of Conogulella (AJ. de Winter 2012, pers. eomm.) and many other West Afriean speeies

eurrently attributed to Gulella or its subgenera (Degner 1934; Sehileyko 2000) in the same respeets. The

observation of “spermatophores” in the penis is unusual in Streptaxidae. The only prreviously published

report appears to be de Winter et al. (1999), although similar struetures have reeently been found in the

penes of other Gulella and former Gulella speeies (Rowson, unpublished; de Winter 2012, pers. eomm.).

It is not eertain that the struetures in Dadagulella gen. nov. are true spermatophores, espeeially if they

are formed attaehed to the penial wall. Muratov (1999: 16) and Sehileyko (2003: 1467) state that the

somewhat similar struetures in eertain Trigonoehlamydoidea Hesse, 1882 remain in the penis but empty

their eontents (perhaps a earbonate buffer; Muratov, 1999) into the partner’s bursa. Muratov (1999)

notes that their eavities eannot be filled with sperm without a eonneetion to the epiphallus, but also that

this does not rule out that Trigonoehlamydoidea also produee true spermatophores. We note only that

in D. pembensis sp. nov., the remains in the bursa resemble the tail of the struetures found in the penis

Molecular data

The available DNA sequenee data suggest that Dadagulella gen. nov. is a sister lineage to the one that

eontains Gulella, and that the two beeame separated in the Cenozoie (Rowson et al. 2010a). Further

sequenee data from additional speeies of Dadagulella gen. nov. and Gulella (Rowson & Herbert

unpublished), ineluding the type speeies of Gulella (Herbert & Rowson 2011), support the eonelusion

that the two lineages are separate, and that Gulella ineludes the type speeies of several nominal subgenera

while Dadagulella gen. nov. does not. The two ean be distinguished by the presenee of an indel in the

mitoehondrial DNA sequenee eoding for COl (eytoehrome oxidase 1) in almost all Gulella speeies

relative to Dadagulella gen. nov. (Rowson et al. 2010a; Rowson & Herbert, unpublished). Another

streptaxid speeies ineluded in the “G. radius group” by van Bruggen (1969, 2000), the Kenyan G.

pretiosa (Preston, 1911), has also been sequeneed (Rowson et al. 2010a). It is elearly not elosely related

to either Dadagulella gen. nov. or Gulella itself, a finding also supported by anatomieal data (Rowson

unpublished).

Fossil record

There are many East Afriean Mioeene fossil taxa attributable to Gulella (e.g. Verdeourt 1963; Piekford

2009). It would be reasonable to expeet Dadagulella gen. nov. to oeeur with them if the two had diverged

by the Mioeene. However, none appear to unambiguously show the features of Dadagulella gen. nov.

Two speeies, G. leakeyi Verdeourt, 1963 and Gulella sp. C of Verdeourt 1963 from Rusinga resemble

Dadagulella gen. nov. in their aeuminate spires, and his figure 53 of G. sp. C (speeimen 631/55)

resembles D. radius eomb. nov. but he does not diseuss its affinities. The speeies G. moja Piekford, 2009

from Korn seems only superfieially similar to Dadagulella gen. nov. All are from a part of East Afriea

from whieh no extant Dadagulella gen. nov. are known, although the distributions of other streptaxids

found in the Rusinga and Korn deposits have evidently ehanged (Verdeourt 1963; Piekford 2009).

Biogeography

Dadagulella gen. nov. is evidently an eastern group that has not yet been reeorded from West Afriea or

the Congo basin (Fig. 84). It eurrently appears most diverse in Tanzania and Kenya, with endemies in

both highlands and lowlands. There are no reeords of Dadagulella gen. nov. from the voleanie highlands

of eentral and northern Kenya, south-western Tanzania, or northern Tanzania, with the exeeption of

D. cresswelli sp. nov. at Ngorongoro. Two speeies oeeur in the Albertine Rift region of endemism, but

the group is otherwise unknown from the Guineo-Congolian forests of western Kenya or Uganda. This

represents a disjunetion of around 500 km aeross an area whose forest molluses have been fairly well-

studied (e.g. Wronksi & Hausdorf 2010; Tattersfield & Rowson unpublished). In eontrast, the genus

oeeurs widely in eastern Kenya and eentral-eastem Tanzania, ineluding throughout the famed Eastern

Are Mountains region of endemism, where eight speeies oeeur. Of these, six appear to be endemie

ROWS ON B. & TATTERSFIELD R, Revision of Dadagulella gen. nov. (Streptaxidae)

to a single mountain range: Taita {D. nictitans eomb. nov.); W. Usambara {D. cuspidata eomb. nov.);

E. Usambara {D. frontier arum sp. nov., D. conoidea eomb. nov.); Ulugum {D. ecclesiola sp. nov.); and

Udzungwa {D. delta sp. nov.). These are all montane or submontane, oeeurring around or above 1000 m

elevation, exeept for D. ecclesiola sp. nov. and D. conoidea eomb. nov. whieh are found in the foothills.

Endemies are not yet known from three other major Eastern Are ranges: Nguru {D. minareta sp. nov.

only), Mahenge {D. radius eomb. nov. only) and Ukaguru (no Dadagulella gen. nov. reeorded). The

lowland U. radius eomb. nov. sd. is apparently replaeed in some areas by other narrow-range endemies,

namely D. pembensis sp. nov. or subspeeies we attribute to D. browni eomb. nov. sd., whieh evidently

ranges into southern Tanzania. However, in three plaees in Tanzania D. radius eomb. nov. has been

found in sympatry with other Dadagulella gen. nov. speeies. We take this in support of the hypothesis

that sueh speeies are good genetie speeies rather than eeophenotypie variants. The apparent greater

interpopulation variability seen in the lowland speeies D. radius eomb. nov. /. and D. browni eomb. nov.

/. may refleet their wider ranges and the larger number of populations sampled. It remains possible that

those populations most similar to typieal D. radius eomb. nov. form a eomplex of Pleistoeene, ineipient.

Indian

Ocean

Comoros Is.

a ^

MAYOTTE

MADAGASCAR

• D. browni browni

★ D. b. ma^ensis

■ D. b. semulikiensis

# D. cresswelli

★ D. delgada

■ D. min. mahorana

A D. meredithae

• D. min. minuscula

★ D. rondoensis

■ D. selene

♦ D. conoidea

★ D. cuspidata

■ D. delta

• D. ecclesiola

★ D. frontieranum

■ D. minareta

A D. nictitans

• D. pembensis

★ D. radius calva

■ D. r. radius

40 E

KENYA

★

Taita

Hills

Tam

Desert

Fig. 84. Distribution of Dadagulella gen. nov. speeies, based on published reeords and material examined

here (for additional reeords of D. meredithae eomb. nov. in northern Malawi see van Bruggen 2000).

The inset on the left eorresponds to the map on the right.

European Journal of Taxonomy 37 : 1-46 ( 2013 )

or hybridizing species, while the more distinctive populations (several of them montane endemics) are

fully isolated species that separated earlier.

Beyond East Africa, Dadagulella gen. nov. is represented by at least one species in Malawi and Zambia,

two in eastern Mozambique, one in eastern South Africa and one on the Comoros archipelago. The

genus does not appear to have been found on Madagascar, although the genus Gulella s.l is richly

represented there (e.g. Emberton 2001). There is some shell resemblance between Dadagulella gen. nov.

and the following Madagascan species: G. ambatovakiae Emberton, 2001; G. benjamini Emberton &

Pearce, 2000; G. hafa Emberton, 2001; G. hafahafa Emberton, 2001; G. mahafinaratra Emberton,

2001; G. manomboae Emberton, 2001; G. michellae Emberton, 2001; G. vakinifia Emberton, 2001

and G. vatosoa Emberton, 2001. This is a heterogenous group of species that does not key out together

in the key of Emberton (2001). In each case the spire is acuminate, although slightly less so than in

most Dadagulella gen. nov.. Juvenile teeth are not mentioned in the text of Emberton (2001) and a

juvenile shell of one species (a paratype of G. hafahafa) is figured and appears to lack apertural teeth.

The genitalia of G. benjamini were figured by Emberton & Pearce (2000) and show an apical caecum

and two longitudinal pilasters with a broad area between them. There is no apical scoop. The genitalia

of G. reeae Emberton & Pearce, 2000 in the same paper have a vas deferens and pilasters more typical

of Gulella (Rowson & Herbert unpublished). The anatomy of G. vatosoa is broadly identical to that of

G. benjamini (unpubl. obs.). The anatomy of G. hafahafa, which is a Gulella based on sequence data

(Rowson et al. 2010) is unlike that of G. benjamini or G. vatosoa in having few, larger hooks (unpubl.

obs.) and unlike that of Dadagulella gen. nov. We conclude that Dadagulella gen. nov. as recognised

here is not yet known from Madagascar. If Dadagulella gen. nov. and Gulella are indeed sister groups,

these ibservations support an African, rather than Madagascan, divergence of the two.

Acknowledgements

We are grateful to A. J. de Winter (RMNH) for discussions and data on Conogulella, and photographs of

RMNH specimens of Dadagulella gen. nov. We thank D.G. Herbert (NMSA) for generously providing

material, J. Ablett (NHMUK), C.F. Ngereza (NMT), J. Slapcinsky (Florida Museum of Natural History,

USA), D. Van den Spiegel and D. De Bakker (MRAC) for lending material and R. Janssen (SMF) for

the photograph of D. rondoensis comb. nov. Comments from TV. Muratov (NMSA), D.G. Herbert and

an anonymous referee helped to improve the manuscript.

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Manuscript received: 30 June 2012

Manuscript accepted: 5 December 2012

Published on: 4 February 2012

Topic editor: Rudy Joeque

Section editor: Kurt Jordaens

Desk editor: Charlotte Thionois