ockraspiris, Bulimulus, Branson & M c C o y , 1965b: 97 (Mexico, Campeche, ...... A n annotated list of the snails of Dallas County, ...... Consul-General, Quito.
SYSTEMATICS, PHYLOGENY AND ZOOGEOGRAPHY OF BULIMULINAE (MOLLUSCA) by
A. S. H . B R E U R E D i v i s i o n o f Systematics and E v o l u t i o n a r y B i o l o g y , c/o R i j k s m u s e u m v a n N a t u u r l i j k e H i s t o r i e , L e i d e n W i t h 182 text-figures, 5 tables and 3 plates CONTENTS I. II. III. IV. V. VI. VII. VIII. IX. X. XL
Abstract Introduction Ecology M a t e r i a l and methods T e r m i n o l o g y and character analysis Acknowledgement s Systematics P h y l o g e n y and zoogeography I n t r a f a m i l i a r relationships Summary/Resumen References Index
3 3 4 8 9 21 22 138 162 165 168 201
ABSTRACT I n this publication a revision is g i v e n o f the genera o f the s u b f a m i l y B u l i m u l i n a e (Gastropoda, P u l m o n a t a , B u l i m u l i d a e ) . T h e morphologica l v a r i a t i o n o f the shell, radula, p a l l i a l organs and genitalia is analized and 21 character transition series are recognized. I n the systematical part the f o l l o w i n g data are presented f o r each genus : description o f shell and anatomy, distribution, ecology, bibliograph y and a list o f t a x a . T h e number o f (sub) genera is reduced f r o m 80 to 43 ( + t w o n o m i n a i n q u i r e n d a ) . A new synonymy is : Paracochlea H y l t o n Scott, 1967 = Eudioptus A l b e r s , 1860. T h e f o l l o w i n g new species names are introduced : Bostryx sophieae, Drymaeus (Drymaeus) marcapatensis, Drymaeus (Drymaeus) sophieae, Drymaeus (Mesembrinus) pseudobesus. Berendtia digueti M a b i l l e is designated type species o f Teneritia M a b i l l e ; Helix zoographica d ' O r b i g n y is designated type species o f Hamadryas A l b e r s . Based on the transition series mentioned above, hypotheses o f phylogenetic relationships are presented f o r the genus groups, u s i n g the methods o f H e n n i g . T h e relationships between the f i v e subfamilies o f the B u l i m u l i d a e are also investigated but remain tentative. I n the zoogeographical section the various theories are reviewed and their relevance f o r the distribution o f the B u l i m u l i n a e is treated, u s i n g the hypotheses o f phylogenetic relationships and C r o i z a t ' s vicariance theory. I.
INTRODUCTION
D u r i n g a r e v i s i o n o f some b u l i m u l i d g e n e r a f r o m the W e s t Indies ( B r e u r e , 1974a, 1975a), I became a w a r e o f the m u l t i p l e t a x o n o m i c p r o b l e m s o f f e r e d
ZOOLOGISCHE V E R H A N D E L I N G E N
4
168
(1979)
b y the m a i n l a n d t a x a . E s p e c i a l l y the publications b y W e y r a u c h (1956-1967) and
V a n M o l (1971) p r o v o k e d m y interest. T h a n k s to grants o f the F o u n -
d a t i o n f o r the A d v a n c e m e n t o f T r o p i c a l R e s e a r c h ( W O T R O ) I c o u l d initiate a r e v i s i o n o f the g e n e r a o f B u l i m u l i d a e i n 1975. B e c a u s e o f the l i m i t e d time available f o r this research, the p r e s e n t p a p e r is e n t i r e l y c o n f i n e d to the B u l i m u l i n a e . D a t a o n the other s u b f a m i l i e s
have
p a r t l y been p u b l i s h e d ( B r e u r e , 1974c) o r w i l l be p u b l i s h e d i n the n e a r f u t u r e (Breure & Schouten, i n preparation). In
m y v i e w the a n a t o m y yields indispensable i n f o r m a t i o n f o r a generic
r e v i s i o n a n d , therefore, as v e r y f e w p r e s e r v e d a n i m a l s w e r e available i n c o l lections, it w a s necessary possible
b y grants
to collect m o s t m a t e r i a l m y s e l f .
T h i s was made
o f the F o u n d a t i o n f o r the A d v a n c e m e n t
of Tropical
R e s e a r c h ( W O T R O ) . T h e specimens w h i c h I collected d u r i n g t w o f i e l d t r i p s to S o u t h A m e r i c a were the basis f o r the results o f m y research. Thanks
to the m o n u m e n t a l m o n o g r a p h
( 1 8 9 5 - 1 9 0 2 ) , a l l description s
previously
o f the B u l i m u l i d a e b y
published were
readily
Pilsbry
available,
w h i c h greatly facilitated the i d e n t i f i c a t i o n o f the m a t e r i a l . I n P i l s b r y ' s m o n o g r a p h the g e n e r i c f r a m e w o r k is based o n one o f the m o s t i m p o r t a n t e x t e r n a l m o r p h o l o g i c a l characters o f the B u l i m u l i d a e , v i z . the sculpture o f the p r o t o conch. U n t i l n o w this character has r e t a i n ed its i m p o r t a n c e f o r identifications, but it p r o v es to be o f less value as i n d i c a t o r o f phylogenetic relationships. T h i e l e (1931) listed 11 g e n e r a a n d subgenera i n the s u b f a m i l y B u l i m u l i n a e ( P a r t u l a a n d Placostylus
n o w excluded). I n Zilch's monograph o n euthyneu-
r a n gastropods the n u m b e r o f ( s u b ) g e n e r i c t a x a i n the s u b f a m i l y h a d g r o w n to 8 0 (also i n d u c e d b y a c h a n g e d concept o f Placostylus
a n d Aspastus
(sub)genera;
Diplomorpha,
n o w e x c l u d e d ) . W i t h three genera c l a s s i f i e d s e n s u
lato this n u m b e r is n o w r e d u c e d to 43 ( p l u s t w o n o m i n a i n q u i r e n d a ) . T h e v a r i o u s classifications a r e s u m m a r i z e d i n T a b l e 1. II. F i e l d observations
ECOLOGY
d u r i n g J a n u a r y - J u n e 1975
(see B r e u r e , 1975b) a n d
A p r i l - M a y 1976 indicate that b u l i m u l i d species f r e q u e n t the f o l l o w i n g m a c r o habitats:
(1)
leaf
litter l a y e r ;
( 2 ) herbaceous vegetation;
(3)
rock-faces;
( 4 ) trees. T h e species o f the leaf litter l a y e r a r e characterized b y t h e i r m o s t l y u n i f o r m b r o w n i s h c o l o u r , the u n e x p a n d e d p e r i s t o m e a n d the straight
r o w s o f the
r a d u l a . T h e species were collected b y r a k i n g t h r o u g h leaves a n d debris o n the ground. Species that live i n herbaceous vegetation m a y less f r e q u e n t l y also be f o u n d i n the leaf litter layer. T h e y a r e characterized b y their light ( o f t e n
whitish)
VO
M
M
ON
w
9
BREURE, BULIMULINAE
T h e f o l l o w i n g abbreviations are u s e d to r e f e r to the location o f the imens: A M , Australian M u s e u m , Sydney; Natural History, Ne w
A M N H ,
York; A N S P , Academy
spec-
American M u s e u m of
o f N a t u r a l Sciences,
Phil-
adelphia; B M N H , B r i t i s h M u s e u m ( N a t u r a l H i s t o r y ) , L o n d o n ; C A S , C a l i f o r n i a A c a d e m y o f Sciences, D e p t . o f G e o l o g y , S a n F r a n c i s c o ; C M , C a r n e g i e M u s e u m , Pittsburgh; C M G , M u s e o Civico di Storia Naturale, Genua; D G M , D i v i s ã o de
Geologia
e Mineralogia, Departamento
N a c i o n a l de
Produção
M i n e r a l , R i o de J a n e i r o ; D Z S P , D e p a r t a m e n t o de Z o o l o g i a , U n i v e r s i d a d e de São Paulo, São Paulo; F M N H , F i e l d M u s e u m of Natural History , Chicago; I M L , Instituto M i g u e l L i l l o , T u c u m á n ; I O C , Instituto O s w a l d o C r u z , R i o de Janeiro;
IRSN,
Institut
R o y a l des
Sciences
Naturales, Brussels;
MACN,
M u s e o A r g e n t i n o de Ciências N a t u r a l e s , B u e n o s A i r e s ; M C Z , M u s e u m o f Comparative
Zoology,
turelle, G e n e v a ; Plata;
Cambridge
M L P , Museo
Janeiro;
(Mass.);
MHNG,
M I H S , private collection o f
M N H N,
La
Plata, L a
Plata;
Musée d'Histoire N a -
D r . M . I. H y l t o n Scott,
M N , Museu
Muséum National d'Histoire Naturelle,
Paris; NMB,
historisches
Victoria,
bourne;
NRS,
Basel;
NMV,
Naturhistoriska
National
Riksmuseet,
M u s e u m of Stockholm;
de
MRCN,
M u s e u R i o - G r a n d e n s e de Ciências N a t u r a i s , P o r t o A l e g r e ; Museum,
La
Nacional, R i o
Natur-
RMNH,
MelRijks-
museum van Natuurlijke Historie, L e i d e n ; S A M , South Australian M u s e u m , Adelaide;
SMF,
Natur-Museum
Senckenberg,
F l o r i d a State M u s e u m , G a i n e s v i l l e ;
UMMZ,
Frankfurt am
Main;
UF,
University of Michigan, M u -
seum o f Z o o l o g y , A n n A r b o r ; U S N M , N a t i o n a l M u s e u m o f N a t u r a l H i s t o r y , Washington; voor
W A M , Western Australian Museum, Perth; Z M A ,
Taxonomische
Zoologisches
Zoologie
(Zoologisch
Amsterdam;
M u s e u m der H u m b o l d t - U n i v e r s i t ä t , B e r l i n ;
sches M u s e u m , U n i v e r s i t ä t Zürich, T h e status o f the type species L T , lectotype; N T , neotype; The
Museum),
ZMUZ,
ZMB, Zoologi-
Zürich. is abbreviated
as follows:
P T , paratype ( s ) ; S T ,
f o l l o w i n g abbreviations
Instituut
HT,
holotype;
syntype(s).
are u s e d to r e f e r to parts o f the
anatomy:
E P , epiphallus; F L , flagellum; P , penis; P S , penis sheath. IV.
TERMINOLOGY AND CHARACTER ANALYSIS
T h e t e r m i n o l o g y o f the shell m o r p h o l o g y is largely i n accordance w i t h that used b y
Pilsbry
(1895-1902). S e v e r a l m o r p h o l o g i c a l structures
are
never-
theless d e f i n e d b y figures, not o n l y to facilitate the descriptions i n c l u d e d i n this paper, but also to p r o v i d e a reference f o r future w o r k . T h e shell shape m a y be ovate ( f i g . 71), conical ( f i g . 161), (fig.
81).
elongate-ovate ( f i g .
i n ) , ovate-
conical, globose ( f i g . 166), t u r r i t e d ( f i g . 126), o r f u s i f o r m
T h e u m b i l i c u s m a y be perforate,
n a r r o w l y perforate ,
rimate o r
IO
ZOOLOGISCHE V E R H A N D E L I N G EN
l68 (l979)
Figs. 1-4. The umbilicus is wide (1), narrow (2), rimate (3) or imperforate ( 4 ) .
Figs. 5-7. The whorls are flat (5), slightly convex (6) or convex (7).
BREURE, BULIMULINAE
11
F i g s . 8-12. T h e aperture is ovate (8), subovate (9), elongate-ovate (10), subcircular (11) or obliquely truncate-ovate (12).
F i g s . 13-16. T h e peristome is simple (13), n a r r o w l y expanded (14), expanded (15) r e f l e x e d (16).
or
12
ZOOLOGISCHE V E R H A N D E L I N G E N 168 (1979)
F i g s . 17-25. D i f f e r e n t types o f central radula teeth; respectively C - i , C-2, C-3, C-4, C-5, C-6, C-7, C-8 and C - 9 (see B r e u r e , 1978b, f o r descriptions).
F i g s . 26-31. D i f f e r e n t types o f lateral radula teeth; respectively L - i , L - 2 , L - 3 , L - 4 , L - 5 and L - 6 (see Breure , 1978b, f o r descriptions).
BREURE, BULIMULINAE
13
F i g . 32-48. D i f f e r e n t types o f lateromarginal radula teeth; respectively L M - i , L M - 2 , L M - 3 , L M - 4 , L M - 5 , L M - 6 , L M - 7 , L M - 8 , L M - o (2x), L M - 1 0 , L M - 1 1 , L M - 1 2 , L M - 1 3 , L M - 1 4 , L M - 1 5 and L M - 1 6 (see B r e u r e , 1978b, f o r descriptions).
BREURE, BULIMULINAE
15
F i g s . 55-60. P a l l i a i organs. F i g . 55. Plectostylus coquimbensis ( B r o d e r i p ) . F i g . 56. Bostryx modestus ( B r o d e r i p ) . F i g . 57. Stenostylus zilchi W e y r a u c h . F i g . 58. Scutalus (Kuschelenia) culmineus ( d O r b i g n y ) . F i g . 59. Bostryx virgultorum ( M o r e l e t ) . F i g . 60. Plekocheilus (Aeropictus) dissimulans ( P r e s t o n ) . Scale — 5 m m .
16
ZOOLOGISCHE V E R H A N D E L I N G E N 168 ( 1 9 7 9 )
imperforate
(figs.
1-4).
T h e protoconch
m a y be s m o o t h ,
granulate,
or
s c u l p t u r e d w i t h a x i a l w r i n k l e s , a x i a l riblets, s p i r a l lines, o r is g r a t e d o r p i t reticulate (pis. 1-3). T h e w h o r l s a r e slightly c o n v e x , c o n v e x o r flat 5-7). T h e aperture is, e.g., subovate, ovate, elongate-ovate, cate-ovate
(figs.
(figs.
subcircular, t r u n -
8 - 1 2 ) . T h e peristome is simple, n a r r o w l y e x p a n d e d , e x -
p a n d e d o r r e f l e x e d (figs. 13-16). I n side v i e w the peristome m a y be parallel to the l e n g t h a x i s o f the shell o r f o r m i n g a n angle
('skewed')
o f ca. 45
degrees at the utmost.
TABLE 2 V a r i a t i o n i n palliai o r g a ns i n B u l i m u l i n a e A d r e c t a l u r e t e r p a r t l y open
Veins weakly to moderately developed
Bostryx, Scutalus Neopetraeus, Leiostracus, Simpulopsis
Veins strongly developed
closed
Bostryx, Thaumastus (Thaumastiella), Bulimulus, Rabdotus, Scutalus (Scutalus), S. (Suniellus), Drymaeus, Newboldius, Leiostracus (Pseudoxychona)
Bostryx, Scutalus (Vermiculatus), Berendtia, Drymaeus, Plectostylus
Veins strongly developed, anteriorly deltoid ramified + parallel veins
Adrectal ureter
Plekocheilus, Thaumastus (Thaumastus), T. (Paeniscutalus), T. (Scholvienia), Bostryx, Bulimulus, Scutalus (Scutalus), Bothriembryon (Bothriembryon), Otostomus, Stenostylus, Auris
Scutalus (Vermiculatus), S. (Kuschelenia), Discoleus
The
Bostryx, Scutalus lenia), Discoleus, Neopetraeus
Naesiotus, (Suniellus), Oxychona, Rhinus,
(KuscheDrymaeus,
structure o f the r a d u l a has b e e n d e s c r i b ed b y B r e u r e (1978b ) a n d
B r e u r e & E s k e n s ( i n p r e p a r a t i o n ) . S e e also figs. 17-48. T w o , m o r e o r less independent, t r e n d s m a y be observed, v i z . ( 1 ) the transverse r o w s v a r y f r o m straight ( i n most species; R R 1) to V - s h a p e d ( i n a l i m i t e d n u m b e r o f species; RR
2)
a n d ( 2 ) the ( c e n t r a l )
teeth
are tricuspid (normally;
R T
1) o r
m o n o c u s p i d ( i n some g r o u p s ; R T 2 ) . S o m e authors, e.g. S o l e m ( 1 9 7 4 ) , hav e discussed the e v o l u t i o n a r y t r e n d s i n the structure o f the r a d u l a a n d they agree
that
the o c c u r r e n c e o f V - s h a p e d transverse
rows
a n d monocuspid
( c e n t r a l ) teeth is rather exceptiona l ( a n d m a y be r e g a r d e d as a p o m o r p h o u s character states; see part V I I ) . I n m o s t g r o u p s the teeth i n the c e n t r a l p a r t
BREURE, BULIMULINAE
17
F i g s . 61-64. Schematic reconstruction o f the penial complex. F i g . 61. Bulimulus guadalupensis (Bruguière). F i g . 62. Rabdotus mooreanus ( P f e i f f e r ) . F i g . 63. Bostryx ignobilis ( P h i l i p p i ) . F i g . 64. Bostryx tumidulus ( P f e i f f e r ) . o f the r a d u l a have ' s u p p o r t i n g denticles' ( R S 2; see S o l e m , 1972, f o r details o n f u n c t i o n i n g ) . I n some genera, however , this structure is absent ( R S
1)
[see p i . 3 ] . The
v a r i a t i o n i n the
palliai o r g a n s
mainly
c o n c e r n the
shape
of
the
n e p h r i d i u m , the l e n g t h o f the p e r i c a r d , the d e v e l o p m e nt o f the veins a n d the structure o f the adrectal ureter. T h e v a r i a t i o n is s h o w n i n figs. 4 9 - 6 0 a n d s u m m a r i z e d i n table 2. C o m p a r e d w i t h m o s t other S t y l o m m a t o p h o r a the genitalia * ) o f idae are relatively
simple. T h e r e a r e n o appendages
Bulimul-
(at least not i n the
B u l i m u l i n a e ) a n d the e x t e r n a l m o r p h o l o g i c a l v a r i a t i o n is l i m i t e d to d i f f e r ences i n relative l e n g t h a n d thickness. T h e i n t e r n a l m o r p h o l o g y a n d histology of
the genitalia
(especially the p e n i s ) , however ,
offer
a useful additional
variation. 1) T e r m i n o l o g y i n accordance w i t h B a y n e (1973). 2
18
ZOOLOGISCHE V E R H A N D E L I N G E N
The
168
penis m a y be w i t h a p r o x i m a l sheath ( P S
(1979)
i ) , o f v a r i a b l e length, o r
w i t h o u t s u c h a sheath ( P S 2 ) . T h e sheath m a y cove r 1/3-1/15 o f the l e n g t h o f the phallus ( =
penis +
epiphallus +
f l a g e l l u m ) . I n Discoleus
has a 'pseudo-sheath/ o v e r its entire l e n g t h
(PW
the penis
2 ) . I n t e r n a l l y the penis
l u m e n m a y be simple ( P L 1), constricted i n its m e d i a n part ( P L 2 ) o r w i t h
F i g s . 65-66. Schematic reconstruction o f penial complex. F i g . 65. Bothriembryon (Bothriembryon) indutus ( M e n k e ) . F i g . 66. Plectostylus coquimbensis ( B r o d e r i p ) . tubes parallel to the m a i n l u m e n / p o u c h e s / p a r a l l e l tubes / c i r c u l a r g l a n d ( P L 3; figs. 6 1 - 7 0 ) . I n several g r o u p s the subepithelial tissue i n the distal part o f the p e n is is m a d e u p o f large, r o u n d e d ( g l a n d u l a r ) cells ( P D 2 ) . T h e epithelium
i n the
penis
g l a n d u l a r cells
(P
is m a d e
up by
one
(Pi)
or two
different
types
of
2 ) . D e t a i l s o n these character states m a y be f o u n d i n
Breure (1978b). The
epiphallus a n d flagellu m are rather constant
i n both external a n d
internal m o r p h o l o g y ; i n several g r o u p s the epiphallus intrudes the distal p a r t of
the penis
( E P 2 ) , w h i l e i n some species the flagellu m is embedde d i n
F i g s . 67-70. Schematic reconstruction (67) a n d dissection o f penial c o m p l e x . F i g . 67. Plekocheilus ( P f e i f f e r ) . F i g . 68. Plekocheilus (Eurytus) elaeodes ( P f e i f f e r ) . F i g . 69. Plekocheilus (Aeropictus) cheilus (Plekocheilus) aurissileni ( B o r n ) .
(Plekocheilus) blainvilleanus loveni delicatus ( P i l s b r y ) . F i g . 70. P l e k o -
w
M VO
W
g s
M W
s
21
BREURE, BULIMULINAE
T h e spermathecal duct is u s u a l l y m o r e o r less s u b c y l i n d r i c a l a n d as l o n g as the s p e r m o v i d u c t
(SD
i ) . I n some g r o u p s this duct is r e d u c e d i n l e n g t h
( S D 2 ) . I n other g r o u p s the p r o x i m a l part o f the duct is relatively
stout,
w i t h a spermathecal a p p e n d i x ( S A 2) a n d a n a r r o w distal p a r t o f the duct. The
vagina,
external
oviduct
and
spermoviduct
a n d interna l m o r p h o l o g y .
are all rather constant
in
both
T h e r e is o n l y one g e n u s i n w h i c h
g l a n d u l a r folds are a r r a n g e d parallel to the l e n g t h o f the s p e r m o v i d u c t
the
(Si);
i n all other genera the g l a n d u l a r folds are p e r p e n d i c u l a r to the length
axis
( S 2 ) . T h e difference s that p r o b a b l y do exist i n the i n t e r n a l m o r p h o l o g y a n d histology
of
the s p e r m o v i d u c t
can o n l y be studied w i t h specialized
histo-
chemical techniques, f o r w h i c h o u r materia l was not suitable. T h i s p a p er o n l y presents figures o f the genitalia o f species o f Auris Leiostracus
(Pseudoxychona),
and
as these t a x a were not treated i n m y p r e v i o u s
papers. D e t a i l s o n the a n a t o m y o f species o f other genera m a y be f o u n d i n B r e u r e (1975e, i 9 7 6 d , 1977a, 1978b) a n d B r e u r e & C o p p o i s ( 1 9 7 8 ) . T h e v a r i a t i o n o f the d i f f e r e n t character states is s u m m a r i z e d i n T a b l e 3, as far as data are available. V.
ACKNOWLEDGEMENTS
T h e facilities o f the R i j k s m u s e u m v a n N a t u u r l i j k e H i s t o r i e w e r e put at m y disposal b y
P r o f . dr. W .
Vervoort,
for which I am very
grateful.
I
am
indebted to D r . E . G i t t e n b e r g e r f o r his m u l t i p l e advices. F o r technical assistance w i t h m y research I a m obliged to M r . S . P l o e g e r o f o u r D i v i s i o n a n d to M r . R . V r o o m o f the L e i d e n m u s e u m . M e s s r s . A . A . C . E s k e n s a n d J . R . S c h o u t e n studied some b u l i m u l i d g r o u p s u n d e r m y s u p e r v i s i o n , as p a r t o f their w o r k r e q u i r e d to obtain a doctorate degree ( D r s . ) at the U n i v e r s i t y o f L e i d e n . F o r assistance w i t h the photograph s a n d d r a w i n g s I w i s h to t h a n k M e s s r s . H . H e i j n a n d A . 't H o o f t . T h e s c a n n i n g electron m i c r o g r a p h s w e r e t a k e n at the G e o l o g i s c h - M i n e r a l o g i s c h Instituut
(University of Leiden)
by
M r . S . P l o e g e r a n d the author, w i t h the exper t assistance o f M e s s r s . F . v a n Sandijk a n d W . C . L a u r i j s s e n . F o r their assistance w i t h m y w i s h to than k F e l i p e a n d A u r a m a r i a G e r e n a
(Bogotá)
field w o r k
and M r . A .
I
Pena
( L i m a ) . P r o f . d r . J . T h . W i e b e s , head o f the D i v i s i o n o f Systematics
and
E v o l u t i o n a r y B i o l o g y , s u p e r v i s e d m y research. T h e f o l l o w i n g persons k i n d l y lent m e materia l o r s u p p l i e d m e w i t h data f r o m the collections u n d e r their charge: D r . Â . A n d e r s s o n ( S t o c k h o l m ) , D r . J . L . B . A r a u j o ( R i o de J a n e i r o ) , D r s . E . B i n d e r a n d C . V a u c h e r Dr.
K . Boss
van Goethem Kilias
(Geneva),
(Cambridge, Mass.), D r . C. C. Christensen ( T u c s o n ) , D r . J . ( B r u s s e l s ) , M r s . D r . M . I. H y l t o n Scott ( L a P l a t a ) , D r . R .
(Berlin), D r . P.
Mordan
a n d his staff
(London), Drs. H . E .
B.
22
ZOOLOGISCHE V E R H A N D E L I N G E N 168 (1979 )
Rezende
a n d A . C . dos Santos
Coelho
( R i o de J a n e i r o ) , D r . A . S o l e m
(Chicago), D r . F . G . T h o m p s o n (Gainesville), M r . S. Tillier ( P a r i s ) , D r s . A . Willink and Z . Tomsic (Tucumán), D r . B . R. W i l s o n (Perth) and D r s . A . Zilch and R . Janssen ( F r a n k f u r t a m M a i n ) . T h i s research was m a d e possible b y grants W WR
87-96,
W R 87-110 a n d
87-135 o f the F o u n d a t i o n f o r the A d v a n c e m e n t o f T r o p i c a l R e s e a r c h
( W O T R O ) , f o r w h i c h I want to express m y sincere gratitude.
VI. In
this p a r t a systematic
SYSTEMATICS
r e v i e w is p r e s e n t e d o f the B u l i m u l i n a e . E a c h
genus is b r i e f l y described a n d data a r e a d d e d o n its s y n o n y m y , d i s t r i b u t i o n a n d ecology. T h e m a i n publications i n w h i c h f u r t h e r data o n the g e n u s a n d its species m a y be f o u n d a r e also indicated. F o r each ( s u b ) g e n u s a list o f the t a x a b e l o n g i n g to that ( s u b ) g e n u s is g i v e n . I n this list the type localities, as g i v e n i n the o r i g i n a l p u b l i c a t i o n , a r e stated a n d data o n type m a t e r i a l a r e given,
i f available.
material seen;
(b)
C r i t e r i a to i n c l u d e t a x a (reputedly)
i n these lists
are:
(a)
c o r r e c t ly i d e n t i f i e d m a t e r i a l seen;
type
(c) o n
a u t h o r i t y o f another a u t h o r ; ( d ) o n account o f the o r i g i n a l d e s c r i p t i o n ; (e) on
account o f a r e d e s c r i p t i o n . A t the e n d o f this p a r t a list o f n o m i n a
i n q u i r e n d a is g i v e n , as well as some data o n fossil species a n d a list o f t a x a that are n o w e x c l u d e d f r o m the B u l i m u l i n a e .
K e y to the gener a o f B u l i m u l i n a e ia.
A u s t r a l i a n species
Bothriembryon
b.
N e o t r o p i c a l species
2
2a.
Protoconch smooth
3
b. 3a. b. 4a.
b.
Protoconch sculptured
b. 6a. b. 7a. b.
Bostryx
S h e l l large (height m o r e t h a n 4 0 m m )
4
S h e l l s u r f a c e granulate o r malleate; aperture elongate-ovate —
Colom-
bia, V e n e z u e l a
Dryptus
S h e l l s u r f a ce plicate, f o l d e d o r granulate; apertur e ( b r o a d l y ) ovate Brazil
5a.
5
S h e l l s m a l l (height less t h a n 4 0 m m )
P r o t o c o n c h granulate o r pit-reticulate Protoconch with axial a n d / o r spiral components P r o t o c o n c h granulate
—
Auris 6 10 Plekocheilus
P r o t o c o n c h pit-reticulate
7
P e r i s t o m e simple
8
Peristome expanded
Scutalus
BREURE, BULIMULINAE
8a. b. 9a. b.
23
S h e l l elongate-ovate, relatively large
Thaumastus
S h e l l globose, relatively s m a l l
Sphaeroconcha
A x i a l a n d s p i r a l component s o f p r o t o c o n c h sculpture equally s t r o n g
or present 10a. b. 11a. b. 12a. b. 13a.
16
A x i a l components c o n s i s t i n g o f w r i n k l e s
.
.
.
.
Simpulopsis
.
.
Newboldius
A x i a l c o m p o n e n t s c o n s i s t i n g o f straight riblets S h e l l s o l i d , large (ca. 70 m m h i g h ) .
.
.
11 .
S h e l l ( r a t h e r ) t h i n , small ( u p to ca. 50 m m h i g h )
12
S h e l l ( b r o a d l y ) conical
13
S h e l l m o r e o r less elongate-ovate
14
S h e l l b r o a d l y conical, i m p e r f o r a t e ; a p e r t u re skewed, triangular
b.
10
E i t h e r a x i a l o r s p i r a l c o m p o n e n t s o f p r o t o c o n c h sculptur e d o m i n a t i n g
Oxychona
S h e l l conical, r i m a t e ; aperture elongate-ovate to t r i a n g u l a r Cochlorina
14a.
b. 15a.
A p e r t u r e n a r r o w e d b y a callous flange at the i n n e r side o f
the
peristome
Otostomus
A p e r t u r e not n a r r o w e d b y a flange Inside
of
aperture
with
a pearly
15 lustre;
species
l i v i n g above
3000 m b. 16a.
Stenostylus
Inside o f aperture lustreless; species l i v i n g below 3000 m .
B o t h a x i a l a n d s p i r a l c o m p o n e n t s i n p r o t o c o n c h sculpture present a n d a x i a l ones d o m i n a t i n g
b. 17a. b.
Drymaeus
.
.
17
E i t h e r a x i a l o r s p i r a l components i n p r o t o c o n c h sculpture present
22
A x i a l components c o n s i s t i ng o f riblets
18
A x i a l components consisting o f w r i n k l e s
20
18a.
P r o t o c o n c h m o r e o r less a n g l e d above; peristome b r o a d l y e x p a n d e d
b.
P r o t o c o n c h r e g u l a r l y r o u n d e d ; peristome not to w e a k l y e x p a n d e d
19a.
Interstices o n p r o t o c o n c h as b r o a d as the riblets, s p i r a l striae n e a r l y
Neopetraeus
as s t r o n g as a x i a l riblets b.
Interstices o n p r o t o c o n c h 1-5
Llaucanianus times as b r o a d as the riblets, s p i r a l
lines relatively wea k 20a. b. 21 a.
Naesiotus
S h e l l ovate-conical; last w h o r l m o r e o r less keeled S h e l l elongate-ovate to globose;
last w h o r l r o u n d e d
Leiostracus .
.
.
.
21
Simpulopsis
L a s t w h o r l not p r o m i n e n t ; interstices o n p r o t o c o n c h b r o a d e r tha n the i r r e g u l a r l y spaced w r i n k l e s
22a.
.
L a s t w h o r l p r o m i n e n t ; interstices o n p r o t o c o n c h as b r o a d as wrinkles
b.
19
P r o t o c o n c h sculpture w i t h o n l y a x i a l c o m p o n e n ts
Bostryx 23
24
ZOOLOGISCHE V E R H A N D E L I N G E N 168 ( 1 9 7 9 )
b. 23a. b. 24a. b. 25a.
P r o t o c o n c h sculptur e w i t h o n l y s p i r a l c o m p o n e n t s
30
A x i a l components c o n s i s t i n g o f w r i n k l e s
24
A x i a l c o m p o n e n ts c o n s i s t i n g o f riblets
Rabdotus
Peristome broadly expanded and reflexed
25
P e r i s t o m e simple o r h a r d l y e x p a n d e d
26
A x i a l w r i n k l e s o n p r o t o c o n c h u n d u l a t i n g ; last w h o r l p r o m i n e n t Plekocheilus
b.
A x i a l w r i n k l e s o n p r o t o c o n c h zigz&g;
last w h o r l not p r o m i n e n t Rhinus
26a.
W r i n k l e s o n p r o t o c o n c h rather w e a k l y developed, r e g u l a r ; shell rather s m a l l (less tha n 50 m m )
b.
27
W r i n k l e s rather s t r o n g , r e g u l a r o r b r o k e n into o b l o n g , shorter w r i n k l e s ; shell rather large ( u s u a l l y m o r e t h a n 50 m m )
2?2l. b. 28a.
S h e l l u n i c o l o u r e d ( w h i t i s h to b r o w n ) ;
species
b. 31a. b. 32a. b.
.
.
l i v i n g below
29 500 m ,
Bulimulus
S h e l l u s u a l l y w i t h a c o l o u r pattern o f spots a n d / o r s p i r a l b a n d s ; 500 m i n the A n d e s
species Scutalus
C e n t r a l teeth o f r a d u l a m o n o c u s p i d ; species l i v i n g below ca. 500 m i n Plectostylus
C e n t r a l teeth o f r a d u l a t r i c u s p i d ; species l i v i n g above 500 m i n A n d e s of
30a.
Thaumastus 28
N o r t h Chile b.
.
S h e l l w i t h a x i a l streaks a n d / o r s p i r a l bands o r variegate d .
l i v i n g above 29a.
.
Shell uniformly coloured
m o s t l y outside the A n d e s b.
.
Argentina, Bolivia, Peru and Ecuador
Scutalus
P r o t o c o n c h w i t h n u m e r o u s s p i r a l lines
31
P r o t o c o n c h w i t h few s p i r a l lines
Lopesianus
C e n t r a l teeth o f r a d u l a t r i c u s p i d
32
C e n t r a l teeth o f r a d u l a m o n o c u s p i d
Leiostracus
P e n i s w i t h a sheath
Bostryx
P e n i s without a sheath
Plekocheilus G u i l d i n g ,
Discoleus
1828
Caprella G u i l d i n g , 1824: 341. T y p e species by m o n o t y p y : Caprella undulata G u i l d i n g [not Caprella L a m a r c k , 1801]. Plekocheilus G u i l d i n g , 1828a : 532. T y p e species by monotypy : Caprella undulata G u i l d i n g . Plecocheilus Swainson, 1833 : explanation p i . 103 [emendation]. Pelekocheilus Beck, 1837: 54. T y p e species by monotypy : Voluta aurissileni B o r n . Plecochilus A g a s s i z , 1846: 297 [emendation]. Pelecocheilus A l b e r s , 1850: 151 [emendation]. Pelecychilus A l b e r s , i 8 6 0 : 188. T y p e species b y subsequent designation ( P i l s b r y , 1896) : Voluta aurissileni B o r n . Pleocheilus M . E . G r a y , 1874 : p i . 74* f i g . 1. T y p e species by monotypy : Caprella undulata G u i l d i n g .
BREURE, BULIMULINAE
25
Plecocochilus Paetel, 1889: 207 [emendation]. Plechocheilus Leme, 1973: 307 [emendation]. Description. —
S h e l l (elongate-)ovate;
rimate to i m p e r f o r a t e ; t h i n to s o l i d .
C o l o u r w h i t i s h to b r o w n i s h , w i t h a x i a l z i g z a g streaks o r oblique s p i r a l series o f spots. S u r f a c e s m o o t h o r malleate, i n several species w i t h cuticula r cavities filled w i t h a i r . P r o t o c o n c h granulate o r w i t h a x i a l w r i n k l e s . W h o r l s slightly convex; to
suture h a r d l y to well i m p r e s s e d , d e s c e n d i n g i n front. A p e r t u r e s u b -
elongate-ovate.
Peristome
thickened
and
more
or
less
expanded
and
reflexed. C o l u m e l l a i n several species w i t h a f o l d . T h e central teeth o f the r a d u l a are m o n o c u s p i d , w i t h t r i a n g u l a r to ovate mesocones a n d h a r d l y developed ectocones. T h e l a t e r o m a r g i n a l teeth are
(1)
b i c u s p i d w i t h acute to truncate, spatulate mesocones a n d acute, ovate to deltoid ectocones;
o r (2)
b i c u s p i d , shifted, w i t h rather b l u n t spatulate to
elongate
mesocones a n d acute, t r i a n g u l a r to deltoid ectocones, w h i c h m a y be b i f i d i n blunt,
ovate
mesocones, acute elongate-ovate endocones a n d acute deltoid ectocones.
the
outermost
teeth;
Half-
row formula: C / i LMx/2
(x =
or
+
L
(3)
x/i
t r i c u s p i d , shifted,
+
M
y/2
(x =
with
1-6,
rather
y = 38-106) o r C / i
+
50-59)·
T h e p e r i c a r d , w h i c h is strongl y t r a n s v e r s a l l y disposed, is as l o n g as the nephridium,
which
is
broadly
triangular.
The
main
pulmonary
vein
is
p r o m i n e n t , the side veins are s t r o n g ly developed, especially at the a n t e r i o r end
w h e re
the veins
are r a m i f i e d a n d where
one o r two
short veins
are
parallel to the m a i n p u l m o n a r y v e i n . T h e adrectal ureter is closed. P e n i s w i t h a p r o x i m a l penis sheath ( i n Eurytus ( i n Plekocheilus
s.str., Eurytus
a n d Eudolichotis).
s u b c y l i n d r i c a l , i n several species epiphallus
without
external
a n d Aeropictus)
p r o x i m a l l y s w o l l e n , a n d p a s s i n g into the
differentiation.
The
flagellum
is slender
rather l o n g , but i n some species it is short a n d stout (especially i n and
Eudolichotis).
and
Aeropictus
T h e v a g i n a is relatively short. T h e spermathecal duct is
m o r e o r less t a p e r i n g , w i t h a n elongate-globose Distribution. —
o r withou t
T h e penis is m o r e o r less
spermatheca at the distal e n d .
W e s t Indies, V e n e z u e l a , B r a z i l , B o l i v i a , P e r u , E c u a d o r ,
Colombia, Panama. Relationships. —
See page 147 f o r a d i s c u s s i o n o f the phylogenetic r e l a t i o n -
ships o f this genus. Remarks. —
T h e d i f f e r e n c e s between the s u b g e n e r a are but slight a n d the
d i v i s i o n o f Plekocheilus status o f Sparnotion Bibliography. —
into five subgenera is o n l y tentative. E s p e c i a l l y the
P i l s b r y , 1944, is u n c e r t a i n . T h e m a i n publications o n this genus are: B r e u r e , 1978b;
H a a s , 1955a; O b e r w i m m e r , 1931; r a u c h , 1967b.
P i l s b r y , 1895, 1939b; S o l e m , i 9 6 0 ;
Wey-
vi
VO
— ..
as . w
. Ä W
00
2
ON
BREURE, BULIMULINAE
F i g . 82. D i s t r i b u t i o n o f Plekocheilus.
27
ZOOLOGISCHE V E R H A N D E L I N G E N 168 (1979)
32 Remarks. —
J u s t i f i c a t i o n o f the s y n o n y m y m a y be f o u n d i n B r e u r e , 1977a.
T a x a . — T h e f o l l o w i n g t a x a are i n c l u d e d i n this subgenus: argenteus, Euritus, Jousseaume, 1900: 41, p i . 1 figs. 20-21 (Venezuela, M e r i d a , 4000 m ) . bulimoides, Succinea, P f e i f f e r , 1842b: 131 [indication, refers to Bulimus succinoides [sic] P e t i t ] . calliostoma, Bulimus (Eurytus), D o h r n , 1882: 103, p i . 3 figs. 1-2 (província A n t i o q u i a N o v a e Granadae). cathcartiae, Bulimus, Reeve, 1848: p i . 42 f i g . 265 ( N e w Granada, P r o v . M e r i d a ) [ L T B M N H 1975288]. cleeforum, Plekocheilus (Aeropictus) succinoides [sic], B r e u r e , 1977a: 260, figs. 19-20 (Colombia, Dept. Cundinamarca, P á r a m o de Sumapaz, Cabeceras R i o Bogotá, A l t o de T o r q u i t a , ca. 3900 m ) [ H T Z M A ] . delicatus, Plectostylus, P i l s b r y , 1935: 84, p i . 6 figs. 6-8 ( C o l o m b i a, Soacha near Bogota) [ H T A N S P 164577a]. dissimulans, Bulimus (Eurytus), Preston, 1909: 509, p i . 10 f i g . 5 (Venezuela, M e r i d a ) [ L T B M N H 19144.3.37]. latilabris, Bulimus, P f e i f f e r , i855f : 7 (Santa F é de B o g o t a [ C o l o m b i a ] ) [ L T B M N H 1975127]. manco, Plekocheilus, P i l s b r y , 1930c: 356, p i . 31 f i g . 4 ( P e r u ) [ H T A N S P 152287]. quadricolor, Bulimus, P f e i f f e r , 1848a: 229 ( N e w Granada, province o f M e r i d a , C h a chopo) [ L T B M N H 1975283]. rhodocheilus, Bulimus, Reeve, 1848: p i . 28 f i g . 173 ( B r a z i l ) [ L T B M N H 1975129]. scytodes, Bulimus, P f e i f f e r , 1853b: 256 (Ande s o f C o l o m b i a ) . succineoides, Bulimus, Petit, 1840: 75 ( [ C o l o m b i a ] les environs de Bogota) [ S T M N H N ] . tenuissimus, Plekocheilus (Orcesiellus), W e y r a u c h , 1967b: 469, figs. 23, 50 (Ecuador, 20 k m a l oeste de Q u i t o , Tandayapa, 2500 m ) [ H T I M L 3364]. veranyi, Bulimus, P f e i f f e r , 1848a: 230 ( N e w Granada , province o f M e r i d a , Chachopo) [ L T B M N H 1975297]. zilchi, Plekocheilus (Aeropictus), Breure, 1977a: 260, figs. 2, 21-22 ( C o l o m b i a , Dept. Boyacá, S W L a b r a n z a Grande, Quebrada Cbmijoque) [ H T S M F 245387].
Plekocheilus (Sparnotion) P i l s b r y ,
1944
Sparnotion P i l s b r y , 1944c : 30. T y p e species by monotypy : Bulimus hauxwelli Crosse. Description. —
Shell f u s i f o r m ; narrowly perforate;
rather s o l i d . C o l o u r
y e l l o w i s h to light b r o w n . S u r f a c e w i t h some incrassate g r o w t h striae, i r r e gularly
spaced
Pgranulate. rowly
papillae
a n d cuticular
W h o r l s slightly
elongate-ovate,
convex;
cavities
filled with
air.
Protoconch
suture well i m p r e s s e d . A p e r t u r e n a r -
the basal m a r g i n p r o d u c e d . P e r i s t o m e
narrowly
re-
flexed. T h e a n a t o m y is u n k n o w n . Distribution. — Ecology. —
P e r u (Dept. Loreto).
T h e species lives i n t r o p i c a l r a i n forest, p r o b a b l y i n the leaf
litter layer. T h e o n l y k n o w n t a x o n is : hauxwelli, Bulimus, Crosse, 1872: 211 Peruviae) [ H T M C Z ] .
( i n v i c i n i o f l u m i n is A m b i y a c u , ad l o c u m Pebas,
36
ZOOLOGISCHE V E R H A N D E L I N G E N
168
(1979 )
F i g s . 83-84. V a r i a t i o n i n shell shape i n Auris. F i g . 83. A. bilabiata ( B r o d e r i p & S o w e r b y ) . F i g . 84. A. illheocola ( M o r i c a n d ) . Scale — 5 m m . F i g . 85. G e n i t a l i a of Auris bilabiata melanostoma ( M o r i c a n d ) ; after J u r b e r g , 1964.
BREURE, BULIMULINAE
F i g . 89. D i s t r i b u t i o n o f Thaumastus and Auris.
39
ZOOLOGISCHE V E R H A N D E L I N G E N 168 (1979)
40
lutea, Orphnus thompsoni, Cousin, 1887: 212 ( [ E c u a d o r ] Cuença). nigricans, Orphnus thompsoni, Cousin, 1887: 212 ( [ E c u a d o r ] C u e n ç a ). olivacea, Orphnus thompsoni, C o u s i n , 1887: 212 ( [ E c u a d o r ] Cuença). ortizianus, Plecocheilus (Eurytus), H a a s , 1955a: 366, f i g . 73 ( P e r u , valley o f R i o Chanay, between C h i c l a y o and C u t e r v o ) . thompsonii Bulimus, P f e i f f e r , 1845b: 74 ( [ E c u a d o r ] Q u i t o ) [ L T B M N H 1975464]. thompsonoides, Thaumastus, O b e r w i m m e r , 1931: 194, figs. 3, 6 ( E c u a d o r , L o j a ) [ H T S M F 5144]. viriatus, Bulimus, M o r e l e t , 1863: 170, p l . 7 f i g . 4 ( [ P e r u , Dept. Cuzco] vallée de Santa Anna, Niguapata). yanamensis, Bulimus, M o r e l e t , 1863: 171, p l . 8 f i g . 3 ( [ P e r u ] Y a n a m a ) [ L T B M N H 1975127]. zebra, Orphnus thompsoni, Cousin, 1887: 212 ( [ E c u a d o r ] près A z a g u e s ) .
Thaumastus (Scholvienia) Strebel,
1910
Scholvienia Strebel, 1910: 20. T y p e species by subsequent designation ( P i l s b r y , 1932a) : Bulimus bitaeniatus N y s t . Description. —
S h e l l elongate-ovate;
rimate ; rather solid. C o l o u r b r o w n
w i t h y e l l o w i s h s p i r a l b a n d ( s ) . S u r f a c e w i t h some incrassate g r o w t h
striae.
Protoconch
waved
(subexcavated)
with
riblets. W h o r l s slightly c o n v e x ;
strong
axial
wrinkles
and
short
suture w e l l i m p r e s s e d . A p e r t u r e
subovate.
P e r i s t o m e t h i n a n d simple. T h e central teeth o f the r a d u l a are t r i c u s p i d , w i t h acute, mesocones
and
hardly
developed
ectocones.
The
wedge-shape d
l a t e r o m a r g i n a l teeth
are
b i c u s p i d , w i t h rather b l u n t to acute, elongate to wedge-shape d mesocones a n d acute, elongate to deltoid ectocones; withou t s u p p o r t i n g denticles i n the outermost teeth. H a l f - r o w f o r m u l a : C / 3 + L M x / 2 ( x =
31-38).
T h e p e r i c a r d is t r a n s v e r s a l l y d i s p o s ed a n d is as l o n g as the n e p h r i d i u m , w h i c h is b r o a d l y t r i a n g u l a r . T h e t r a n s i t i o n o f the a d r e n a l a n d adrectal ureters is relatively
p o s t e r i o r l y situated.
T h e m a i n p u l m o n a r y v e i n is p r o m i n e n t ,
r a m i f i e d at the a n t e r i o r e n d , where a parallel v e i n is situated. T h e adrectal ureter is closed o v e r its entire length. T h e penis has a p r o x i m a l sheath, is rather thick a n d s u b c y l i n d r i c a l a n d is p a s s i n g without e x t e r n al d i f f e r e n t i a t i o n into the epiphallus. T h e flagellum is tapering. T h e v a g i n a is relatively short. T h e spermathecal duct is m o r e o r less subcylindrical,
its
distal p a r t t a p e r i n g
towards
the
elongate-globose
sper -
matheca. Distribution. —
P e r u (Depts. A p u r i m a c , Ayacucho, J u n i n , Pasco, Huánuco,
Cajamarca and Amazonas). Ecology. —
T h e species live i n o p e n montan e forest a n d steppe
vegetations,
i n the leaf litter layer. T h e vertical d i s t r i b u t i o n is ( 8 0 0 - ) 1800-3500 m . alutaceus, Bulimus, Reeve, 1850b: 99 ( P e r u , C u z c o ) [ L T B M N H 1975148]. argentinus, Thaumastus (Scholvienia), Bequaert, 1949: 114, p i . 7 f i g . 6 ( A r g e n t i n a , P r o v .
42
ZOOLOGISCHE V E R H A N D E L I N G E N 168 ( 1 9 7 9 )
T h e p e r i c a r d is t r a n s v e r s a l ly dispose d a n d is as l o n g as the n e p h r i d i u m , w h i c h is b r o a d l y t r i a n g u l a r . T h e m a i n p u l m o n a r y v e i n a n d the side v e i n s are moderately developed. T h e adrectal ureter is closed o v e r its entire length . P e n i s w i t h a sheath (ca. 1/6
the l e n g t h o f the p h a l l u s ) , slightly
swollen
above the distal e n d o f the sheath, but otherwise m o r e o r less s u b c y l i n d r i c a l a n d p a s s i n g withou t e x t e r n al d i f f e r e n t i a t i o n into the epiphallus. T h e flagellum is relatively short a n d t a p e r i n g . Distribution. — Ecology. —
P e r u (Depts. L a Libertad, Cajamarca).
T h e species live i n ' s a v a n n a h forest', u n d e r stones. T h e vertical
d i s t r i b u t i o n is ( 1200-) 1600-2750 m . T a x a . — T h e f o l l o w i n g t a x a are placed i n this subgenus: debilisculptus, Thaumastus (Thaumastiella) occidentalis, W e y r a u c h , 1960a: 30, p i . 3 f i g . 15 ( N - P e r u , bei L l a m a , an der Autostrasse v o n C h i c l a y o nach Cutervo, ca. 85 k m nö Chiclayo, 2000-2250 m ) [ H T S M F 162029]. glyptocephalus, Bulimulus, P i l s b r y , i 8 9 7 d : 21 ( P e r u ) [ H T A N S P 25675]. koepckei, Thaumastus (Scholvienia), Z i l c h , 1953: 53, figs. 7-9, p i . 14 f i g . 3 ( P e r u , [Dept. Cajamarca] H a c i e n d a Monteseco, ± 1200 m ) [ H T S M F 111487]. occidentalis, Thaumastus (Thaumastiella), W e y r a u c h , 1960a: 28, p i . 3 figs. 13-14 ( N - P e r u , U m g e b u n g v o n Còntumazá, n o k m nö T r u j i l l o , 2750 m ) [ H T S M F 162026]. sarcochrous, Bulimulus, P i l s b r y , i897d : 21 ( P e r u ) [ H T A N S P 4705].
Thaumastus (Paeniscutalus)
W u r t z , 1947
Paeniscutalus W u r t z , 1947: 12. T y p e species b y monotypy : Megalobulimus (Microborus) incarum P i l s b r y . Description. —
S h e l l ovate; rimate ; rather solid . C o l o u r u n i f o r m w h i t i s h
to light b r o w n . S u r f a c e w i t h m o r e o r less incrassate g r o w t h striae. P r o t o c o n c h w i t h indistinct a x i a l w r i n k l e s , w h i c h are p a r t l y b r o k e n into o b l o n g granules. W h o r l s slightly c o n v e x ;
suture crenulate, h a r d l y i m p r e s s e d. A p e r t u r e s u b -
ovate. P e r i s t o m e slightly t h i c k e n e d a n d n a r r o w l y e x p a n d e d . The
central teeth o f
the
r a d u l a are t r i c u s p i d , w i t h rather acute,
ovate
mesocones a n d elongate-truncate ectocones. L a t e r o m a r g i n a l teeth are b i c u s p i d , w i t h rather blunt to acute, elongate mesocones a n d elongate-truncate to ovate ectocones;
without
s u p p o r t i n g denticles
formula: C / 3 + L M x / 2 (x =
i n the outermost
teeth.
Half-row
38).
T h e p e r i c a r d is t r a n s v e r s a l l y disposed a n d is as l o n g as the n e p h r i d i u m , w h i c h is b r o a d l y t r i a n g u l a r . T h e m a i n p u l m o n a r y v e i n a n d the side v e i n s are p r o m i n e n t ; the veins are r a m i f i e d at the a n t e r i o r e n d , w h e r e a v e i n parallel to the m a i n p u l m o n a r y v e i n is situated. T h e adrectal ureter is closed o v e r its entire length. Penis
with
a
sheath,
more
or
less s u b c y l i n d r i c a l a n d p a s s i n g
without
external d i f f e r e n t i a t i o n into the epiphallus. T h e flagellu m is rather short a n d
43
BREURE, BULIMULINAE
relatively thick. T h e spermathecal duct is thick a n d about h a l f as l o n g as the s p e r m o v i d u c t ; the spermatheca is Distribution. — Ecology. —
elongate-globose.
P e r u (Depts. A n c a s h , L a L i b e r t a d ) .
T h e species lives near cultivated g r o u n d s , u n d e r stones.
The
vertical d i s t r i b u t i o n is 1850-3300 m . Remarks. — mulus cutalus
T h i s t a x o n was o r i g i n a l l y described as a subgenus o f
L e a c h , 1814.
Zilch
(i960)
and Parodiz
(1962) c o n s i d e r e d
Buli-
Paenis-
as a separate genus, but W e y r a u c h ( i n I M L - c o l l e c t i o n ) suggested its
classification as a subgenus o f Thaumastus
A l b e r s , i 8 6 0 ; I agree w i t h this
suggestion. T a x a . — T h e f o l l o w i n g t a x a are i n c l u d e d i n
Paeniscutalus:
crenellus, Bulimus, P h i l i p p i , 1867: 67 ( P e r u [Dept. L a L i b e r t a d , D i s t r . Santiago de Chuco] hacienda de U n i g a m b a l ) . incarum, Megalobulimus (Microborus), P i l s b r y , 1944c: 29, p i . 1 figs. 8-9 ( P e r u , [Dept. A n c a s h ] H u a r a z , 3000-3200 m ) [ H T A N S P 180677a].
Thaumastus (Thaumastus) Description. —
S h e l l elongate-ovate;
Albers, i860
imperforate;
solid. C o l o u r light
d a r k b r o w n , m o s t l y w i t h d a r k e r a x i a l streaks o r light s p i r a l b a n d ( s ) . w i t h incrassate
growth
W h o r l s hardly convex;
striae. P r o t o c o n c h w i t h
fine, close a x i a l
to
Surface wrinkles.
suture w e l l i m p r e s s e d, m o r e o r less crenulate. A p e r -
ture relatively small, subovate. P e r i s t o m e slightly e x p a n d e d . The
central teeth
of
the r a d u l a are t r i c u s p i d , w i t h rather acute,
mesocones a n d elongate-truncate
ovate
ectocones. T h e l a t e r o m a r g i n a l teeth are b i -
c u s p i d , w i t h rather blunt, elongate mesocones, r u d i m e n t a r y w i n g - l i k e
endo-
cones a n d elongate-truncate
meso-
ectocones
o r w i t h acute, rather elongate
a n d ectocones; without s u p p o r t i n g denticles i n the outermost teeth. H a l f - r o w f o r m u l a : C / 3 4- L M x / 2 ( x =
32-43).
T h e p e r i c a r d is transversall y d i s p o s e d a n d is about as l o n g as the n e p h r i d i u m , w h i c h is b r o a d l y t r i a n g u l a r . T h e t r a n s i t i o n o f a d r e n a l a n d adrectal ureters is relatively p o s t e r i o r l y situated. T h e m a i n p u l m o n a r y v e i n is p r o m inent a n d r a m i f i e d at the a n t e r i o r e n d , wher e a parallel v e i n is situated. T h e adrectal ureter is closed o v e r its entire length. T h e penis has a p r o x i m a l sheath a n d is rather thick a n d m o r e o r less s u b c y l i n d r i c a l ; p a s s i n g without external d i f f e r e n t i a t i o n into the epiphallus. T h e flagellum
is s u b c y l i n d r i c a l to t a p e r i n g , u p to h a l f as l o n g as the
phallus
length. T h e v a g i n a is relatively short. T h e spermathecal duct is s u b c y l i n d r i c a l , w i t h a n elongate-truncate to globose spermatheca at the distal e n d . Distribution. — Ecology. —
Brazil, Bolivia, Peru, Ecuador.
I n P e r u the species live i n c l o u d a n d montan e forests, m a i n l y
46
ZOOLOGISCHE V E R H A N D E L I N G E N 168
(1979)
the u p p e r w h o r l s paler. S u r f a c e w i t h some incrassate g r o w t h striae. P r o t o conch with convex;
fine
axial
suture
well
wrinkles impressed.
a n d short w a v e d Aperture
riblets. W h o r l s
sub-ovate.
Peristome
slightly
thin
and
simple. [ A f t e r S t r e b e l ] . Distribution. — Ecology. —
P e r u (Dept. J u n i n ) .
Unknown.
Remarks. —
O n l y the type s p e c i m e n was
k n o w n a n d this is
probably
destroyed, together w i t h m o s t o f S t r e b e r s types, d u r i n g the 1939-1945 w a r ( D a n c e , 1966: 3 0 2 ) . J u d g i n g f r o m the d e s c r i p t i on this t a x o n is mos t p r o b a b l y a s y n o n y m o f Scholvienia
S t r e b e l, 1910 (page 4 0 ) .
T h e sole t a x o n i n this g e n u s is: claritae, Thomsenia, Strebel, 1910: 27, p i . 2 f i g . 16 ( P e r u , Chanchamayo) . N o m e n inquirendum
Lopesianus W e y r a u c h , Lopesianus W e y r a u c h , Weyrauch. Description. —
1958:
120. T y p e
1958
species by m o n o t y p y :
S h e l l elongate-ovate;
narrowly
Lopesianus crenulatus
perforate;
thin.
Colour
u n i f o r m b r o w n i s h . S u r f a c e w i t h slightly incrassate g r o w t h striae. P r o t o c o n c h w i t h relatively
few,
indistinct s p i r a l lines. W h o r l s slightly c o n v e x ;
weakly crenulate, well i m p r e s s e d . A p e r t u r e elongate-ovate.
suture
Peristome thin and
simple. Distribution. —
Brazil.
E c o l o g y . — U n k n o w n . T h e species lives at sea level. Relationships.
—
The
sculpture
relationship w i t h Bostryx,
of
the
protoconch
o r possibl y Leiostracus,
suggests
a
remote
but n o t h i n g c a n be said
u n t i l the a n a t o m y is k n o w n . Remarks. —
O n l y the type m a t e r i al o f Lopesianus
crenulatus
Weyrauch
is available a n d , despite all efforts, n o f u r t h e r specimens have been f o u n d at the type locality ( R e z e n d e & A r a u j o , p e r s o n a l c o m m u n i c a t i o n ) . T h e sole t a x o n i n this genus is: crenulatus, Lopesianus, W e y r a u c h , 1958: 121, p l . 6 figs. Arrairal-Praínha, E s t a d o do R i o ) [ H T S M F 156376].
Bostryx T r o s c h e l , Bostryx T r o s c h e l , Troschel.
1847:
49. T y p e
7-8
( B r a s i l i e n , Cabo
Frio,
1847
species by m o n o t y p y :
Bulimus
(Bostryx)
solutus
Peronaeus A l b e r s , 1850: 163. T y p e species b y subsequent designation ( A l b e r s , i860) : Bulinus pupiformis B r o d e r i p . Ataxus A l b e r s , 1850: 164. T y p e species by m o n o t y p y : Bulimus umbilicaris Souleyet. Pyrgus A l b e r s , 1850: 177. T y p e species b y monotypy : Bulinus turritus B r o d e r i p .
F i g s . 90-109. V a r i a t i o n i n shell shape i n Bostryx. F i g . 90. B. bermudezae W e y r a u c h . F i g . 91. B. elatus ( P h i l i p p i ) . F i g . 92. B. hamiltoni (Reeve). F i g . 93. B. obeliscus Z i l c h . F i g . 94. B. arcuatus B r e u r e . F i g . 95. B. tschudii ( T r o s c h e l ) . F i g . 96. B. pustulosus ( B r o d e r i p ) . F i g . 97. B. obliquiportus W e y r a u c h . F i g . 98. B. tubulatus scalaricostus ( M o r e l e t ) . F i g . 99. B. infundibulum infundibulum ( P f e i f f e r ) . F i g . 100. B. chagualensis P i l s b r y . F i g . 101. B. scalriformis ( B r o d e r i p ) . F i g . 102. B. solutus ( T r o s c h e l ) . F i g . 103. B. frederici Breure . F i g . 104. B. rhodolarynx apurimacensis ( D a l i ) . F i g . 105. B. megomphalus P i l s b r y . F i g . 106. B. conspersus ( S o w e r b y ) . F i g . 107. B. erythrostomus ( S o w e r b y ) . F i g . 108. B. reentsi ( P h i l i p p i ) . F i g . 109. B. planissimus P i l s b r y & Olsson. Scale = 5 m m .
48
ZOOLOGISCHE V E R H A N D E L I N G E N 168 ( 1979)
F i g . n o . D i s t r i b u t i o n of Bostryx sensu lato.
49
BREURE, BULIMULINAE
Geopyrgus P i l s b r y , 1896a: 114. N e w name f o r Pyrgus A l b e r s , 1850 not Hübner, 1816. Lissoacme P i l s b r y , 1896a: 114. T y p e species by o r i g i n a l designation: Bulinus erythrostoma Sowerby. Platybostryx P i l s b r y , 1896b: 129. T y p e species b y m o n o t y p y : Bulimulus eremothauma Pilsbry. Geoceras P i l s b r y , 1896b : 136. T y p e species by o r i g i n a l designation : Bulimus columellaris Reeve. Dentaxis P i l s b r y , 1902a : x x x i . T y p e species by monotypy : Bulimulus dentaxis P i l s b r y . Phenacotaxus D a l l , 1912a: 7. T y p e species by o r i g i n a l designation: Bulimulus infundibulum umbilicatellus P i l s b r y . Ataxellus D a l l , 1912a: 7. T y p e species by o r i g i n a l designation: Phenacotaxus (Ataxellus) spiculatus pectinatus D a l l . Scansicohlea P i l s b r y , 1930c : 358. T y p e species by o r i g i n a l designation : Bulimulus (Scansicohlea) bromeliarum P i l s b r y . Scansicochlea T h i e l e, 1931: 656. T y p e species by m o n o t y p y : Bulimulus (Scansicohlea) bromeliarum P i l s b r y . Discobostryx P i l s b r y & O l s s o n , 1949: 11. T y p e species by m o n o t y p y : Bostryx (Discobostryx) planissimus P i l s b r y & Olsson. Vermetellus H a a s , 1951: 520. T y p e species by m o n o t y p y : Bostryx (Vermetellus) metagyra P i l s b r y & Olsson . Pseudoperonaeus W e y r a u c h , 1958: i n . T y p e species b y m o n o t y p y : Bostryx (Pseudoperonaeus) bermudezae W e y r a u c h . Elatibostryx W e y r a u c h , 1958: 112. T y p e species by o r i g i n a l designation '.Bostryx (Elatibostryx) imeldae W e y r a u c h . Pampasinus W e y r a u c h , 1958: 113. T y p e species by o r i g i n a l designation : Bostryx weyrauchi P i l s b r y . Multifasciatus W e y r a u c h , 1958: 116. T y p e species b y o r i g i n a l designation : Bulimus subroseus P f e i f f e r . Bilamelliferus W e y r a u c h , 1958: 118. T y p e species by m o n o t y p y : Bulimus tschudii Troschel. Kionoptyx H a a s , 1966: 239. T y p e species by o r i g i n a l designation : Kionoptyx sagasteguii Haas. Naesiotellus W e y r a u c h , 1967a : 414. T y p e species b y monotypy : Naesiotus (Naesiotellus) late columellaris W e y r a u c h . Floreziellus W e y r a u c h , 1967b: 488. T y p e species by m o n o t y p y : Floreziellus florezi Weyrauch. Description. — discoid;
Shell
( d e p r e s s e d - ) c o n i c a l to
elongate-ovate,
globose
or
( n a r r o w l y ) p e r f o r a t e to r i m a t e ; rather t h i n . C o l o u r w h i t i s h , b r o w n -
ish o r b l u i s h , u n i f o r m o r w i t h d a r k e r s p i r a l lines a n d / o r spots. P r o t o c o n c h w i t h n u m e r o u s , fine s p i r a l lines, sometimes s m o o t h o r w i t h indistinc t a x i a l w r i n k l e s o r riblets ( n e v e r as s t r o n g a n d r e g u l a r as i n Naesiotus
species).
W h o r l s slightly c o n v e x ( i n some species the last w h o r l is k e e l e d ) ; suture w e l l impressed. A p e r t u r e ( s u b ) o v a t e to t r i a n g u l a r - o v a t e ( i n some species a d n a t e ) . P e r i s t o m e simpl e o r slightly e x p a n d e d , i n some species continuous. C o l u m e l l a u s u a l l y simple, sometimes w i t h a l a m e l la w i t h i n the last w h o r l s . Central
teeth
of
the
r a d u l a are
tricuspid
(mesocones
lanceolate
and
ectocones ovate to deltoid) o r m o n o c u s p i d ( w i t h blunt, deltoid c o n e s ) . L a t e r o m a r g i n a l teeth b i c u s p i d ( w i t h elongate to lanceolate mesocones a n d ovate to 4
50
ZOOLOGISCHE V E R H A N D E L I N G E N 168 ( 1 9 7 9 )
deltoid ectocones, w h i c h m a y be serrate i n the outermost teeth) o r m o n o cuspid (x =
(with
blunt, conical c o n e s ) .
Half-row
formula:
16-31) o r C / i + L x / i + M y / 2 ( x = 6-19, y =
C/3
+
L M
x/2
13-31).
P e r i c a r d h a l f as l o n g as to as l o n g as the n e p h r i d i u m , w h i c h is ( n a r r o w l y ) t r i a n g u l a r . T h e m a i n p u l m o n a r y v e i n is m o d e r a t e l y to w e l l developed, side veins are w e a k l y to w e l l developed. T h e adrectal u r e t e r is closed o r p a r t i a l l y o p e n ( 1 / 1 0 - 1 / 3 o f its l e n g t h ) . P e n i s u s u a l l y w i t h a p r o x i m a l sheath. T h e l u m e n o f the penis is l i n e d w i t h two types o f e p i t h e l i u m a n d is rather n a r r o w , especially at the t r a n s i t i o n to the distal p a r t o f the penis. I n the distal p a r t a c i r c u l a r g l a n d is present, w h i c h is e x t e r n a l l y
visible as a swelling. T h e t r a n s i t i o n to the epiphallus, both
i n t e r n a l l y a n d externally, is g r a d u a l . T h e flagellum is slender, w i t h a distally attached retractor muscle. T h e spermathecal duct is m o r e o r less s u b c y l i n d r i c a l . T h e spermatheca is globose. Distribution. —
Venezuela ( ? ) , N - A r g e n t i n a , Bolivia, Chile, P e r u , E c u a -
dor. Ecology. —
T h e species u s u a l l y live o n the g r o u n d i n leaf litter o r i n
s h r u b s , w e r e they are g e n e r a l l y
f o u n d i n d o r m a n c y . S o m e species live o n
rock-faces. T h e v e r t i c al d i s t r i b u t i o n is 0-3600 m . Relationships. — o n page
Remarks. — of
T h e phylogenetic relationships o f this g e n u s are discussed
151. It has b e e n m e n t i o n e d b e f o r e ( B r e u r e , 1978b) that a n u m b e r
species g r o u p s m a y be r e c o g n i z e d w i t h i n Bostryx
r e s p o n d m o r e o r less
with
some
of
the
( s e n s u lato) that c o r -
'subgenera'
listed i n the
above-
m e n t i o n e d s y n o n y m y . T h e r e are, however, a rather large n u m b e r o f t a x a that can not be allocated to one o f these species g r o u p s a n d it is p r e f e r r e d , therefore, to treat the genus here s e n s u lato. Bibliography. — Dall,
1912a; H a a s ,
1896b, 1930C, 1932,
T h e m a i n publications o n this genus a r e : B r e u r e , 1978b; 1955b;
H y l t o n Scott,
1944b; R e h d e r , 1945;
1967b;
P a r o d i z , 1947;
W e y r a u c h , 1958,
Pilsbry,
1960a,
1960c,
1964, 1967a, 1967b. T a x a . — T h e f o l l o w i n g t a x a are p l a c e d i n this g e n u s :
abancayensis, Bostryx, P i l s b r y , 1944b: 123, p i . 11 f i g . 20 ( P e r u , A b a n c a y , 2300 m ) [ H T A N S P 180001]. acalles, Bulimus, P f e i f f e r , 1853b : 258 ( P e r u v i a n A n d e s ) . acme, Bulimulus (Peronaeus), H a a s , 1955b: 325, f i g . 68 ( P e r u , A p u r i m a c , Andahuaylas, O n g o y , a l t u r a de l a H a c i e n d a M o z o b a m b a) [ H T F M N H 51355]. acromelas, Bulimus, M o r e l e t , 1863: 202, p i . 11 f i g . 1 ( [ P e r u ] vallée d ' A y a c u c h o et dans celle de r U r u b a m b a ) . aequicostata, Peronaeus, Rehder, 1945: 5 ( N e w name f o r Bulimus scalarioides Pfeiffer, 1867, not Bulimus scalaroides Reeve, 1849).
BREURE, BULIMULINAE pericanus, Orthalicus, A d a m s
& A d a m s , 1855:
159
[emendation
57 f o r piuranus A l b e r s ,
1854]· peristomatus, Scutalus, Döring, 1879: 66 ( [ A r g e n t i n a ] S i e r r a de P o c h o , Quebr. de Y a t a n , de M e r m u l a , etc.). peruvianus, Drymaeus torallyi, P i l s b r y , 1944b: 126, p l . 11 f i g . 13 ( P e r u , Santa valley, H u a r a z , 3100 m ) [ H T A N S P 180008]. philippi, Bulimus, P f e i f f e r , 1842b: 120 [indication]. philippii, Peronaeus, Rehder, 1945: 3 ( C h i l e , near Copiapó) [ H T U S N M 537830]. pictus, Bulimus, P f e i f f e r , 1855a: 58 ( P e r u ) [ L T B M N H 1975545]. piuranus, Bulimus, A l b e r s , 1854a: 31 ( P e r u v i a septentrionali, prope oppidum P i u r a ) [ S T Z M B 10289]. placitus, Bostryx, Breure, 1978b: n i , figs. 153-154 ( P e r u , Dept. Huánuco, 7.6 k m S A m b o , 2360-2380 m ) [ H T R M N H 55207]. planissimus, Bostryx (Discobostryx), P i l s b r y & Olsson , 1949: 12, f i g . 13 ( P e r u ? ) [ H T A N S P 184269a]. platycheilus, Neopetraeus, H a a s , 1955b: 311, figs. 60-61 ( P e r u , A p u r i m a c Dept., A n d a huaylas P r o v . , H a c i e n d a P a l m i r a ) [ H T F M N H 51315]· ploegerorum, Bostryx, B r e u r e , 1978b: 115, figs. 163-166, p l . 2 f i g . 4 ( P e r u , Dept. A n c a s h , 5 k m S W C h a v i n de H u a n t a r , 3300 m ) [ H T U F 22791]. poveli, Bostryx huanucensis, Breure, 1978b: 115, f i g . 131, p l . 1 f i g . 5 ( P e r u , Dept. Pasco, 42.5 k m N N E C e r r o de Pasco, 2800 m ) [ H T R M N H 55109]. productus, Bulimus, P h i l i p p i , 1867: 77 ( [ P e r u ] S i e r r a Cotahuasi). pruinosus, Bulinus, Sowerby, 1833b: 36 ( [ C h i l e ] Coquimbo) [ S T B M N H ] . ptyalum, Bulimulus (Lissoacme), D a l l , 1910: 181, f i g . 3 ( P e r u , banks o f the R i o Pampas) [ H T U S N M 209271]. pumilio, Peronaeus, Rehder, 1945: 5 ( N e w name f o r Bulimus nanus Reeve, 1848, not L a m a r c k , 1804). pumilus, Bostryx spiculatus, B r e u r e , 1978b : 126, figs. 187-100 ( P e r u , Dept. Cuzco, near C a l c a on the left side o f the R i o V i l c a n o t a (ca. 2900 m ) ) [ H T I M L 1707a]. punctilineatus, Bulimulus (Scutalus), H a a s , 1951: 517, f i g . 105 ( P e r u , U r u b a m b a valley, Sahuayaco, 800 m ) [ H T F M N H 30914]. puntanus, Peronaeus (Lissoacme), P a r o d i z , 1947: 13, f i g . 8 ( A r g e n t i n a , prov. S a n L u i s , C e r r o de M o r r o ) [ H T M A C N 9917]. pupiformis, Bulinus, B r o d e r i p i n B r o d e r i p & Sowerby, 1832b: 105 ( C h i l i , H u a s c o ) . pustulosus, Bulinus, B r o d e r i p i n B r o d e r i p & Sowerby, 1832b: 105 ( C h i l i , H u a s c o ) . pygmaeus, Bostryx (Bostryx), W e y r a u c h , 1960c: 121, p l . 11 figs,
6o
ZOOLOGISCHE V E R H A N D E L I N G E N 168 (1979)
willinki, Bostryx (Bostryx), W e y r a u c h , 1964: 54, f i g . 12 ( A r g e n t i n a , P r o v . Catamarca, Quebrada de Tinogasta) [ H T I M L 121a]. zilchi, Bostryx (Bostryx), W e y r a u c h , 1958: 108, p l . 9 f i g . 41-42 ( M - P e r u , L a r a o s , 3600 m , i m T a l e des R i o Canete, 155 k m ösö L i m a [corrected to Quichao, 5 k m v o n L a r a o s ( W e y r a u c h , 1960c: 124)]) [ H T S M F 156348].
Bulimulus L e a c h ,
1814
Bulimulus L e a c h , 1814: 42. T y p e species by o r i g i n a l designation: Helix exilis G m e l i n . Siphalomphix Rafinesque, 1833: 165. T y p e species by monotypy : Siphalomphix bonariensis Rafinesque. Loboa Ihering, 1917: 121. T y p e species by monotypy : Loboa brunoi Ihering. Description. —
S h e l l ovate to o b l o n g ; n a r r o w l y p e r f o r a t e to rimate. C o l o u r
u n i f o r m l y b r o w n to y e l l o w i s h , sometimes w i t h bands o f d a r k e r ( r e d d i s h - ) b r o w n . S u r f a c e w i t h g r o w t h striae a n d o f t e n w i t h fine s p i r a l striae. P r o t o c o n c h w i t h a x i a l w r i n k l e s , sometimes w i t h g r a n u l a t i o n o r pit-reticulate o n its l o w e r part. A p e r t u r e
(sub)
ovate to squarish-ovate.
P e r i s t o m e simple, u n -
e x p a n d e d o r slightly e x p a n d e d . T h e central teeth o f the r a d u l a are t r i c u s p i d , w i t h lanceolate to wedgeshaped mesocones a n d ovate to deltoid ectocones. T h e central teeth are slightly smaller tha n the l a t e r o m a r g i n a l teeth, w h i c h are b i c u s p i d w i t h elongate to lanceolate mesocones a n d deltoid ectocones that m a y be serrate i n the outermost teeth. H a l f - r o w f o r m u l a : C / 3 + L M x / 2 ( x =
20-31).
P e r i c a r d ca. 3 / 4 the l e n g t h o f the n e p h r i d i u m , w h i c h is (elongate) t r i a n gular. T h e m a i n p u l m o n a r y v e i n is p r o m i n e n t a n d b r o a d . S i d e veins are well developed, especially at the a n t e r i o r e n d . T h e adrectal ureter is closed. T h e mantle collar is well developed. Penis
usually
swollen;
with
a p r o x i m a l sheath;
the
distal part
of
the
penis
the l u m e n o f the penis is d i v i d e d into several parallel tubes.
epiphallus, w h i c h internally deeply penetrates
is
The
into the penis, is slender a n d
about two times as l o n g as the penis. T h e flagellum is slender, w i t h a distally attached retractor muscle. T h e v a g i n a is relatively short. T h e m e d i a n part o f the spermathecal duct is w i d e n e d ; the duct tapers t o w a r d the distal e n d . T h e spermatheca is globose. Distribution. — Brazil,
Paraguay,
Antilles, Venezuela, Guyana, Surinam, F r e n c h Guyana, Uruguay,
Argentina,
Bolivia, Peru,
Colombia,
Central
America, Mexico. Ecology. —
T h e species are u s u a l l y g r o u n d - d w e l l i n g o r live i n s h r u b s , o n
stems a n d cacti. T h e y are generally c o n f i n e d to secondar y vegetation.
The
vertical d i s t r i b u t i o n is o-ca. 500(-ca. 800) m . Remarks. — (1979)
n
a
s
Zilch
(i960)
listed 25 subgenera u n d e r Bulimulus.
Breure
s h o w n that these t a x a s h o u l d either be g i v e n generic status o r
considered as s y n o n y m s . T w o other t a x a have been r e f e r r e d to Bulimulus
as
66
ZOOLOGISCHE VERHANDELINGEN 168 (1979) P e r i c a r d n e a r l y as l o n g as the n e p h r i d i u m , w h i c h is ( n a r r o w l y ) triangular.
M a i n p u l m o n a r y v e i n moderately developed, strongest at the a n t e r i o r e n d ; side veins weakly to moderately developed. T h e adrectal ureter is closed. P e n i s w i t h a p r o x i m a l sheath. T h e l u m e n o f the p e n i s is d i v i d e d into several parallel tubes o r sac-like cavities a n d l i n e d b y t w o types o f epithelium. T h e epiphallus is n o t to h a r d l y i n t r u d i n g the penis. E x t e r n a l l y the penis is m o r e o r less s u b c y l i n d r i c a l ; a m e d i a n o v o i d s w e l l i n g is present i n several species. T h e flagellum is s u b c y l i n d r i c al o r d i v i d e d into a stout a n d a slender part. T h e spermathecal
duct is m o r e o r less s u b c y l i n d r i c a l , w i t h a globose
spermatheca at its distal e n d .
F i g s . 116-124. V a r i a t i o n i n shell shape i n Naesiotus. F i g . 116. N. rhabdotus ( H a a s ) . F i g . 117. N. cutisculptus ( A n c e y ) . F i g . 118. N. gracillimus W e y r a u c h . F i g . 119. N. deletangi ( P a r o d i z ) . F i g . 120. N. fernandezae W e y r a u c h . F i g . 121. N. ochsneri ( D a l l ) . F i g . 122. N. quitensis ( P f e i f f e r ) . F i g . 123. N. carlucioi (Rezende & L a n z i e r i ) . F i g . 124. N. martinicensis ( P f e i f f e r ) . Scale = 5 m m .
BREURE, BULIMULINAE
71
perrus, Bulimulus (Naesiotus), D a l l , 1917b: 376 (Galapagos, N a r b o r o u g h Island, r i m o f the crater, 2000-4500 ft.) [ S T C A S ] . perspectivus, Bulimus, P f e i f f e r , 1846a: 33 ( L o c a l i t y u n k n o w n ) [ L T B M N H 1975166]. phlegonis, Bulimulus (Naesiotus) ustulatus, D a l l & Ochsner, 1928: 160, p i . 9 figs. 11-12, 15-17 (Galapagos, Charles L s l a n d , S W o f S p r i n g M o u n t a i n , 1650 ft.) [ S T C A S ] . pileatus, Bulimulus (Naesiotus) eschariferus, D a l l , 1896: 434 ([Galapagos] Chatham Island). pilosus, Buliminus, Guppy, 1871: 310, p l . 17 f i g . 9 ( [ W e s t Indies] T r i n i d a d ) [ S T B M N H 1875.2.8.3]. pilsbryi, Naesiotus, W e y r a u c h , 1956b : 6, p i . 1 f i g . 4 ( N o r t h e r n P e r u , R i o M a r a n o n , near Chagual, P a m p a Calquiche, 1200 m ) [ H T S M F 155698]. pinzonensis, Naesiotus, V a g v o l g y i , 1977: 772, p i . 1 figs. 6a-b, p i . 2 f i g . 2 (Ecuador , Archipiélago de C o l o n , Pinzón Island, northern and western c l i f f s o f m a i n peak, 1400-1500 ft.) [ H T U S N M 757718]. pinzonopsis, Naesiotus, V a g v o l g y i , 1977: 774, p i . 1 figs. 7a-b (Ecuador, Archipiélago de Colon, Pinzón Island, northern and western c l i f f s o f m a i n peak, 1400-1500 ft.) [ H T U S N M 757719]. planospira, Bulimulus rugulosus, Ancey, 1887: 294 (He Chatham, archipel des Galapagos). pollonerae, Bulimulus, A n c e y , 1897: 17, f i g . 10 ( S a n L o r e n z o , province de J u j u y , R é publique A r g e n t i n e ) . prepinguis, Naesiotus, V a g v o l g y i , 1977: 775, p i . 1 figs. 2a-b, p i . 2 f i g . 5 (Ecuador, Archipiélago de C o l o n , Pinzón Island, secondary peak (864 ft.), n o r t h o f the crater) [ H T U S N M 757720]. punctustriatus, Protoglyptus, P a r o d i z , 1946b: 5, f i g . 2, p i . 1 figs. 5-6 ( A r g e n t i n a , P r o v . J u j u y , Puesto V i e j o ) [ H T M A C N 091]. quitensis, Bulimus, P f e i f f e r , 1848a : 230 ( Q u i t o ) [ L T B M N H 1975320]. rabidensis, Bulimulus (Naesiotus), D a l l , 1917b: 381 (Galapagos, J e r v i s Island, 9001000 ft.) [ S T C A S ] . ramosae, Protoglyptus, H y l t o n Scott, 1952: 23, p i . 1 f i g . 6 ( [ A r g e n t i n a ] P r o v . Salta, Pocitos) [ H T M I H S ] . reibischii, Bulimulus (Naesiotus), D a l l , 1895: 126 (Galapagos, Indefatigable Island). rhabdotus, Bulimulus (Protoglyptus), H a a s , 1951: 512, f i g . 100 ( P e r u , A m b o , near Huánuco, 2000 m ) [ H T F M N H 30915]. rivasii, Bulimus, d O r b i g n y , 1836: 276, p i . 34 figs. 8-10 ( [ B o l i v i a ] sur les coteaux des derniers contreforts des A n d e s boliviennes, avant de descendre dans les pleins de Santa C r u z de l a S i e r r a , principalement à l a Cuesta de P e t a c a ) . rocayana, Helix, d O r b i g n y , 1835: 13 (província Santa C r u z de l a S i e r r a , republica Boliviana). rufescens, Bulimulus (Scutalus) quitensis, G e r m a i n , 1910: C 35, p l . 4 figs. 1-2 ( [ E c u a d o r ] C u j u j a ; valle de T u m b a c o ) . rugatinus, Bulimulus (Naesiotus), D a l i , 1917a: 10 ( N e w name f o r Bulimulus acutus Reibisch, 1892, not L e a c h , 1814). rugifer, Cochlicellus, Beck, 1837: 63 [indication]. rugiferus, Bulinus, Sowerby, 1833b: 36 (Galapagos, James Island) [ L T B M N H 1975178]. rugulosus, Bulinus, Sowerby, 1838? [1832-1841] : f i g . 87 (Galapagos) [ L T B M N H 1975176]. saeronius, Bulimulus (Naesiotus), D a l l , 1917a: 9 (Galapagos, Indefatigable Island) [ H T U S N M 274097]. sanctaeluciae, Bulimus (Leptomerus), E . A . S m i t h , 1889: 403 ( [ W e s t Indies] St. L u c i a ) [HT B M N H ] . scalesiana, Naesiotus, A . G . S m i t h , 1972: 17, figs. 19-25 (Galapagos, I s l a Santa C r u z , H o r n e m a n F a r m area) [ H T C A S 13745]. sculpturatus, Bulimus, P f e i f f e r , 1846a: 29 (Galapagos Is.) [ L T B M N H 1975174].
BREURE, BULIMULINAE
73
Orthotomium Crosse & Fischer, 1875 [1870-1894] : 473. T y p e species by o r i g i n a l designation : Bulimus sufflatus G o u l d . Globulinus Crosse & Fischer, 1875 [1870-1894] : 475. T y p e species by o r i g i n a l designation : Bulimus sufflatus G o u l d . Leptobyrsus Crosse & Fischer, 1875 [1870-1894] : 475. T y p e species by o r i g i n a l designation : Bulimus spirifer Gabb. Columna Cooper, 1892a: 215. T y p e species by m o n o t y p y : Rhodea californica ramentosa Cooper [not Columna P e r r y , 1811]. Plicolumna Cooper, 1895: 164. N e w name f o r Columna Cooper, 1892, not P e r r y , 1811. (June). Pseudorhodea D a l l , 1895b: 51. T y p e species by o r i g i n a l designation: Rhodea californica ramentosa Cooper. (September). Sonorina P i l s b r y , 1896a: 114. N e w name f o r Leptobyrsus Crosse & Fischer, 1875, Leptobyrsa Stâl, 1873. Puritanina Jacobson, 1958: 7. T y p e species by o r i g i n a l designation : Bulimulus (Scutalus) montezuma D a l l . Hannarabdotus E m e r s o n & Jacobson, 1064: 325. T y p e species by o r i g i n a l designation: Bulimulus slevini H a n n a . n o t
Description. —
S h e l l elongate-ovate
to elongate-globose
or turrited; per-
forate; t h i n to rather solid. C o l o u r w h i t i s h to yellowish, u n i f o r m l y coloured o r w i t h a x i a l b r o w n streaks. S u r f a c e smooth. P r o t o c o n c h w i t h straight a x i a l riblets. W h o r l s slightly elongate-ovate.
convex;
suture well impressed. A p e r t u r e s u b -
to
P e r i s t o m e t h i n , simple to e x p a n d e d a n d m o r e o r less reflexed.
C o l u m e l l a w i t h o r without a fold. C e n t r a l teeth o f the r a d u l a are t r i c u s p i d , w i t h lanceolate to
wedge-shaped
mesocones a n d ovate to deltoid ectocones. L a t e r o m a r g i n a l teeth are b i c u s p i d , with elongate to lanceolate mesocones a n d ovate to deltoid ectocones. H a l f - r o w formula: C / 3 + L M x / 2 (x =
28-36).
P e r i c a r d is ca. h a l f as l o n g as the n e p h r i d i u m , w h i c h is n a r r o w l y triangular. The
m a i n p u l m o n a r y v e i n is p r o m i n e n t a n d the
side-veins
are
strongly
developed, especially at the anterior e n d . T h e adrectal ureter is closed o v e r its entire length. P e n i s w i t h a p r o x i m a l sheath. T h e l u m e n o f the penis is d i v i d e d i n a p r o x i m a l part ( b r o a d , w i t h pouches) a n d a distal part ( n a r r o w ) . T h e penis is swollen above the distal e n d o f the sheath a n d the transitio n to the e p i p h a l lus is, both internally a n d externally, g r a d u a l. T h e epiphallus is
relatively
long. T h e flagellum is slender, w i t h a distally attached retractor muscle. T h e spermathecal duct is s u b c y l i n d r i c a l , n a r r o w , a n d w i t h a globose
spermatheca
at the distal e n d . D i s t r i b u t i o n . — M e x i c o , southern U n i t e d States. Ecology. —
T h e species live o n rocks o r " u p o n bushes a n d other vegeta-
t i o n " ( P i l s b r y , 1946a: 6 ) . Relationships. — T h e phylogenetic relationships o f this genus are discussed
.—.
VJ
vo
M
s
ZOOLOGISCHE VERHANDELINGEN 168 (1979)
78 Description. —
" S h e l l rimate t u r r i t e d , straightly t a p e r i n g to a n
obtuse
r o u n d e d a p e x w h i c h is retained i n adult shells. W h o r l s about n , the first 2 vertically costellate,
the n e x t
h a v i n g the
riblets
cut into s p i r a l series
of
granules; last w h o r l b e c o m i n g free i n front , acutely keeled above. A p e r t u r e oblique, semicircular-ovate, the peristome b r o a d l y e x p a n d e d a n d subreflexed . Internal a x i s
imperforate, very
slender a n d w e a k l y
s i g m o i d w i t h i n each
w h o r l " ( P i l s b r y , 1902c: 57). C e n t r a l teeth o f the r a d u l a are t r i c u s p i d , w i t h lanceolate to
wedge-shaped
mesocones a n d ovate to deltoid ectocones. L a t e r o m a r g i n a l teeth are b i c u s p i d , w i t h elongate to lanceolate mesocones a n d ovate to deltoid ectocones. H a l f row formula: C / 3 + L M x / 2 (x =
30-34).
P e r i c a r d n e a r l y as l o n g as the n e p h r i d i u m , w h i c h is t r i a n g u l a r . T h e m a i n p u l m o n a r y v e i n is p r o m i n e n t , especially at the a n t e r i o r e n d , wher e the side veins are also strongly developed. T h e adrectal ureter is o p e n at the anterior e n d o v e r 1/5-1/6 o f its length. T h e peni s has a p r o x i m a l sheath a n d is s w o l l e n above the distal e n d o f the sheath. T h e epiphallus a n d especially the flagellum are relatively l o n g . T h e spermathecal duct is n a r r o w a n d s u b c y l i n d r i c a l , w i t h a globose
sper-
matheca at the distal e n d . Distribution. — Ecology.
Mexico (Baja California).
— T h e species aestivates sealed to rocks ( C h r i s t e n s e n , i n l i t t . ) .
T h e o n l y t a x o n i n c l u d e d i n this genus is: taylori, Clausilia? (Balea?), P f e i f f e r , 1861a: 27, p i . 2 f i g . 7 (Localitas ignota).
Spartocentrum D a l l ,
1895
Spartocentrum D a l l , 1895b: 51. T y p e species by o r i g i n a l designation: Cylindrella (Urocoptis) irregularis Gabb. Teneritia M a b i l l e , 1897: 79. T y p e species by present designation: Berendtia digueti Mabille. Description. —
" S h e l l m a n y - w h o r l e d , slender, c y l i n d r i c below,
above, r e t a i n i n g the a p e x entire. A p e x bulbous, the f i r s t 2
tapering
w h o r l s vertically
costellate, f o l l o w i n g 2 w h o r l s decussated, granose, subsequent w h o r l s r i b b e d , the last w i t h n o trace o f a s u b p e r i p h e r al c o r d , adnate o r b e c o m i n g free. I n ternal a x i s hollow, smooth, somewhat sinuous w i t h i n each w h o r l , h a v i n g a s p i r a l swellin g o r c o n v e x i t y " ( P i l s b r y , 1902c:
51).
T h e central teeth o f the r a d u l a are t r i c u s p i d , w i t h lanceolate to wedgeshaped mesocones a n d ovate to deltoid ectocones. L a t e r o m a r g i n a l teeth are b i c u s p i d , w i t h elongate to lanceolate mesocones a n d deltoid ectocones, w h i c h are serrate i n the outermost teeth. (x = 24).
H a l f - r o w formula:
C/3
+
L M
x/2
82
ZOOLOGISCHE V E R H A N D E L I N G E N
168
(1979)
s t r o n g l y developed a n d f o r m a deltoid network . T h e adrectal ureter is closed. T h e penis has a p r o x i m a l sheath a n d is rather stout a n d m o r e o r less s u b c y l i n d r i c a l i n f o r m . T h e l u m e n o f the penis is n a r r o w ; i n the distal part o f the penis, w h e r e a d i f f e r e n t type o f e p i t h e l i u m is f o u n d , a short b l i n d sac is present. T h e t r a n s i t i o n to the epiphallus, both i n t e r n a l l y a n d externally, is g r a d u a l . T h e flagellum is rather slender, w i t h a distally attached
Fig. 141. Distribution of
Scutalus.
retractor
BREURE, BULIMULINAE w i t h s p i r a l bands o r a variegate
pattern. S u r f a c e
85 rather s h i n i n g ,
growth
striae incrassate o r w i t h a x i a l riblets. P r o t o c o n c h w i t h a x i a l w r i n k l e s . W h o r l s rather flat, the last w h o r l m o r e o r less inflated. A p e r t u r e (sub)ovate.
Per-
istome t h i n to slightly thickened, simple o r h a r d l y e x p a n d e d . T h e central teeth o f the r a d u l a are t r i c u s p i d , w i t h lanceolate to w e d g e shaped mesocones a n d ovate to deltoid ectocones. T h e l a t e r o m a r g i n a l teeth are b i c u s p i d , w i t h elongate to lanceolate mesocones
a n d deltoid
ectocones,
w h i c h are serrate i n the outermost teeth. H a l f - r o w f o r m u l a : C / 3 + L M (x =
x/2
26-33).
T h e p e r i c a r d is as l o n g as the n e p h r i d i u m , w h i c h is ( b r o a d l y ) t r i a n g u l a r . The
main
pulmonary
vein
is
p r o m i n e n t a n d b r o a d , the
side
veins
are
moderately to well developed, especially at the anterior e n d , w h e r e also a v e i n parallel to the m a i n p u l m o n a r y v e i n m a y be f o u n d . T h e adrectal ureter is partially o p e n (1/10-1/2 o f its l e n g t h ) . T h e penis has a p r o x i m a l sheath a n d is s u b c y l i n d r i c a l i n f o r m . T h e l u m e n of the p r o x i m a l part is rather n a r r o w b y i n f o l d i n g s . M o r e distally the l u m e n is wider, but n a r r o w again at the transitio n to the distal p a r t o f the penis. T h e t r a n s i t i o n to the epiphallus, both internally a n d externally, is g r a d u a l . T h e flagellum
is slender a n d rather l o n g ; the retractor m u s c le is distally
attached. T h e spermathecal duct is m o r e o r less tapering , w i t h a (elongate-) globose spermatheca at its distal e n d . Distribution. — Bolivia, P e r u , Ecuador. Ecology. —
T h e species live m a i n l y o n shrubs, mostl y nea r r o c k y outcrops.
T h e vertical d i s t r i b u t i o n is 2600-5000 m . T a x a . — T h e f o l l o w i n g t a x a are i n c l u d e d i n this subgenus: achrous, Bulimulus (Scutalus), H a a s , 1952: 126, f i g . 24 ( B o l i v i a , Dept. Cochabamba, T a r a t a , 2800 m ) [ H T F M N H 39720]. aequatorius, Bulimus, P f e i f f e r , 1853d : 420 (reipublicae Aequatoris, monte Schinchulagua) [ L T B M N H 1975377]. altorum, Bulimulus (Scutalus) revinctus, H a a s , 1951: 516, f i g . 104 ( P e r u , puna between Andahuaylas and Abancay, 400 m) [ H T F M N H 30912]. angrandi, Bulimus, Morelet, i 8 6 0 : 372 (Pérou, Huancabelica). anthisanensis, Bulimus, P f e i f f e r , 1853d: 406 (reipublicae Aequatoris, monte A n t h i s a n a ) [ L T B M N H 1975372]. aquilus, Bulimus, Reeve, 1848: p i . 22 f i g . 138 ( P e r u , T a c n a ) [ L T B M N H 1975376]. aureus, Scutalus (Vermiculatus), Breure, 1978b: 170, p i . 10 f i g . 12 ( P e r u , Dept. C a j a marca, 26 k m N E Encanada, 3650 m ) [ H T U F 22754]. badius, Bulinus, Sowerby, 1835 : 141 (província P e r u v i a e X a g u a ) . bicolor, Bulinus, Sowerby, 1835: 141 (província P e r u v i a e X a g u a ) [ L T B M N H 1975151]. bolivianus, Bulimulus (Scutalus), M a r s h a l l , 1932: 2, p l . 1 figs. 3-4 ( B o l i v i a , A y a p a y o R i v e r , T a m a n a n i , 85 miles N E O r u r o ) [ H T U S N M 382216]. coagulatus, Bulimus, Reeve, 1849: p l . 77 f i g . 558 ( P e r u ) [ L T B M N H 1975351]. confusus, Bulimus, Reeve, 1848: p l . 48 f i g . 316 [no type locality g i v e n ; L T B M N H 1975194].
92
ZOOLOGISCHE VERHANDELINGEN 168 ( 1979)
colours, sometimes w i t h s p i r a l bands. S u r f a c e w i t h u n e v e n , w r i n k l e d g r o w t h striae, usually stronger below the suture, o f t e n variable strength.
crossed b y s p i r a l lines o f
Protoconch with axial wrinkles
o r reticulate.
Aperture
ovate. P e r i s t o m e t h i n a n d simple. Distribution. —
A u s t r a l i a : N o r t h e r n T e r r i t o r y , P a l m V a l l e y near H e r -
m a n n s b u r g (fossi l r e c o r d , see M c M i c h a e l , 1968); T a s m a n i a ; S o u t h A u s t r a l i a , K a n g a r o o Island, E y r e P e n i n s u l a a n d w e s t w a r d ;
W e s t e r n A u s t r a l i a , entire
southern coast, n o r t h w a r d to the H a m e r s l e y R a n g e . Remarks. —
Z i l c h ( i 9 6 0 ) o n l y listed the t a x a described b y Iredale a n d d i d
not place them i n his system. B u r c h (1976) considered Iredale's of Bothriembryon
subgenera
as usable, but study o f the shell m o r p h o l o g y a n d o f the
anatomy (see B r e u r e , 1978b) has c o n v i n c e d m e that the t a x a are s y n o n y m o u s w i t h Bothriembryon
s.str., except
Tasmanembryon.
F i g s . 142-145. V a r i a t i o n i n shell shape i n Bothriembryon. F i g . 142. B. (B) dux ( P f e i f f e r ) . F i g . 143. B. (B.) inflatus ( L a m a r c k ) . F i g . 144. B. (B.) indutus ( M e n k e ) . F i g . 145. B. (Tasmanembryon) gunnii ( S o w e r b y ) . Scale = 5 m m . K e y to the subgenera o f a.
P r o t o c o n c h pit-reticulate m o r e o r less globose
b.
Bothriembryon
o r w i t h anastomosin g
w r i n k l e s.
Bothriembryon
Spermatheca (Bothriembryon)
P r o t o c o n c h w i t h oblique a x i a l w r i n k l e s , the intervals about as b r o a d as the w r i n k l e s , crossed b y weaker s p i r a l lines. S p e r m a t h e c a b r o a d l y elongateovate. Spermatheca l duct relatively short Bothriembryon
Bothriembryon (Bothriembryon) P i l s b r y , Description. —
(Tasmanembryon) 1894
S h e l l elongate-ovate to o b l o n g - c o n i c a l; rather solid. C o l o u r
variable, w i t h shades o f yellow, b r o w n , r e d , lillac a n d white, o f t e n i n patterns
96
ZOOLOGISCHE VERHANDELINGEN 168 (1979)
brachysoma, Bothriembryon gunni, P i l s b r y , 1900b: 19, p l . 3 f i g . 53 ( E a s t coast o f Tasmania) [ H T A N S P 8461]. gunnii, Bulinus, Sowerby, 1845: 298, p i . 19 f i g . 5 [Tasmania, near H o b a r t ; P l i o c e n e ; teste P i l s b r y , 1900b: 18]. tasmanicus, Bulimus, P f e i f f e r , 1853b : 260 ( V a n Diemen's L a n d ) .
Oxychona M ö r c h ,
1852
Oxychona Mörch, 1852: 14. T y p e species by monotypy : Trochus bifasciatus B u r r o w . Description. —
Shell conical; imperforate; (rather) thin. C o l o u r u n i f o r m l y
w h i t i s h o r w i t h p u r p l i s h - b r o w n s p i r a l bands. S u r f a c e (malleate,) w i t h s p i r a l lines. P r o t o c o n c h w i t h a g r a t i n g sculpture o f a x i a l riblets a n d s p i r a l striae that are o f equal strength. W h o r l s ( n e a r l y ) flat, the last w h o r l s h a r p l y keeled a n d w i t h a flat to concave base;
suture h a r d l y impressed. A p e r t u r e v e r y
oblique a n d skewed, t r i a n g u l a r. P e r i s t o m e p a r t l y e x p a n d e d a n d reflexed . C e n t r a l teeth o f the r a d u l a m o n o c u s p i d , w i t h rather blunt, spatula-shaped mesocones. T h e lateral teeth are b i c u s p i d , w i t h b l u n t spatula-shaped m e s o cones a n d small, acute, ovate to t r i a n g u l a r ectocones situated at the basal plate. H a l f - r o w f o r m u l a : C / i +
that are p o s t e r i o r ly
L M x/2 (x =
37).
P e n i s w i t h a p r o x i m a l sheath, m o r e o r less s u b c y l i n d r i c a l a n d p a s s i n g without e x t e r n a l d i f f e r e n t i a t i o n into the epiphallus. T h e flagellu m is s u b c y l i n d r i c a l a n d relatively l o n g , internally w i t h a d o u b l e - c u r v e d longitudina l fold. Distribution. —
Brazil.
E c o l o g y . — T h e species live o n trees. T a x a . — T h e f o l l o w i n g t a x a are i n c l u d e d i n this genus: bifasciatus, Trochus, B u r r o w , 1815: 188, p i . 27 f i g . 2 [publication not seen]. blanchetiana, Helix, M o r i c a n d , 1833: 539, p i . 1 f i g . 3 (Brésil, aux environs de B a h i a ) [ST Z M B ] . bosciana, Helix, Férussac, 1821: 37 [nomen n u d u m ] . currani, Oxychona pyramidella, B a r t s c h , 1916: 53 ( B r a z i l , B a h i a , R i o G r u n g u g y [ G r u n g o g y ] ) [ H T U S N M 322281]. gyrina, Helix, 'Valenciennes' Deshayes, i n Férussac & Deshayes, 1820-1851 : p l . 63B f i g . 4 (Brésil). lonchostoma, Caracolla, M e n k e , 1828: 76 (intes[sic] R i o et Campo, B r a s i l i a ) . minarum, Drymaeus (Oxychona) bifasciatus, A n c e y , 1901a: 93 ( [ B r a z i l ] M i n a s Geraes). pyramidella, Helix, W a g n e r i n S p i x , 1827: 22 (sylvis mediterraneis inter montem sanctum et flumen S. F r a n c i s c i , i n Província B a h i e n s i ) .
Otostomus B e c k ,
1837
Otostomus Beck, 1837: 55. T y p e species by subsequent designation ( G r a y , 1847) : Auris signata S p i x . Description. —
S h e l l obliquely elongate-ovate;
perforate ;
solid. C o l o u r
whitish, w i t h b r o a d b r o w n s p i r a l bands o n the last w h o r l . S u r f a c e
with
granules a n d incrassate g r o w t h striae. P r o t o c o n c h w i t h a g r a t i n g sculpture
98
ZOOLOGISCHE VERHANDELINGEN 168
t a p e r i n g a n d relatively
(1979)
l o n g , internally w i t h a d o u b l e - c u r v e d
longitudinal
fold. Distribution. — Ecology. —
Brazil.
T h e species
live o n trees. T h e vertical d i s t r i b u t i o n is o-ca.
500 m . Relationships. —
T h i s g e n us was classified as a subgenus o f Drymaeus
P i l s b r y (1898) o r considered closely related to Drymaeus [Weyrauch
(1958) e r r o n e o u s l y s y n o n y m i z e d Drymaeus
A n a t o m i c a l research shows that Cochlorina Otostomus
by
( W e y r a u c h , 1960a) with
Cochlorina],
closely resembles Oxychona
and
( B r e u r e & E s k e n s , i n p r e p a r a t i o n ) . See page 158 f o r a discussion
o f the phylogenetic relationships. T a x a . — T h e f o l l o w i n g t a x a are placed i n this genus: aurisleporis, Bulimus, Bruguière, 1792: 346 (l'île de Madagascar [sic]). aurismuris, Helix, S. M o r i c a n d , 1839: 140, p l . 3 figs. 1-3 ( [ B r a z i l ] province de B a h i a , à l a fazenda de P a l m e i r i n h a , entre C a x o e i r a et Jacobina). auritum, Stenostoma, S p i x , 1827: 18, p l . 13 figs. 1-2 ( [ B r a z i l ] Província Sebastianopolitana). fasciata, Navicula, S p i x , 1827: p l . 15 figs. 2-3 ( [ B r a z i l ] sylvis aborginibus P r o v i n c i a e Bahiensis). intensior, Drymaeus (Zaplagius) aurisleporis, P i l s b r y , 1898: 190, p l . 28 f i g . 4 [no type locality g i v e n ] . involutus, Bulimulus, M a r t e n s , 1867a: 63 ( B r a s i l i e n ) . lagotis, Bulimus, M e n k e , 1828: 15 [indication]. lateralis, Bulimus, M e n k e , 1828: 76 ( B r a s i l i a ) . lateritius, Drymaeus (Zaplagius), P i l s b r y , 1898: 320 ( B r a z i l , P r o v . o f B a h i a ) [ H T A N S P 73553]. leporis, Auricula, L a m a r c k , 1822: 138 (Madagascar [sic]). lyonetianus, Bulimus, K ü s t e r i n K ü s t e r et al., 1840-1865: 23, p l . 5 figs. 5-7 (Insel Frankreich [?]). myotis, Otostomus, Beck, 1837: 55 [indication]. navicula, Helix, W a g n e r i n S p i x , 1827: 22 ( [ B r a z i l ] sylvis aborginibus P r o v i n c i a e Bahiensis). uranops, Drymaeus (Zaplagius), P i l s b r y , 1898: 188, p l . 27 figs. 24-27 ( B r a z i l ) .
Newboldius P i l s b r y ,
1932
Newboldius P i l s b r y , 1932: 398. T y p e species by o r i g i n a l designation: Newboldius inca Pilsbry. Description. —
S h e l l elongate-ovate; i m p e r f o r a t e ; thick a n d solid. C o l o u r
w h i t i s h to l i g h t b r o w n , the last w h o r l w i t h 3-4 b r o w n i s h s p i r a l bands.
Surface
w i t h m o r e o r less incrassate g r o w t h striae; sometimes w i t h s p i r al lines a n d / o r malleation. P r o t o c o n c h [ w i t h a g r a t i n g sculpture o f a x i a l riblets a n d spiral striae, w h i c h are o f equal s t r e n g t h ] . W h o r l s h a r d l y c o n v e x;
suture
weakly
impressed. A p e r t u r e elongate-ovate, slightly oblique, a sulcus at the basalcolumellar m a r g i n . P e r i s t o m e b r o a d l y e x p a n d e d a n d reflexed, thick; b r o w n i s h to p i n k coloured. A callus at the parietal region.
100
ZOOLOGISCHE VERHANDELINGEN 168 (1979)
w i t h rather blunt, elongate to lanceolate mesocones a n d t r i a n g u l a r to deltoid ectocones. T h e m a r g i n a l teeth are t r i c u s p i d , shifted, w i t h rather blunt, ovate mesocones, acute elongate-ovate endocones a n d acute, deltoid ectocones, w h i c h m a y be b i f i d i n the outermost teeth. H a l f - r o w f o r m u l a : C / i M y / 3 (x = 2-6,y =
+
L x/2
+
76-84) .
T h e n e p h r i d i u m is b r o a d l y t r i a n g u l a r , its p e r i c a r d i a l side strongly c u r v e d . T h e m a i n p u l m o n a r y v e i n is p r o m i n e n t , r a m i f i e d at its anterior e n d . T h e adrectal ureter is closed except f o r a short distance nea r the u r i n a r y o p e n i n g . P e n i s w i t h a v e r y short sheath, m o r e o r less s u b c y l i n d r i c a l a n d p a s s i n g without external d i f f e r e n t i a t i o n into the epiphallus, w h i c h is slightly tapering. T h e flagellum is ( r a t h e r ) slender. T h e spermathecal duct is s u b c y l i n d r i c a l, w i t h a globose spermatheca at the distal e n d . Distribution. — Ecology. — and
P e r u (Dept. A n c a s h, L a Libertad, Cajamarca, A m a z o n a s ) .
T h e ecology o f o n l y one species is k n o w n , w h i c h lives o n trees
shrubs i n x e r o p h y t i c savannah vegetation.
8oo-i9oo("320o)
T h e vertical d i s t r i b u t i o n is
m.
Relationships. —
T h e phylogenetic relationships o f this genus are discussed
o n page I58ff. T h e genus is characterized b y the shell shape, the p r o t o c o n ch sculpture a n d the e x p a n d e d peristome. T a x a . — T h e f o l l o w i n g t a x a are i n c l u d e d i n
Neopetraeus:
altoperuvianus, Bulimus, Reeve, 1849: p i . 72 f i g . 521 (Chachapoyas, A l t o - P e r u ) [ L T B M N H 1975437]. arboriferus, Neopetraeus, P i l s b r y , 1898: 175, p i . 32 figs. 32-33 (Andes o f P e r u ) [ L T A N S P 4684a]. atahualpa, Bulimulus, D o h r n , 1863 '· 153 [no type locality g i v e n ; S T B M N H ] . binneyanus, Bulimus, P f e i f f e r , 1857dl 229 ( A n d i b u s prov. Patas, P e r u ) [ L T B M N H 1975426]. brownii, Neopetraeus decussatus, P i l s b r y , 1898: 179, p i . 32 figs. 40-41, p i . 33 f i g . 39 ( P e r u ) [ L T A N S P 4692a]. camachoi, Neopetraeus, W e y r a u c h , 1967a : 418, figs. 68-70 ( N o r t e de P e r u , r i o Chotano, Cuesta de C h u g u i d , 1500 m ) [ H T I M L 1541a]. catamarcanus, Bulimus, P f e i f f e r , 1858: 256, p l . 42 f i g . 5 ( P e r u , A n d e s o f the province C a x a m a r c a [ s i c ] ). cora, Helix, d O r b i g n y , 1835 : 15 (republica P e r u v i a n a ) . cremnobates, Neopetraeus, P i l s b r y , 1949 : p i . 3 f i g . 9 [nomen nudum] ; 'Pilsbry* H . B . B a k e r , 1963: 227 (hydroelectric plant on R i o Santa, near southern edge o f Dept. de L i b e r t a d , P e r u ) [ H T A N S P 185841a]. decussatus, Bulimus, Reeve, 1849: p i . 72 f i g . 519 (Andes o f C a x a m a r c a [sic] P e r u ) [ L T B M N H 1975180]. excoriatus, Bulimus, P f e i f f e r , 1855b: 123 (Andes o f P e r u ) [ L T B M N H 1975500]. filiola, Drymaeus (Neopetraeus), P i l s b r y , 1897a: 22 ( P e r u ) [ H T A N S P 25724]. gracilior, Neopetraeus altoperuvianus, ' P f e i f f e r ' P i l s b r y , 1898: 173, p i . 33 figs. 37-38 [no type l o c a l i t y ; H T A N S P 63081]. heterogyrus, Bulimus, P h i l i p p i , 1869: 42 ( [ P e r u ] inter Sartimbamba et Chusgon i n departamento de l a L i b e r t a d ) . latistrigatus, Neopetraeus arboriferus, P i l s b r y , 1898: 176, p i . 32 figs. 34-35 (Andes o f P e r u ) [ L T A N S P 4685a].
BREURE, BULIMULINAE
101
lobbii, Bulimus, Reeve, 1849 : p l . 72 f i g . 516 ( B a n k s of the M a r a n o n near Balsas, P e r u ) [ L T B M N H 1975431]. millegranus, Otostomus, M a r t e n s , 1883: 177, p i . 32 figs. 1-4 (Balzas, P e r u v i a e orientalis, 963 meter) [ H T Z M B 36493]. myristicus, Bulimus, Reeve, 1849: p i . 72 f i g . 520 (Andes o f C a x a m a r c a , P e r u ) [ L T B M N H 1975433]. obesus, Neopetraeus arboriferus, W e y r a u c h , 1967a: 416, figs. 62-63 ( P e r u ) [ H T I M L 1244a]. orientalis, Neopetraeus catamarcanus, Breure, 1978b: 212, f i g . 364, p i . 4 figs. 3-4, p i . 5 ( P e r u , Dept. Amazonas, 18 k m E N E Balsas, 1880 m ) [ H T U F 22779]. patasensis, Bulimus, P f e i f f e r , 1858: 257, p l . 42 f i g . 6 ( P r o v i n c e o f Patas, A n d e s o f P e r u ) [ L T B M N H 1975439]· paucistrigatus, Neopetraeus arboriferus, W e y r a u c h , 1967a: 417, figs. 65-67 ( N o r t e de P e r u , valle del r i o M a r a n o n , rut a de H u a m a c h u co a P a t a z , Chagual, 1300 m ) [ H T I M L 1239a]. perincrassatus, Neopetraeus tessellatus, P i l s b r y , 1898: 169, p i . 31 figs. 18-19, p i . 33 f i g . 48 ( P e r u ) [ H T A N S P 72113]. platystomus, Bulimus, P f e i f f e r , 1858: 256, p l . 42 f i g . 2 ( P r o v i n c e o f Patas, A n d e s o f P e r u ) [ L T B M N H 1975428]. ptychostylus, Bulimus, P f e i f f e r , 1858: 256, p l . 42 f i g . 7 ( P r o v i n c e o f Patas, A n d e s o f P e r u ) [ L T B M N H i97543o]. rectistrigatus, Neopetraeus arboriferus, P i l s b r y , 1898: 176, p i . 32 figs. 36-37 (Andes o f P e r u ) [ L T A N S P 4683a]. tessallatus, Bulimus, Shuttleworth, 1852: 200 [no type locality g i v e n ] . unicolor, Bulimus cora, P f e i f f e r i n P f e i f f e r & Clessin, 1881: 245 [indication]. vadum, Neopetraeus, P i l s b r y , 1898: 165, p i . 29 figs. 32-34 ( P e r u ) [ L T A N S P 5260a]. weyrauchi, Neopetraeus, P i l s b r y , 1944a: 88, p i . 9 f i g . 4 ( P e r u , Santa valley, H u a r a z , 3200 m ) [ H T A N S P 170080].
Llaucanianus W e y r a u c h , Llaucanianus W e y r a u c h , 1967a: 420. T y p e Weyrauch. Description. — uniformly
1967
species by m o n o t y p y : Llaucanianus haasi
S h e l l ovate-conical; n a r r o w l y p e r f o r a t e ; solid. C o l o u r p i n k ,
coloured o r w i t h oblique b r o w n streaks
o n the u p p e r w h o r l s .
S u r f a c e w i t h v e r y fine s p i r a l lines. P r o t o c o n c h w i t h a x i a l riblets a n d less prominent
spiral striae. L a s t w h o r l relatively large a n d slightly
inflated.
A p e r t u r e elongate-ovate, skewed, orange c o l o u r e d inside. P e r i s t o m e e x p a n d e d , white. T h e anatomy is u n k n o w n . Distribution. — P e r u (Dept. Cajamarca). E c o l o g y . — T h e species lives i n rock-chasm s at 2700 m altitude. Remarks. — Neopetraeus of Neopetraeus
T h i s genus,
a n d Drymaeus,
w h i c h is m o r e o r less intermediate
between
m a y p r o v e to be better classified as a subgenus
once the anatomy has been studied.
T a x a . — T h e o n l y t a x o n is: haasi, Llaucanianus, W e y r a u c h , 1967a: 421, figs. 34-36 ( P e r u , [Dept. Cajamarca] P e n a R o t a , margen izquierda del r i o L l a u c a n , 8 k m N E Bambamarca, 2700 m ) [ H T S M F 162046].
102
ZOOLOGISCHE VERHANDELINGEN 168 (1979)
Stenostylus P i l s b r y , 1898 Stenostylus P i l s b r y , 1898: 184. T y p e species b y subsequent designation ( P i l s b r y , 1898) : Bulimus nigrolimbatus P f e i f f e r . Description. —
S h e l l ovate; i m p e r f o r a t e ;
(rather) thin. C o l o u r yellowish
to light b r o w n , w i t h a x i a l streaks o f ( d a r k e r ) b r o w n . S u r f a c e w i t h incrassate g r o w t h striae a n d / o r s p i r a l lines. P r o t o c o n c h w i t h a g r a t i n g sculpture o f a x i a l riblets a n d s p i r a l striae, w h i c h a r e o f equal strength. W h o r l s slightly c o n v e x ; suture slightly crenulate, well impressed. A p e r t u r e (elongate-)ovate, w i t h a pearly layer inside. P e r i s t o m e t h i n a n d simple. T h e central teeth o f the r a d u l a ar e m o n o c u s p i d , relatively small, w i t h blunt, elongate to t r i a n g u l a r mesocones. L a t e r a l teeth a r e b i c u s p i d , slightly shifted, w i t h ( r a t h e r ) blunt, elongate to lanceolate mesocones a n d small, t r i a n g u l a r to deltoid ectocones. T h e m a r g i n a l teeth ar e t r i c u s p i d , shifted, w i t h rather blunt, ovate mesocones, acute elongate-ovate endocones a n d acute deltoid ectocones. H a l f - r o w f o r m u l a : C / i + L x / 2 + M y / 3 ( x = 7-8, y = 64). P e n i s w i t h a short sheath (1/5-1/11 the length o f the p h a l l u s ) , m o r e o r less
subcylindrical a n d passing
without
external
differentiation
into the
epiphallus. T h e flagellum is (rather) short, s u b c y l i n d r i c a l. T h e spermathecal duct is m o r e o r less s u b c y l i n d r i c a l , w i t h a n elongate-globose
spermatheca at
the distal e n d . D i s t r i b u t i o n . — PVenezuela, P e r u , F E c u a d o r , PColombia. E c o l o g y . — T h e species live u n d e r stones a n d i n rock-chasms . T h e vertical d i s t r i b u t i o n is 3000-ca. 4000 m . Remarks. —
Stenostylus
w a s described as a section o f Drymaeus,
but
W e y r a u c h (1956a) has c o n v i n c i n g l y s h o w n that it is j u s t i f i e d to r e g a r d this t a x o n as a separate genus. S o m e o f the species i n c l u d e d b y P i l s b r y (1898) i n Stenostylus
p r o v e d to belong to Scutalus
latter t a x o n closely resembles Stenostylus
(Suniellus)
B r e u r e , 1978. T h i s
i n the shell m o r p h o l o g y , b u t m a y
be r e a d i l y distinguished b y the sculpture o f the p r o t o c o n c h, a n d the structure o f the r a d u l a. Relationships. —
T h e phylogenetic
relationships a r e discussed o n page
I58ff. T h e genus is characterized b y the shell shape, the c o l o u r a n d the sculpture o f the protoconch. B i b l i o g r a p h y . — T h e m a i n publications are: B r e u r e , 1978b; P i l s b r y , 1898; W e y r a u c h , 1956a. T a x a . — T h e f o l l o w i n g t a x a are i n c l u d e d i n
Stenostylus:
colmeiroi, Bulimus, H i d a l g o , 1872: 122 (Baeza, República del E c u a d o r ) . [?] guttula, Bulimus, P f e i f f e r , 1854b: 154 (Gualea, reipublicae Aequatoris) m a y prove to belong to Simpulopsis (Eudioptus)]. meleagris, Bulimus, P f e i f f e r , 1854b: 157 (Andes o f N e w G r a n a d a ) .
[this t a x o n
BREURE, BULIMULINAE
105
teeth b.
Drymaeus
s.str.
P e r i s t o m e u s u a l l y s i m p l e ; mandíbula w i t h m o r e t h an 20 plates, w h i c h are ca. 8 times as l o n g as w i d e ; transverse r o w s o f r a d u l a V - o r W - s h a p e d , w i t h relatively small t r i - to m u l t i c u s p i d c e n t r a l a n d l a t e r o m a r g i n a l teeth Drymaeus
Drymaeus (Drymaeus) A l b e r s , Description. —
S h e l l elongate-ovate;
(Mesembrinus)
1850
( n a r r o w l y ) p e r f o r a t e;
t h i n to solid.
C o l o u r w h i t i s h , y e l l o w i s h o r p i n k , w i t h a x i a l streaks, s p i r a l bands
and/or
spots o f b r o w n , r e d , black o r yellow. S u r f a c e w i t h incrassate g r o w t h striae. Aperture
elongate-
to
obliquely
ovate, i n v e r s e d e a r - s h a p ed
o r triangular.
P e r i s t o m e ( s i m p l e to) b r o a d l y e x p a n d e d . Mandíbula w i t h 13-18 verse
rows
of
plates, w h i c h are 4-5 times as l o n g as wide. T r a n s -
r a d u l a u s u a l l y straight,
C e n t r a l teeth are (1)
slightly
V-shaped.
t r i c u s p i d , w i t h elongate-ovate to lanceolate
mesocones
a n d rather blunt, t r i a n g u l a r ectocones; deltoid mesocones. (rather)
i n some
species
o r (2) m o n o c u s p i d , w i t h t r i a n g u l a r to
L a t e r a l teeth are b i - to t r i c u s p i d , slightly shifted,
b l u n t elongate-ovate mesocones,
(curved,
acute,
lanceolate
with endo-
cones) a n d t r i a n g u l a r to deltoid ectocones that are p o s t e r i o r l y situated o n the basal plate. M a r g i n a l teeth t r i c u s p i d , shifted, w i t h lanceolate to elongate-ovate mesocones, c u r v e d lanceolate endocones a n d relatively small deltoid ectocones that m a y be b i f i d . H a l f - r o w f o r m u l a : C / y -
40-67) o r C / 3 + L M x / 3 ( x =
P e n i s w i t h a (relatively)
1+
L
x/2
+
M
y/3
(x =
1-4,
5 2 - 8 7 ).
short sheath, w h i c h is absent i n a few
species;
s u b c y l i n d r i c a l a n d p a s s i n g without externa l d i f f e r e n t i a t i o n into the epiphallus. F l a g e l l u m , s u b c y l i n d r i c a l , relatively
short;
the retractor muscle
is
distally
inserted. V a g i n a relatively long. S p e r m a t h e c a l duct s u b c y l i n d r i c a l o r t a p e r i n g, usually as l o n g as s p e r m o v i d u c t, but i n some species r e d u c e d i n length. T h e spermatheca is m o r e o r less globose, but n o r m a l l y not d i f f e r e n t i a t e d i n species w i t h a r e d u c e d spermathecal duct. Distribution. —
Venezuela,
Brazil,
Uruguay,
Argentina, Bolivia,
Peru,
E c u a d o r , Colombia, Panama , Costa R i c a , Nicaragua, H o n d u r a s , Guatemala, Mexico. Ecology. —
T h e species live o n shrubs ( a n d t r e e s ? ) . T h e vertical d i s t r i -
b u t i o n is u p to 2900 m . Taxa. —
T h e f o l l o w i n g t a x a are placed i n this subgenus
(taxa
tentatively
r e f e r r e d to this subgenus are m a r k e d b y a n asterisk) : *abruptus, Bulimulus 1947.2.10.1]. abscissus, Bulimus, 1975497].
(Drymaeus),
Pfeiffer,
1855h:
Rolle, 116
1905: (Prov.
35 of
(Peru,
Huancabamba)
Quito, Ecuador)
[LT
[HT BMNH
BREURE, BULIMULINAE bolivianos, Bulimus, P f e i f f e r , 1846a: 34 ( M e r i d a , A n d e s o f B o l i v i a [sic]) [ L T
107 BMNH
1975444]. botterii, Bulimulus (Drymaeus), Crosse & Fischer, 1875: 52 ( i n v i c i n i o civitatis O r i z a b a , reipublicae M e x i c a n a e ) . boucardi, Drymaeus, D a Costa, 1907: 305, p i . 26 figs. 5~5a ( [ P a n a m a ] C h i r i q u i ) [ H T B M N H 1007.11.21.26]. bourcieri, Bulimus, P f e i f f e r , 1853d: 314 ( P i c h i n c h a , reipublicae Aequatorius) [LT B M N H 1975446]. brachystoma, Helix, d O r b i g n y , 1835: 18 (província Santa C r u z de l a S i e r r a , republica Boliviana). branneri, Drymaeus, F . B a k e r, 1914: 637, p i . 23 figs. 1-4 (280 k m above P o r t o V e l h o , B r a z i l ) [ H T A N S P 109308]. bucia, Bulimus, P f e i f f e r , 1859: 39 ( B r a s i l i a ) . buckleyi, Bulimus (Drymaeus), Sowerby, 1895: 214, p i . 13 figs. 3-4 ( E c u a d o r) [ L T B M N H 1007.11.21.48]. canaliculars, Bulimus, P f e i f f e r , 1845a: 68 ( B o l i v i a ) [ L T B M N H 1975514]. canarius, Bulimus, P f e i f f e r , 1867: 76 ( [ P e r u ] T r u j i l l o ) . cantatus, Bulimus, Reeve, 1848 : p i . 56 f i g . 375 [no type locality g i v e n ] . carandaitiensis, Bulimulus (Drymaeus), Preston, 1907: 491, f i g . 4 (Carandaiti, province of C o r d i l l e r a , B o l i v i a , 1000 m ) . castaneostrigatus, Drymaeus, D a Costa, 1906b: 98, p i . 11 f i g . 5 ( E a s t e r n P e r u , P o z u z o ) [ H T B M N H 1007.11.21.19]. catamayensis, Mormus, M i l l e r , 1879: 120, p i . 12 f i g . 4 (Catamayo (prov. L o j a ) [Ecuador]). catenae, Drymaeus (Drymaeus), H a a s , 1952: 118, f i g . 20 ( P e r u , Dept. Cuzco, P r o v . Quispicanchi, H a c i e n d a Cadena, 1000 m ) [ H T F M N H 38121]. caucaensis, Bulimulus (Drymaeus), D a Costa, 1898: 81, p i . 6 f i g . 3 (Colombia, valley of the R . Cauca) [ H T B M N H 1007.11.21.43]. cecileae, Bulimus, J . M o r i c a n d , 1858: 452, p i . 14 f i g . 4 ( [ P e r u ] T a r a p o t o ) . celendinensis, Drymaeus, W e y r a u c h , 1956a: 151, p i . 11 figs. 10-11 ( N - P e r u , H ü g e l 2 k m w. Celendin, 2700 m ) [ H T S M F 155307]. chamaeleon, Bulimus, P f e i f f e r , 1855h: 116 ( Q u i t o [ E c u a d o r ] ) [ S T B M N H ] . championi, Otostomus, M a r t e n s , 1893: 222, p i . 14 f i g . 5 (Guatemala, H a c i e n d a de las Nubes, C e r r o Z u n i l ) [ L T B M N H 1001.6.22.451]. chanchamayensis, Bulimus, H i d a l g o , 1870: 49 (Chanchamayo, Pérou) [ S T Z M B ] . chaperi, Bulimulus, Crosse & Fischer, 1893: 31, p i . 1 figs. 1-2 ( M e x i q u e , île de Mescala, sur le lac Chapala dans l ' E t a t de J a l i s c o ) . chenui, Bulimus, P h i l i p p i , 1867: 72 ( [ P e r u ] Pachicama c prope L i m a ) . chiapasensis, Bulimus, P f e i f f e r , 1866: 81 (Chiapas, republica M e x i c a n a e ) . chiapensis, Otostomus, M a r t e n s , 1893: 205, p l . 15 f i g . 12 ( E . M e x i c o , C o r d o v a ) . chicoensis, Drymaeus fallax, Breure, 1977a : 261, f i g . 4 (Colombia, Cundinamarca, Bosque de Chico , 2700-2800 m ) [ H T S M F 245405]. chimborasensis, Bulimus, Reeve, 1848: p l . 44 f i g . 275 (Chimborazo, C o l u m b i a [sic, E c u a d o r ] , N e w Granada) [ S T B M N H ] . chiriquiensis, Drymaeus, D a Costa, 1901: 238, p l . 24 f i g . 1 (Panama , C h i r i q u i , Boqueti) [ H T B M N H 1007.11.21.119]. chrysomelas, Bulimulus (Thaumastus), M a r t e n s , 1867b: 145 [oberen A m a z o n e n s t r o m gebiets; S T Z M B ] . citrinellus, Bulimus scitulus, 'Philippi * P f e i f f e r , 1853d: 411 (inter M a c a n y a et f l u v i u m Maranon). clarus, Bulimus, P f e i f f e r , 1857a: 330 (Meobamba, P e r u ) . clathratus, Bulimus, P f e i f f e r , 1858: 258 ( P r o v . of Patas, A n d e s of P e r u ) [ L T B M N H 1975449].
114
ZOOLOGISCHE VERHANDELINGEN 168 (1979)
rugistriatus, Drymaeus (Drymaeus), H a a s , 1952: 120, f i g . 21 ( P e r u , Dept. Cuzco, P r o v . Quispicanchi, H a c i e n d a Cadena, 1000 m ) [ H T F M N H 38123]. rugosa, Lymnaea, Valenciennes, 1827: 250, p l . 56 f i g . 5 [no type locality g i v e n ] . saccatus, Bulimus, P f e i f f e r , 1855g: 94, p l . 31 f i g . 2 (Meobamba, E . P e r u ) [ L T B M N H 1975127]. sachsei, Bulimus, A l b e r s , 1854a : 30 ( C o l u m b i a australi [sic, P e r u ] ad f l u v i u m M a r a n h o n ) [ST Z M B ] . *sanctaemarthae, Drymaeus, P i l s b r y , 1901: 161, p l . 48 figs. 49-50 (Colombia, N W slope o f Santa M a r t a M t s . , Jiracasaca) . santanensis, Drymaeus poecilus, D a l l , 1912a : 4 ( P e r u , Santa A n a ) . *schmidti, Bulimus, P f e i f f e r , 1854a : 65 [no type locality g i v e n ] . schmidti, Drymaeus (Drymaeus), H a a s , 1955b: 314, f i g . 62 ( P e r u , Dept. Cuzco, M a r c a pata, Ccachabamba) [ H T F M N H 51323]. schunkei, Drymaeus (Drymaeus), H a a s , 1949: 237, f i g . 50b ( P e r u , Dept. L o r e t o , R i o U c a y a l i , C e r r o A z u l ) [ H T F M N H 30040]. scitulus, Bulimus, Reeve, 1849: p i . 71 f i g . 513 ( P e r u , Chachapoyas). scitus, Otostomus, H . A d a m s , 1867: 442, p i . 38 f i g . 5 ( E . P e r u ) . scoliodes, Drymaeus, Dautzenberg, 1901c: 309 ( R i o M i x i o l l o , province H u a l l a g a , P é r o u ) . selli, Bulimulus (Drymaeus), Preston, 1909: 511, p i . 10 f i g . 3 ( B r i t i s h Guiana) [ H T B M N H 1915.1.6.36]. semistriatus, Drymaeus (Drymaeus), H a a s , 1955a: 374, f i g . 77 ( B r a z i l , São P a u l o , V a r n h a g e n D i s t r . , F a z e n d a Ipanema) [ H T F M N H 49784]. serratus, Bulimus, P f e i f f e r , 1855g: 94, p l . 31 f i g . 6 ( P e r u , Moyobamba ) [ L T B M N H 1975475]. silvanus, Drymaeus (Orodrymaeus), Z i l c h , 1953: 56, p i . 14 f i g . 5 ( P e r u , [Dept. C a j a marca] B e r g u r w a l d der H a c i e n d a T a u l i s , 1700 m ) [ H T S M F 108567]. similaris, Bulimus, J. M o r i c a n d , 1856: 177, p i . 6 f i g . 8 ( [ P e r u ] M o y o b a m b a ) . siolii, Drymaeus (Drymaeus), H a a s , 1952: 108, f i g . 14 ( B r a z i l , Pará, Cacaual Grande, F u r o Piapó) [ H T F M N H 38170]. smithii, Bulimulus (Drymaeus), D a Costa, 1898: 81, p i . 6 f i g . 8 ( [ C o l o m b i a ] Bogota) [ L T B M N H 1907.11.21.52]. solidus, Bulimulus (Drymaeus), Preston, 1907: 494, f i g . 9 (Colombia, Bogota) [ST BMNH]. sophieae, Drymaeus (Drymaus). N e w name f o r Drymaeus (Drymaeus) pergracilis H a a s , 1952, not Bulimulus pergracilis R o l l e , 1905 [secondary h o m o n y m y ] . souzalopesi, Drymaeus (Drymaeus), W e y r a u c h , 1965: 74, figs. 4-5 ( B r a s i l , Estado de Goyaz, P l a n a l t i n a N o v a ) [ H T I M L 10622a]. spadiceus, Drymaeus, D a Costa, 1906b: 97, p l . 11 figs. 2-3 ( [ C o l o m b i a] Bogota) [ H T B M N H 1907.11.21.15]. spectatus, Bulimus, Reeve, 1849: p i . 81 f i g . 601a ( N e w Granada) [ L T BMNH 1874.12.11.226]. sporadicus, Bulimus, Reeve, 1848: p i . 49 f i g . 325 ( B o l i v i a , [ A r g e n t i n a ] P a t a g o n i a [sic]). steyermarki, Plecocheilus (Eurytus), H a a s , 1955a: 377, f i g . 79 (Venezuela, State o f B o l i v a r , northwest part o f Chimantá-massif, plateau below Apacará-tepui, 1800 m ) [ F M N H 49735]. stolli, Otostomus ghiesbreghti, M a r t e n s , 1893: 209, p i . 13 figs. 5-10 ( C e n t r a l Guatemala, L l a n o o f Quezaltenango, 6000 to 9000 feet). striyatus, Bulimus, Reeve, 1848 : p i . 44 f i g . 280 ( [ P e r u ] H u a l l a g a ) . strigatus, Bulinus, Sowerby, 1833a: figs. 95-96 ( P e r u , H u a l l a g a ) . subeffusus, Bulimus, P h i l i p p i , 1869 ' 36 ( i n nemoribus P e r u v i a e de H u a n c a y o dictis, loco Coyllorbamba). subhybridus, Gonyostomus, D a Costa, 1906b: 97, p i . 9 f i g . 1 ( E a s t e rn P e r u , P o z u z o ) [ H T B M N H 1007.11.21.127].
116
ZOOLOGISCHE VERHANDELINGEN 168 (1979)
sticht, Drymaeus (Drymaeus), H a a s , 1955b: 333, f i g . 73 ( P e r u , Dept. Tumbes, H u a s i m o , 220 m ) [ H T F M N H 51927]. zoogeographicus, Drymaeus, Z i s c h k a , 1953: 82 [emendation f o r zoographica d O r b i g n y ] . zoographica, Helix, d O r b i g n y , 1835: 19 (província Yungacensi, republica B o l i v i a n a ) .
Drymaeus (Mesembrinus)
A l b e r s , 1850
Mesembrinus A l b e r s , 1850: 157. T y p e species by subsequent designation ( A l b e r s , i860) : Helix virgulata Férussac. Antidrymaeus G e r m a i n , 1907: 59. T y p e species by subsequent designation ( P i l s b r y , 1926b) : Bulimulus (Drymaeus) inusitatus F u l t o n . Leptodrymaeus P i l s b r y , 1946a : 23. T y p e species by o r i g i n a l designation : Bulimus dominicus Reeve. Leptomormus W e y r a u c h , 1958: 136. T y p e species by o r i g i n a l designation: Drymaeus bequaerti W e y r a u c h . Diaphanomormus W e y r a u c h , 1964: 57. T y p e species by o r i g i n a l designation: Drymaeus (Diaphanomormus) coelestini obesus W e y r a u c h . Description. —
S h e l l elongate-ovate to ovate; n a r r o w l y p e r f o r a t e to rimate;
t h i n to rather solid. C o l o u r w h i t i s h to yellowish, u n i f o r m l y c o l o u r e d o r w i t h b r o w n i s h s p i r a l bands o r b r o w n i s h a x i a l streaks (that m a y be d i v i d e d into series o f spots). S u r f a c e s m o o t h o r w i t h incrassate g r o w t h striae a n d / o r fine s p i r a l lines. A p e r t u r e elongate- to subovate.
Peristome thin a n d simple
(to
expanded). Mandíbula w i t h m o r e t h a n 2 0 plates, w h i c h are ca. 8 times as l o n g as wide. T r a n s v e r s e r o w s o f the r a d u l a are V - to W - s h a p e d . C e n t r a l teeth t r i c u s p i d , w i t h lanceolate mesocones a n d but slightly smaller t r i a n g u l a r to ectocones.
L a t e r o m a r g i n a l teeth
t r i c u s p i d , shifted,
with
lanceolate
elongate-ovate
to
lanceolate mesocones, c u r v e d lanceolate endocones a n d t r i a n g u l a r to deltoid ectocones that m a y be b i f i d o r serrate. H a l f - r o w f o r m u l a : C / 3 (x =
+
L M
x/3
56-146).
P e n i s w i t h o r without a p r o x i m a l sheath, s u b c y l i n d r i c a l a n d p a s s i n g without external d i f f e r e n t i a t i o n into the epiphallus. T h e flagellum is s u b c y l i n d r i c al and
(usually)
relatively
short. V a g i n a relatively
long. Spermathecal
duct
t a p e r i n g , w i t h a m o r e o r less globose spermatheca o r reduced i n length, i n w h i c h case the spermatheca m a y be u n d i f f e r e n t i a t e d . Distribution. —
Venezuela,
Guiana, Surinam, F r e n c h Guyana,
Brazil,
P e r u , Colombia, Panama, Costa Rica, Nicaragua, E l Salvador, H o n d u r a s , G u a t e m a l a , B e l i z e , M e x i c o , U S A , W e s t Indies. Ecology. —
T h e species live o n trees a n d shrubs. T h e vertical d i s t r i b u t i o n
is o-ca. 1000 m . Taxa. —
T h e f o l l o w i n g t a x a are placed i n this subgenus ( t a x a
tentatively
r e f e r r e d to this subgenus are m a r k e d b y a n asterisk) : albescens, Bulimulus aureolus, Guppy, 1871: 308 [no type locality g i v e n ; T r i n i d a d ] . albicans, Bulimulus multifasciatus, M a z e 1874: 163 ( [ W e s t Indies, M a r t i n i q u e ] Saint P i e r r e , hauteurs du Parnasse, 200 mètres environs).
BREURE, BULIMULINAE fidustus, Bulimus, Reeve, 1849: p l . 76 f i g . 557 ( N e w Granada, Sebundoi) [ L T
119 BMNH
1975517]. flavescens, Helix liliacea, Férussac, 1821: 54 (les A n t i l l e s , P o r t o R i c o ) . flavidulus, Bulimus (Liostracus), E . A . S m i t h , 1877b: 364, p i . 39 f i g . 3 ( S o u t h E c u a d o r , Z a r a m a ) [ L T B M N H 1975134]. flavidus, Bulimus, M e n k e , 1829: 6 [not seen]. flavotinctus, Drymaeus vincentinus, P i l s b r y , 1899: 18, p i . 12 f i g . 11 ( [ W e s t Indies] T r i n i d a d ) [ H T A N S P 25859]. floridanus, Bulimus, P f e i f f e r , 1857a: 330 ( F l o r i d a ) [ L T B M N H 1975*99] · floridianus, Bulimus, Binney, 1859: 134 ( F l o r i d a ) . fluctuatus, Bulimulus, Beck, 1837: 66 [indication]. *funeralis, Bulimus, Bruguière, 1789: 321 (l'intérieur de l'Amérique méridionale). fuscobasis, Bulimus (Liostracus), E . A . S m i t h , 1877b: 365, p l . 39 f i g . 6 (Andes o f P e r u , Tarapoto) [ L T B M N H 1975139]. gabbi, Bulimus, A n g a s , 1879: 477, p l . 40 f i g . 3 ( [ C o s t a R i c a ] P i c o Blanco, 3000-6000 ft) [ L T B M N H 1879.7.22.23]. gabbianus, Bulimulus, Binney, 1884: 124, p l . 12 f i g . F (Costa R i c a ) . gereti, Drymaeus, A n c e y , 1901a: 93 ( [ B r a z i l ] P r o v . de G o y a z ) . gorgonensis, Drymaeus, H a a s , 1966: 233, f i g . 49 (Colombia, Cauca, Island o f Gorgona) [ H T F M N H 114164]. gracilior, Otostomus (Drymaeus) sulfureus, M a r t e n s , 1893: 225 (Guatemala, N i c a r a g u a ) . gratus, Bulimus (Mesembrinus), P f e i f f e r , 1856a: 159 [ N e w name f o r Bulimus columbiensis P f e i f f e r , 1855, not L e a , 1838]. grenadensis, Bulimus, P f e i f f e r , 1848a : 231 [no type locality g i v e n ] . griffini, Drymaeus (Drymaeus), H a a s , 1955a: 383, f i g . 83 (Venezuela, western side o f Abacapa-tepui, Chimantá-massif, State o f Bolívar, 1300 m ) [ H T F M N H 49734]. gruneri, Bulimus, P f e i f f e r , 1846a: 30 ( M e x i c o ) . guttulatus, Bulimus knorri, Schaufuss, 1881: 178 (Venezuela). hachensis, Bulimus, Reeve, 1850a : p l . 85 f i g . 627 (Guatemala, banks o f the R i o H a c h a ) [ L T B M N H 1975392]. haitensis, Drymaeus sallei, P i l s b r y , 1899: 12, p l . 39 f i g . 4 ( H a i t i , P o r t - a u - P r i n c e , F o r t Jacques) [ L T A N S P 3552]. havanensis, Drymaeus multilineatus, Jaume & B o r r o , 1941: 404 (Cuba, província de l a H a b a n a , cerca de S a n F r a n c i s c o de P a u l a , Reparto " D i e z m e r o " ) . hegewischi, Bulimus, P f e i f f e r , 1842: 46 ( M e x i c o , M i c h o a c a n , P a r q u a r o ) . hemphilli, Bulimulus, W r i g h t , 1889: 8 [indication]. herrerae, Drymaeus, B a r t s c h, 1907: 119 (Bonanz a Z i m a p a n , H i d a l g o , M e x i c o ) [ H T U S N M 192992]. heterogeneus, Bulimus, P f e i f f e r , 1866b: 83 ( [ M e x i c o ] i n regione "savannarum" prope Veracruz). heynemanni, Bulimus, P f e i f f e r , 1866b: 83 ( M e x i c o , O r i z a b a ) . hjalmarsoni, Bulimus, P f e i f f e r , 1856b: 51 (in insula P o r t o r i c o , plantatione " P a j a s " prope M a n a t i ) . hoffmanni, Otostomus tripictus, M a r t e n s , 1893: 225, p l . 14 figs. 1 1 - n a ( C e n t r a l Costa R i c a , woods of S a n L o r e n z o de Bota, 1300 m ) . honduranus, Otostomus, M a r t e n s , 1893: 232 ( H o n d u r a s ) . hondurasanus, Bulimus, P f e i f f e r , 1846a: 29 ( H o n d u r a s ) [ L T B M N H 1975265]. honduratianus, Bulimus, T r i s t a m , 1861: 230 (Guatemala). hyalinoalbida, Bulimulus (Drymaeus) moricandi, Crosse & Fischer, 1875 [1870-1894] : 497, p l . 24 figs. 9'9a (in província Chiapas, reipublicae M e x i c a n a e ) . hypozonus, Otostomus (Drymaeus) palpaloensis, M a r t e n s , 1893: 223 ( E a s t M e x i c o [several localities mentioned]). ictericus, Otostomus depictus, M a r t e n s , 1873: 183, p i . 1 figs. 16-17 ([Venezuela] Caracas).
I20
ZOOLOGISCHE VERHANDELINGEN 168 (1979)
immaculatus, Bulimus, C . B . A d a m s i n Reeve, 1850: p l . 85 f i g . 631 (Jamaica) [ L T B M N H 1975540]. imperfectus, Bulimus multifasciatus, Guppy, 1866a: 49 (southern parts of the island o f Trinidad). incarnatus, Bulimus, P f e i f f e r , 1855g: 95 (Venezuela) [ L T B M N H 1975566]. indistinctus, Bulimulus, Guppy, 1868a: 436 [ W e s t Indies, G r e n a d a ] . indistinctus, Bulimus, P f e i f f e r , 1852a: 63 [ N e w name f o r Bulimus monilifer Reeve, 1848, not Bulimus moniliferus G o u l d , 1846]. inglorius, Bulimus, Reeve, 1848: p i . 55 f i g . 368 [no type locality g i v e n ; L T B M N H 1975536]. insulaecygni, Drymaeus, Clapp, 1914: 98, p i . 6 f i g . 5 ( [ W e s t Indies] S w a n Island) [ H T M C Z 22877]. interpunctus, Bulimulus, M a r t e n s , 1887: 161 (Piracicaba, S ã o P a u l o , B r a s i l i e n ) [ H T Z M B 38952]. interruptofasciatus, Drymaeus, Vernhout , 1914a: n , p i . 1 figs. 5-6 ( S u r i n a m , environs of Paramaribo) [ H T R M N H ] . interruptus, Bulimulus (Drymaeus), Preston, 1909: 511, p i . 10 f i g . 1 (Venezuela, M e r i d a ) [ H T B M N H 1914.4.3.38]. interstitialis, Otostomus ghiesbreghti, M a r t e n s , 1893: 209, p i . 13 figs. 5-10 ( C . Guatemala, Cumbr e de S a n M a r t i n , on the S W slope o f the C o r d i l l e r a , 6000 feet). intrapictus, Drymaeus, P i l s b r y , 1930b: 341, p i . 30 f i g . 8 ( P a n a m a, L o s Santos P r o v . , T o n o s i ) [ H T A N S P 140834]. inusitatus, Bulimulus (Drymaeus), F u l t o n , 1900: 87 (Costa R i c a ) [ H T BMNH 1901.4.25.28]. irazuensis, Bulimus, A n g a s , 1878a: 73, p i . 5 figs. 17-20 (volcano o f Irazu, Costa R i c a ; Mexico). jonast, Bulimus, P f e i f f e r i n P h i l i p p i , 1847: 125, p i . 5 f i g . 4 ( A m e r i c a e centralis, V e r a Cruz) [ S T B M N H ] . juquilensis, Drymaeus alternans, ' M a r t e n s ' P i l s b r y , 1899: 88, p i . 15 f i g . 40 ( S . M e x i c o , State o f O a x a c a , J u q u i l a ) . kaemmereri, Bulimus, Mörch, 1852: 23 [nomen nudum] . keppelli, Bulimus, P f e i f f e r , 1853d: 654 (in A n d i b u s P e r u v i a n u s ) . knorri, Bulimus, P f e i f f e r i n P h i l i p p i , 1846: 115, p i . 4 f i g . 3 ( L a G u y a n a ) . koppeli, Bulimus, Sowerby, 1892: 297, p i . 23 figs. 9-12 ( [ C o l o m b i a ] Bogota) [ L T B M N H 1907.11.21.133]. lacteus, Bulimus, L e a , 1838: 65, p i . 23 f i g . 100 (about one hundred miles up the Magdalena R i v e r , Colombia) [ L T A N S P 192930]. laetus, Bulimus, Reeve, 1849: p i . 83 f i g . 616 ( N e w Granada, Sebundoi) [ L T B M N H 1975534]. lascellesiana, Bulimulus (Drymaeus) binominis, E . A . S m i t h , 1895: 316, p i . 21 f i g . 14 ( [ W e s t Indies] Grenada, 1000-2000 feet). laticinctus, Bulimulus, Guppy, 1868a: 431 ( [ W e s t Indies] D o m i n i c a ) . latizonatus, Drymaeus multilineatus, P i l s b r y , 1936: 69 ( F l o r i d a , L o w e r Matecumbe K e y ) [ L T A N S P 160880a]. lentiginosus, Bulimus, P h i l i p p i , 1869: 32 ( [ P e r u ] inter C a j a m a r c a et Contumaza). leucomelas, Bulimus, A l b e r s , 1854b: 219 ( C o l u m b i a [sic, P e r u ] ad f l u v i u m M a r a n h o n ) [ H T Z M B 101785]. liliacea, Helix, Férussac, 1821: 54 (les A n t i l l e s , P o r t o R i c o ) . limpidus, Bulimus, Drouêt, 1859: 64, p i . 2 figs. 23-24 (Guyane française, Ilet-la-Mère). lineolatus, Bulimus, Conrad, 1856: 32 (volcano o f Cartago, Costa R i c a ) . lineolatus, Otostomus recluzianus, M a r t e n s , 1893: 213 ( C e n t r a l Costa R i c a , S a n José). lirinus, Bulimus, Morelet, 1851: 11 ([Guatemala] Petenensium S a n L u i s ) [ S T B M N H ] . lituratus, Bulimulus fluctuatus, Beck, 1837: 66 [indication].
121
BREURE, BULIMULINAE
livescens, Bulimus, P f e i f f e r , 1842: 175 ( M e x i c o ) . lividus, Bulimus, Reeve, 1850: p l . 85 f i g . 626 (Venezuela) [ L T B M N H 1975208]. loxanus, Otostomus, H i g g i n s , 1872: 685, p i . 56 figs. 2-2a ( [ E c u a d o r ] L o x a ) [ L T B M N H 1975552]. loxensis, Bulimus, P f e i f f e r , 1846c: 85 ( E l Catamaija prope L o x a reipublicae A e q u a torius) [ L T B M N H 1975553]· lucidus, Bulimus, Reeve, 1848: p i . 40 f i g . 245 ( [ W e s t Indies] St. V i n c e n t s ) [ S T B M N H ] . lusorius, Bulimus, P f e i f f e r , 1855c: 291 ( B a n k s of A m a z o n , B r a z i l s ) [ L T B M N H 1975543]. lynchi, Drymaeus, P a r o d i z , 1946b: 1, p i . 1 figs. 1-3
( B o l i v i a , P o z o de V a r g a s )
[HT
M A C N 1344]. maculatus, Bulimus, L e a , 1838: 86, p i . 23 f i g . 112 (near Carthagena [ C o l o m b i a ] ) . manupictus, Bulimus, Reeve, 1848: p i . 55 f i g . 369 (Andes of Columbia) [ L T B M N H 1975522]. marielinus, Bulimus, Poey, 1851: 204 [Cuba, teste P i l s b r y , 1899]. martensianus, Drymaeus recluzianus, P i l s b r y , 1899: 56 ( N e w name f o r Otostomus recluzianus lineolatus M a r t e n s , 1893, not Bulimus lineolatus Conrad, 1856). mayaorum, Drymaeus, Rehder, 1966: 287, figs. 3-4 (Isla M u j e r e s , Quintana R o o , M e x i c o ) [ H T U S N M 251656]. membranaceus, Bulimus, P h i l i p p i , 1846: 126, p i . 5 f i g . 2 [no type locality given]. membranaceus, Bulimus, Reeve, 1849 : p i . 75 f i g . 544. membranaceus, Otostomus (Mormus), M a r t e n s , 1873: 186 (Venezuela, Caracas). menkei, Bulimus, Gruner, 1841: 277, p l . 11 f i g . 2 (Reipublicae Venezuela, prov. O r i n o c o ) . meridanus, Bulimus, P f e i f f e r , 1846a: 33 ( M e r i d a , B o l i v i a n A n d e s [sic]) [ S T B M N H ] . mexicanus, Bulimus, Reeve, 1848 : p i . 40 f i g . 244 ( M e x i c o ) . miliaris, Bulimus, P h i l i p p i , 1867: 74 ( [ P e r u , Dept. L a L i b e r t a d ] hacienda de U n i g a m b a l ) . miltochrous, Bulimus, A l b e r s , 1854b: 217 ( C o l u m b i a [sic, P e r u ] ad f l u v i u m M a r a n h o n ) [ H T Z M B 101791]. misellus, Drymaeus translucens, P i l s b r y , 1926b : 83, f i g . 14b (Panama, P r o v . L o s Santos, Tonosi) [ H T A N S P 140281a]. modesta, Bulimus knorri, Schaufuss, 1881: 178 (Venezuela). monachus, Bulimus, P f e i f f e r , 1857a: 333 (Meobamba, P e r u ) . monilifer, Bulimus, Reeve,
1848: p i . 48 f i g . 318 [no type locality g i v e n ; L T
BMNH
1975403]. montagnei, Bulimus, d O r b i g n y , 1836: 286, p i . 32 figs. 5-7 ( [ B o l i v i a ] principalmente à la côte de Petaca [ = Dept. Santa C r u z , P r o v . F l o r i d a , Cuevas Petacas, ca. 23 k m E Samaipata]). montagnei, Bulimus, Reeve, 1848: p l . 23 f i g . 146 ( B o l i v i a , C h i l o n ) [not montagnei dOrbigny]. montaguae, Otostomus sargi, M a r t e n s , 1893: 218 ( C . Guatemala, valley of the R i o Montagua). montanus, Drymaeus roseatus, P i l s b r y , 1901: 161, p i . 48 f i g . 51 (Colombia, western part of Santa M a r t a M t s . , L a s P a n t i d a s ) . *morenoi, Bulimulus (Drymaeus), P r e s t o n, 1007: 494, f i g . 7 ( A r g e n t i n a ) . moricandi, Bulimus, P f e i f f e r , 1847a: 113 ( M o u n t Coban, C - A m e r i c a ) [ L T B M N H 1975212]. moritinctus, Otostomus, M a r t e n s , 1893: 228, p i . 14 figs. 9-10 ( M e x i c o , State of Guerrero, Chilpancingo) [ L T B M N H 1901.6.22.841]. mossi, Bulimulus (Drymaeus), E . A . S m i t h , 1896: 243, p i . 8 f i g . 8 ( [ W e s t Indies] Trinidad). moussoni, Bulimus, P f e i f f e r , 1853a: 147 ( [ W e s t Indies] St. D o m i n g o ) [ L T B M N H 1975210]. *muliebris, Bulimus, Reeve, 1849: p i . 81 f i g . 598 ( N e w Granada) .
124
ZOOLOGISCHE VERHANDELINGEN 168 ( 1979)
sporlederi, Bulimus, P f e i f f e r , 1866: 83 ( [ M e x i c o ] M i r a d o r prope V e r a c r u z ) . stigmaticus, Bulimus, P h i l i p p i , 1867: 74 ( [ P e r u , Dept. L a L i b e r t a d ] hacienda de Unigambal). stramineus, Bulimulus, G u i l d i n g , 1824: 340 ( [ W e s t Indies] S ti V i n c e n t i i ) . studeri, Bulimus, P f e i f f e r , 1847a: 112 (Venezuela, M e r i d a ) [ L T B M N H 1975480]. subfasciatus, Bulimulus dormani, Cockerell, 1891: 18 [no type locality g i v e n ] . subfasciatus, Bulimulus fluctuatus, Beek, 1837 : 66 [nomen n u d u m ]. subfloccosus, Drymaeus translucens, P i l s b r y , 1809: 90, p l . 24 figs. 26-27 ( N i c a r a g u a and H o n d u r a s ) [ L T A N S P 25949a]. subpellucidus, Bulimus (Liostracus), E . A . S m i t h , 1877b: 364, p l . 39 f i g . 2 ( E c u a d o r ) . subunicolor, Otostomus fenestrellus, M a r t e n s , 1893: 215 [no type locality g i v e n ] . sulphureus, Bulimus, P f e i f f e r , 1857a: 318, p i . 35 f i g . 11 ( [ M e x i c o ] C o r d o v a ) . surinamensis, Drymaeus, Vernhout , 1914a: 13, p i . 1 f i g . 3 ( S u r i n a m , P o s t G r o n i n g e n) [HT R M N H ] . sykesi, Drymaeus, D a Costa, 1906a: 7, p i . 1 f i g . 1 ( [ C o l o m b i a ] Bogota) [ H T B M N H 1907.11.21.4]. tenuilabris, Bulimus, P f e i f f e r , 1866a: 831 (Venezuela) [ L T B M N H 1975338]. tepecensis, Otostomus uhdeanus, M a r t e n s , 1893: 233, p i . 15 figs. 1-6 ( W e s t M e x i c o , Tepic, State of J a l i s c o ) . tonosiensis, Drymaeus translucens, P i l s b r y , 1930b: 340 ( P a n a m a, L o s Santos P r o v . , T o n o s i ) [ H T A N S P 151288]. tortugensis, Drymaeus bahamensis, P i l s b r y & Vanatta, 1928: 477, p i . 27 figs. 11-13 ( [ W e s t Indies] H a i t i , T o r t u g a Island) [ H T A N S P 146700a]. totonacus, Bulimulus, Strebel i n Strebel & P f e f f e r , 1882: 84, p l . 5 figs. I3-I3a ( [ M e x i c o ] Rancho de Quilate und Aguacaliente, U m g e g e n d M i s a n t l a ) . translucens, Bulinus, B r o d e r i p i n B r o d e r i p & Sowerby, 1832a: 31 (in A m e r i c a m e r i dionali, K i n g s and Saboga Islands, B a y o f P a n a m a ) . *tribalteatus, Bulimus, Reeve, 1848 : p l . 43 f i g . 269 (Santa F é d i Bogota) . tricingulatus, Bulimus, A n t o n , 1839: 43 (Saboja Islands). tricolor, Bulimus knorri, Schaufuss, 1881: 178 (Venezuela). trigonostomus, Bulimus, Jonas, 1844: 36 ( P r o v i n c i a e Cumana, Reipublicae Venezuela). trimarianus, Otostomus, M a r t e n s , 1893: 216, p l . 13 f i g . 17 ( M e x i c o , T r e s M a r i a s Islas) [ L T B M N H 1901.6.22.950]. trinitarius, Bulimulus (Drymaeus), E . A . S m i t h , 1896: 242, p l . 8 f i g . 7 ( [ W e s t Indies] Trinidad). tripictus, Bulimus, A l b e r s , 1857 : 97 (Costa R i c a ) . tristis, Bulimus, P f e i f f e r , 1855b: 124 ( N e w Granada) [ S T B M N H ] . tropicalis, Bulimus, Morelet, 1849: 9 ( [ M e x i c o ] ad plagam civitatis Campeche) [ L T B M N H 1893.2.4.210]. tryoni, Bulimulus (Thaumastus), Crosse & Fischer, 1875 [1870-1894] : 565 ( [ M e x i c o ] província T a b a s c o ; P a p a n t l a ; M i s a n t l a ; O a j a c a ; C i n a l o a ) . uhdeanus, Bulimulus (Mesembrinus), M a r t e n s , 1864: 541 [no type locality g i v e n ; S T ZMB]. umbraticus, Bulimus, Reeve, 1849: p i . 77 f i g . 559 ( C e n t r a l A m e r i c a ) [ L T B M N H 1975184]. undulatus, Bulimulus, G u i l d i n g , 1828b: 169 ( [ W e s t Indies] S t i V i n c e n t i i , radices montis "Bon Homme"). unicolor, Otostomus lilacinus, M a r t e n s , 1893: 192 ( W . Guatemala). venezuelensis, Otostomus, M a r t e n s , 1893: 224 ( M i r a d o r , M e x i c o ) . venosus, Bulimus, Reeve, 1848: p i . 45 f i g . 285 ( A n g o s t u r a, B a n k s o f the O r i n o c o ) . veracruzensis, Drvmaeus herrarae, Bartsch , 1907: 119 (Cordova, V e r a C r u z , M e x i c o ) [ H T U S N M 192093]. vernhouti, Drymaeus, Breure, I976d: i n , f i g . 4 ( N e w name f o r Drymaeus quadrifasciatus Vernhout, 1914, not Bulimus quadrifasciatus A n g a s , 1878).
BREURE, BULIMULINAE
125
vexillum, Bulinus, B r o d e r i p i n B r o d e r i p & Sowerby, 1832b : 105 ( S i n u Panamae, K i n g s and Saboga I d ) . vincentinus, Bulimus, P f e i f f e r , 1846a: 30 ( [ W e s t Indies] St. V i n c e n t ) [ L T B M N H 1975219]. virginalis, Bulimus, P f e i f f e r , 1856b: 46 ([Venezuela] C a r a c a s ) . virgo, Bulimus, L e a , 1838: 84, p l . 23 f i g . 97 (near Carthagena [ C o l o m b i a ] ) . virgulata, Helix, Férussac, 1821: 54 [indication]. waldoschmitti, Drymaeus, P a r o d i z , 1962: 436, p l . 2 f i g . 16 ( P e r u ) [ H T U S N M 609317]. wintlei, Drymaeus, F i n c h , 1929: 275, figs. ( E c u a d o r ) [ H T B M N H 1929.6.11.1]. xantholeucus, Otostomus castus, M a r t e n s , 1893: 206, p l . 12 figs. 16-21 ( N . Guatemala, Saboma, valley o f the r i v e r P o l o c h i c , 3800 feet). ziegleri, Bulimus, P f e i f f e r , 1847a: 113 [no type locality g i v e n ] . ziegleri, Bulimus, Reeve, 1849 : pi- 58 f i g . 389 ( C e n t r a l A m e r i c a ) .
Sphaeroconcha B r e u r e ,
1978
Sphaeroconcha B r e u r e , 1978b: 153. T y p e species by m o n o t y p y : Bulimulus araozi W e y r a u c h .
(Bulimulus)
D e s c r i p t i o n . — S h e l l globose; r i m a t e; t h i n . C o l o u r u n i f o r m b r o w n . S u r f a c e w i t h e p i d e r m a l s p i r a l striae. P r o t o c o n c h pit-reticulate. W h o r l s rather c o n v e x ; suture
w e ll
impressed.
Aperture
s u b c i r c u l a r,
relatively
large.
Peristome
slightly t h i c k e n e d, simple. T h e central teeth o f the r a d u l a a r e t r i c u s p i d , w i t h v e r y acute, conical to lanceolate mesocones
a n d relatively small, acute, t r i a n g u l a r ectocones. T h e
l a t e r o m a r g i n a l teeth a r e b i - to m u l t i c u s p i d , w i t h acute, elongate t o lanceolate mesocones a n d 1-5 acute, t r i a n g u l a r to deltoid ectocones. H a l f - r o w f o r m u l a : C / 3 + L M X / 2 - 6 ( x = 53). P e n i s w i t h o u t a sheath, m o r e o r less s u b c y l i n d r i c a l . Distribution. — P e r u (Dept. Ecology. —
Huánuco).
T h e species lives i n t r o p i c a l r a i n forest, p o s s i b ly o n leaves, at
500-700 m . Relationships. —
T h i s genus
is no t closely
related to a n y o f the other
b u l i m u l i d genera. It is characterized b y the globose shell shape, the sculpture o f the p r o t o c o n c h , the s u b c i r c u l a r shape o f the aperture, the absence o f a penis sheath, the absence o f parallel tubes i n the penis a n d the structure o f the r a d u l a . T h e o n l y k n o w n t a x o n is: araozi, Bulimulus (Bulimulus), W e y r a u c h , 1956a: 149, p i . 11 f i g . 8 ( M - P e r u , T i n g o M a r i a , 670 m , auf der l i n k e n Seite des R i o H u a l l a g a ) [ H T S M F 155304].
Leiostracus A l b e r s ,
1850
Leiostracus A l b e r s , 1850: 156. T y p e species by subsequent designation ( A l b e r s , i 8 6 0 ) : Bulimus vittatus S p i x . Liostracus Mörch, 1852: 26 [emendation f o r Leiostracus A l b e r s ] .
126
ZOOLOGISCHE VERHANDELINGEN 168 (1979)
Description. — forate;
S h e l l ovate-conical;
w i t h straight sides;
(narrowly)
per-
( r a t h e r ) thin. S u r f a c e smoot h o r w i t h e p i d e r m a l s p i r a l striae. P r o t o -
conch w i t h a x i a l w r i n k l e s a n d / o r fine s p i r a l lines. W h o r l s n e a r l y flat to slightly c o n v e x , last w h o r l m o r e o r less keeled;
suture h a r d l y impressed.
A p e r t u r e oblique, sub- to elongate-ovate. P e r i s t o m e n a r r o w l y e x p a n d e d . Distribution. — Guyana, Surinam, Brazil. K e y to the subgenera o f a.
Leiostracus
S h e l l surface smooth; p r o t o c o n c h w i t h fine s p i r a l lines a n d occasionally some l o w a x i a l w r i n k l e s o n u p p e r part o f w h o r l ; lateral teeth o f the radula bicuspid
b.
Leiostracus
(Leiostracus)
S h e l l surface mostly w i t h e p i d e r m a l s p i r a l striae; p r o t o c o n c h w i t h a x i a l w r i n k l e s a n d fine s p i r a l lines;
lateral teeth o f the r a d u l a m o n o c u s p i d Leiostracus
(Pseudoxychona)
Leiostracus (Leiostracus) A l b e r s , 1850 Description. — uniformly
S h e l l perforate ;
rather t h i n . C o l o u r w h i t i s h to y e l l o w i s h,
c o l o u r e d o r w i t h a x i a l a n d / o r s p i r a l b r o w n i s h bands.
Surface
smooth. P r o t o c o n c h w i t h fine s p i r a l lines a n d occasionally some l o w a x i a l wrinkles o n the u p p e r part o f w h o r l . A p e r t u r e elongate-ovate.
Peristome
narrowly expanded. T h e central teeth o f the r a d u l a are m o n o c u s p i d , relatively s m a l l , w i t h blunt, elongate to t r i a n g u l a r mesocones. L a t e r a l teeth a r e b i c u s p i d , w i t h spatulashaped to truncate mesocones a n d t r i a n g u l a r to deltoid ectocones. T h e m a r g i n a l teeth a r e t r i c u s p i d , shifted, w i t h elongate mesocones, c u r v e d , elongate endocones a n d t r i a n g u l a r ectocones. H a l f - r o w f o r m u l a : C / i + L x / 2 +
M
y / 3 ( x = 8, y = 17-29). T h e p e r i c a r d is ca. %
the l e n g t h o f the n e p h r i d i u m , w h i c h is n a r r o w l y
t r i a n g u l a r. T h e m a i n p u l m o n a r y v e i n is well developed, b u t n o t p r o m i n e n t , while the side veins a r e w e a k l y developed. T h e adrectal ureter is closed o v e r its entire length. T h e penis is without a sheath, m o r e o r less s u b c y l i n d r i c a l a n d p a s s i n g without external d i f f e r e n t i a t i o n into the epiphallus. T h e flagellum is relatively short. T h e p r o x i m a l part o f the spermathecal duct is thick, w i t h a spermathecal
appendix;
the distal part o f the spermathecal
duct is l o n g a n d
n a r r o w . T h e spermatheca is (elongate-)globose. D i s t r i b u t i o n . — G u y a n a , S u r i n a m , B r a z i l ( B a h i a , Espírito S a n t o ) . Ecology. —
T h e species live o n trees. T h e vertical d i s t r i b u t i o n is o-ca.
500 m . R e m a r k s . — P i l s b r y (1899) was the first to restrict the name
Leiostracus
128
ZOOLOGISCHE VERHANDELINGEN 168 ( l Ç 7 9 )
unicolor, Helix (Cochlogena) vittata, S. M o r i c a n d , 1836: 433 ( [ B r a z i l , B a h i a ] les forêts de Illheos). viminea, Helix (Cochlogena), S. M o r i c a n d , 1833: 540, p l . 1 f i g . 5 (Brésil, province de Bahia) [ S T Z M B ] . vitreus, Bulimus, S p i x , 1827 : p l . 8 f i g . 2 ( B r a s i l i a ) . vittatoezonata, Helix (Cochlogena) coxeirana, S. M o r i c a n d , 1836: 433 ( [ B r a z i l , B a h i a ] les forêts de Illheos). vittatozonata, Helix (Cochlogena) vittata, S. M o r i c a n d , 1836: 432 ( [ B r a z i l , Bahia] les forêts de Illheos). vittatus, Bulimus, S p i x , 1827: 7, p l . 7 f i g . 4 ( [ B r a z i l ] s y l v i s . . . P r o v i n c i a r u m Bahiensis et Pernambucanae). zebra, Bulimus, S p i x , 1827: p l . 7 f i g . 5 ( [ B r a z i l ] sylvis mediterraneis P r o v i n c i a e Bahiensis).
Leiostracus (Pseudoxychona) Pseudoxychona P i l s b r y , 1930c: 356. T y p e spiritualis Ihering.
P i l s b r y , 1930
species by o r i g i n a l designation:
Oxychona
D e s c r i p t i o n . — S h e l l ovate-conical; w i t h straight sides; n a r r o w l y perforate to rimate; rather t h i n . C o l o u r y e l l o w i s h to b r o w n i s h , u n i f o r m l y c o l o u r e d o r w i t h d a r k e r s p i r a l b a n d ( s ) . S u r f a c e w i t h e p i d e r m a l s p i r a l striae. P r o t o c o n c h w i t h s p i r al lines a n d a x i a l riblets, w h i c h a r e ca. three times as s t r o n g as the lines. W h o r l s h a r d l y to slightly convex, the last w h o r l keeled; suture h a r d l y impressed. A p e r t u r e oblique, truncate-ovate.
P e r i s t o m e h a r d l y to n a r r o w l y
expanded. T h e central teeth o f the r a d u l a a r e m o n o c u s p i d , w i t h deltoid to t r i a n g u l a r mesocones
o f w h i c h the a p e x is acute a n d pointed. T h e lateral teeth a r e
m o n o c u s p i d , w i t h acute a n d pointed, t r i a n g u l a r mesocones. t r i c u s p i d , shifted, w i t h elongate mesocones, t r i a n g u l a r ectocones.
slightly
H a l f - r o w formula: C / i +
M a r g i n a l teeth
c u r v e d endocones a n d
L x/i +
M y / 3 ( x = 8,
y = 16-18). T h e p e r i c a r d i u m is 1/2-4/5 * h length o f the n e p h r i d i u m , w h i c h is n a r r o w l y e
triangular. T h e m a i n p u l m o n a r y v e i n is moderately, the side veins a r e weakly to moderately developed. T h e adrectal ureter is closed over its entire length. P e n i s without a sheath, s u b c y l i n d r i c a l a n d slightly swolle n at the transitio n to the epiphallus, w h i c h is ca. h a l f as l o n g as the penis. T h e flagellum is relatively v e r y short. T h e v a g i n a is rather long. T h e p r o x i m a l p a rt o f the spermathecal duct is s u b c y l i n d r i c a l a n d rather thick, w i t h a n elongate s p e r m a thecal a p p e n d i x ;
the distal part o f the duct is n a r r o w . T h e spermatheca is
elongate-ovate. D i s t r i b u t i o n . — B r a z i l ( B a h i a , Espírito S a n t o ) . Ecology. — U n k n o w n . Remarks. —
Z i l c h ( i 9 6 0 ) considered this t a x o n a subgenus o f
Bulimulus.
T h e anatomy, however, demonstrates that this classification is incorrect a n d
BREURE, BULIMULINAE
129
F i g s . 161-163. V a r i a t i o n i n shell shape i n Leiostracus. F i g . 161. L. (Pseudoxychona) spec. F i g . 162. L. (L.) perlucidus ( S p i x ) . F i g . 163. L. (L.) onager ( B e c k ) . Scale = 5 m m . F i g . 164. G e n i t a l i a of Leiostracus (Pseudoxychona) spiritualis ( I h e r i n g ). A f t e r a d r a w i n g by courtesy o f D r . H . E . B . Rezende. Scale = 2 m m . F i g s . 165-166. V a r i a t i o n i n shell shape i n Simpulopsis. F i g . 165. S. (S.) decussata P f e i f f e r . F i g . 166. S. (S.) atrovirens ( M o r i c a n d ) . 9
130
ZOOLOGISCHE V E R H A N D E L I N G E N 168
that Pseudoxychona
(1979 )
is m o r e p r o p e r l y placed w i t h Leiostracus,
Rhinus
and
See
phylogenetic
Simpulopsis. Relationships. —
page
relationships. T h e subgenus
i6iff
for a discussion of
the
is characterized b y the sculpture o f the p r o t o -
conch , the shape o f the aperture a n d the structure o f the r a d u l a . Bibliography. —
T h e m a i n publications are: I h e r i n g , 1912;
P i l s b r y , 1930c.
T a x a . — T h e f o l l o w i n g t a x a are i n c l u d e d i n this subgenus : dulcis, Oxychona pileiformis, Ihering , 1912: 485, p i . 42 f i g . 14 [ B r a z i l , E s p i r i t o S a n t o ; teste M o r r e t e s , 1949]. pileiformis, Helix, S. M o r i c a n d , 1836: 420, p i . 2 f i g . 2 ( [ B r a z i l , B a h i a ] Illheos) [ S T MHNG]. polytricha, Oxychona, Ihering, 1912: 486, p i . 42 figs. 16-18 ( [ B r a z i l ] E s p i r i t o Santo, Cachoeira a m R i o Doce) [ H T D Z S P ] . spiritualis, Oxychona, Ihering, 1912: 485, f i g . 1, p i . 41 figs. 10-13, p i . 42 f i g . 15 ( [ B r a z i l ] E s p i r i t o Santo, Cachoeira a m R i o D o c e ) [ H T D Z S P ] .
Rhinus A l b e r s ,
i860
Rhinus A l b e r s , i 8 6 0 : 223. T y p e species by o r i g i n a l designation: Bulimus heterotrichus Moricand. Description. —
Shell
(elongate-)ovate to globose;
r a t h e r t h i n to solid. C o l o u r b r o w n i s h to y e l l o w i s h . incrassate g r o w t h
perforate Surface
to
rimate;
with
slightly
striae a n d s p i r a l series o f h a i r s . P r o t o c o n c h w i t h
w a v e d o r z i g z a g w r i n k l e s . A p e r t u r e ( r o u n d e d ) ovate. P e r i s t o m e
axial
expanded
and usually narrowly reflexed. T h e c e n t r al teeth o f the r a d u l a a r e m o n o c u s p i d , relatively s m a l l , w i t h b l u n t elongate to t r i a n g u l a r mesocones. L a t e r o m a r g i n a l teeth are b i - to t r i c u s p i d , w i t h b l u n t spatula-shaped mesocones, acute, c u r v e d elongate endocones a n d ( r a t h e r ) acute, ovate to t r i a n g u l a r ectocones, w h i c h are p o s t e r i o r l y situated o n the basal plate. H a l f - r o w f o r m u l a : C / i + L M x / 2 - 3
(x =
32).
T h e p e r i c a r d is slightly shorter t h a n the n e p h r i d i u m , w h i c h is t r i a n g u l a r and
slightly c u r v e d . T h e p u l m o n a r y v e i n is rather p r o m i n e n t , but the side
veins are w e a k l y developed. T h e adrectal ureter is closed o v e r its entire length. T h e p e n i s is without a sheath a n d club-shaped. T h e epiphallus is as l o n g as a n d h a l f as b r o a d as the distal p a r t o f the penis, s u b c y l i n d r i c a l a n d slightly s w o l l e n t o w a r d s the t r a n s i t i o n to the flagellum. T h e flagellu m is short (ca. 1/3
the length o f the e p i p h a l l u s ) a n d t a p e r i n g t o w a r d s the distal e n d . T h e
v a g i n a is v e r y short. T h e p r o x i m a l p a r t o f the spermathecal duct is h a l f as b r o a d as the m e d i a n part, w h i c h has a spermathecal a p p e n d i x . T h e distal p a r t o f the duct, w h i c h bears the o v o i d spermatheca, is relatively n a r r o w . Distribution. — Ecology. —
Venezuela, Brazil, PArgentina.
Unknown.
Relationships. —
T h e phylogenetic relationships o f this g e n u s are discussed
o n page i 6 i f f . T h e g e n u s is characterized b y the shell shape, the s p i r a l series
BREURE, BULIMULINAE
of
h a i r s , the
sculpture
of
the
protoconch,
the
131
expanded
peristome,
the
structure o f the r a d u l a a n d the presence o f a spermathecal a p p e n d i x . Bibliography. —
T h e m a i n publications o n this g e n u s a r e : B r e u r e , 1978b;
Pilsbry, i897d. T a x a . — T h e f o l l o w i n g t a x a are i n c l u d e d i n this g e n u s : angosturensis, Bulimus, G r u n e r , 1841: 278, p l . 11 f i g . 3 (republicae Venezuela, província Orinoco). argentinus, Bulimulus (Rhinus), A n c e y , 1901: 92 ( [ A r g e n t i n a ] province d ' E n t r e r i o s , Gualeguaychu) [ S T I R S N ] . ciliatus, Bulimus, G o u l d , 1846b: 191 ( B r a z i l , O r g a n M o u n t a i n s ) . constrictus, Bulimus, P f e i f f e r , 1841: 43 ( A n g o s t u r a [Venezuela, C i u d a d B o l i v a r ] ) . heterogramma, Helix (Cochlogena), S. M o r i c a n d , 1836: 437, p i . 2 figs. 15-17 ( [ B r a z i l , Bahia] Caxoeira). heterotricha, Helix (Cochlogena), S. M o r i c a n d , 1836: 430, p i . 2 figs. 5-6 [no type locality g i v e n ] . hirtus, Bulimus, Beck, 1837: 51 [indication]. hyaloideus, Bulimus, P f e i f f e r , 1855c: 292 ( M e n d e z , A n d e s o f N e w Granada) [ L T B M N H 1975412]. koseritzi, Bulimus (Rhinus), Clessin, 1888: 168 ( B r a s i l i e n ) . longiseta, Helix (Bulimus), S. M o r i c a n d , 1846: 156, p i . 5 figs. 18-20 ( [ B r a z i l ] l a province de B a h i a ) . obeliscus, Bulimulus (Rhinus), H a a s , 1836: 150, figs. 15-16 ( B r a s i l i e n , Staat S t a . Catharina) [ H T S M F 10078]. ovulum, Bulimus, Reeve, 1849: p l . 46 f i g . 556 ( P h i l i p p i n e Islands [sic]) [ L T B M N H 1975416]. pubescens, Helix (Bulimus), S. M o r i c a n d , 1846: 157, p i . 5 figs. 21-23 ( [ B r a z i l ] les environs de B a h i a ) . rochai, Bulimulus (Rhinus), F . B a k e r , 1914: 636, p i . 23 figs. 19-20 ( [ B r a z i l ] Jacoco, 7 k m f r o m Ceará-Mirim ) [ H T A N S P 100058]. scobinata, Helix, W o o d , 1828 : p i . 8 f i g . 77 [publication not seen]. subtenuis, Bulimulus heterotrichus, P i l s b r y , i 8 9 7 d : 76, p i . 13 figs. 2, 25, p i . 15 f i g . 19 [ B r a z i l , teste C l e n c h & T u r n e r , 1962; L T A N S P 25658a]. suturalis, Bulimulus (Rhinus) rochai, F . B a k e r , 1914: 637, p i . 23 figs. 13-14 ( [ B r a z i l ] Mongúba, Ceará & Baturité R . R . , about 27 k m f r o m Ceará) [ H T A N S P 109322a]. taipuensis, Bulimulus (Rhinus) rochai, F . B a k e r , 1914: 636, p i . 23 f i g . 17 ( [ B r a z i l ] fossil beds on the C e n t r a l R . R . , 46 k m f r o m N a t a l ) [ H T A N S P 109321]. tateanus, Bulimus constrictus, Guppy, 1875b: 322 (Venezuelan G u i a n a ) . thomei, Bulimulus (Rhinus), W e y r a u c h , 1967b: 481, figs. 3-5 (Sureste de B r a s i l , estado R i o Grande do S u l , L i v r a m e n t o ) [ H T M R C N 1021a]. velutinohispida, Helix (Cochlogena), S. M o r i c a n d , 1836: 429, p l . 2 f i g . 6 [no type locality g i v e n ; S T M N H N ] .
Simpulopsis B e c k ,
1837
Simpulopsis Beck, 1837: 100. T y p e species b y subsequent designation Helix sulculosa Férussac. Simulopsis G r a y , 1847: 171 [emendation f o r Simpulopsis B e c k ] . Description. —
S h e l l elongate-ovate to globose;
narrowly
( A l b e r s , i860) :
perforat e
to
i m p e r f o r a t e ; thin . S u r f a c e s m o o th o r corrugate. P r o t o c o n c h w i t h fine s p i r a l lines that m o r e o r less cut the low, oblique riblets o r w r i n k l e s into granules. W h o r l s slightly to moderately c o n v e x , the last w h o r l p r o m i n e n t ; suture w e l l
132
ZOOLOGISCHE V E R H A N D E L I N G E N 168
F i g . 167. D i s t r i b u t i o n of
(1979)
Simpulopsis.
BREURE, BULIMULINAE impressed. A p e r t u r e (oblique)
(elongate-)ovate. P e r i s t o m e t h i n a n d simple.
T h e central teeth o f the r a d u l a are (1) to t r i a n g u l a r mesocones; mesocones;
or
(3)
m o n o c u s p i d , w i t h blunt, elongate
(2) m o n o c u s p i d , w i t h rather blunt,
tricuspid, with
t r i a n g u l a r ectocones.
133
elongate to
L a t e r a l teeth are (1)
lanceolate
spatula-shaped mesocones
and
t r i c u s p i d , w i t h acute, spatula- to
wedge-shaped mesocones, small t r i a n g u l a r endocones a n d t r i a n g u l a r to deltoid ectocones;
o r (2)
b i - to t r i c u s p i d , w i t h acute lanceolate mesocones,
t r i a n g u l a r endocones)
a n d t r i a n g u l a r to deltoid ectocones.
lateromarginal) teeth are (1) t r i c u s p i d , w i t h blunt, spatula-shaped acute, c u r v e d elongate endocones
(2)
(and
mesocones,
a n d acute, ovate to t r i a n g u l a r
w h i c h m a y be serrate i n the outermost teeth;
(small
Marginal
ectocones,
t r i c u s p i d , shifted,
with
elongate mesocones, c u r v e d elongate endocones a n d t r i a n g u l a r ectocones; (3) b i c u s p i d , w i t h elongate mesocones a n d t r i a n g u l a r , b i f i d ectocones. row formulas: ( x = 7-10,
C/i
+
L M x/3
y = 24-31), C / 3
or C / 3 + L M x / 2 (x =
to
deltoid.
=
L x/3
35-48), C / i +
M
y/3
+
L x/2-3
+
M
y/3
( x = 10, y = ca. 20-28)
24).
T h e p e r i c a r d is 1/2-4/5 triangular
+
(x
or
Half-
The
t
n
e
l e n g th o f the n e p h r i d i u m , w h i c h is b r o a d l y
main pulmonary
vein
a n d the
side veins
are
moderately to weakly developed. T h e adrectal ureter is closed o v e r its entire length. P e n i s without
a sheath
( i n some
species
w i t h a t u n i c a ) , m o r e o r less
swollen. T h e epiphallus is slender a n d s u b c y l i n d r i c a l , the flagellum is t a p e r i n g o r subcylindrical . T h e p r o x i m a l part o f the spermathecal duct is m o r e o r less s u b c y l i n d r i c a l a n d relatively thick, w i t h a spermathecal a p p e n d i x ; part o f the duct is n a r r o w ; the spermatheca is Distribution. —
Mexico,
Guatemala,
the distal
(elongate-)globose.
West
Indies, V e n e z u e l a ,
Surinam,
F r e n c h Guyana, Brazil, Paraguay, Argentina, P e r u , Ecuador, Colombia. Remarks.
—
The
d i v i s i o n i n subgenera
is m a i n l y based
o n the
shell
m o r p h o l o g y ; the structure o f the r a d u l a a n d the m o r p h o l o g y o f the genitalia are r e m a r k a b l y variable a n d do not parallel the subgeneric d i v i s i o n . Relationships. —
The
phylogenetic
relationships are discussed o n page
i 6 i f f . T h e genus is characterized b y the t h i n shell, the sculpture o f the p r o t o conch, the presence
o f a spermathecal
appendix
a n d the structure o f
the
radula. Bibliography. — &
T h e m a i n publications o n the genus are: A r a u j o ,
1975;
Araujo
1977;
H y l t o n Scott, 1967;
B r e u r e , 1977;
B r e u r e , 1975e, 1978b; B r e u r e &
P i l s b r y , 1899; V a n M o l ,
K e y to the subgenera o f
1971,
Ploeger,
1971.
Simpulopsis
a.
S h e l l globose; surface m o r e o r less corrugate
b.
S h e l l elongate-ovate; surface smooth .
.
Simpulopsis .
Simpulopsis
(Simpulopsis) (Eudioptus)
136
ZOOLOGISCHE VERHANDELINGEN 168 (1979)
callosus, Bulimus, P f e i f f e r , 1846: 128 [locality u n k n o w n ] . cinereus, Bulimus, Reeve, 1848 : p l . 56 f i g . 372 ( B o l i v i a ) . erosus, Bulinus, B r o d e r i p i n B r o d e r i p & Sowerby, 1832b : 106 (Peruviâ, H u a n t a j a y a near Iquiqui) [?Bostryx]. fasciata, Scutalus conospirus, Döring, 1879: 67 ( [ A r g e n t i n a ] S i e r r a de T u c u m a n ) [ ? Bostryx stelzneri]. felipponei, Bulimulus (Scutalus), Ihering, 1928: 95 (Republic of U r u g u a y , Canelones) [ ? ? Naesiotus]. flossdorfi, Bulimulus, H o l m b e r g , 1909a: 11 ( [ A r g e n t i n a ] Chaco, Território de F o r m o s a , prope N u e v a Pompeya) [?Bostryx]. fonsecanus, Bulimulus (Rhabdotus), H a a s , 1961: 20, figs, i i a - b ( S a n Salvador o r N i c a r a g u a , G u l f o f Fonseca) [ H T F M N H 106702]. fourmiersi, Bulimus, d O r b i g n y , 1836: 273, p l . 30 figs. 12-14 ( [ A r g e n t i n a ] l a province de Corrientes, non loin du R i o de Santa L u c i a , au lieu nommé P a s t o reito) [ INaesiotus]. fusca, Bulimulus heloicus, A n c e y , 1901a: 82 (aux environs de Gualeguaychu, province E n t r e r i o s , République A r g e n t i n e ) . guttatus, Bulinus, B r o d e r i p i n B r o d e r i p & Sowerby, 1832a: 31 (Peruviâ, Cobija o r P u e r t o de l a M a r ) [ ? Bostryx]. heloica, Helix, d O r b i g n y , 1835: 11 (província Chiquitensi, republica B o l i v i a n a ) . joergenseni, Bulimulus, H o l m b e r g , 1912b: 150, figs. 7-8 ( A r g e n t i n a , M i s i o n e s , Colónia Bonpland). metamorphus, Bulimulus, P i l s b r y , 1896b : 157, p l . 1 figs. 6-7 ( C h i l i ) [ L T A N S P 69455a] [} Bostryx]. munsterii, Bulimus, d O r b i g n y , 1836: 278, p l . 34 figs. 4-7 ( [ B o l i v i a ] côte de Petaca). nivalis, Helix, d O r b i g n y , 1835 [no type locality given ; P o t o s i , B o l i v i a (cf. d O r b i g n y , 1836: 260) ] [ ?Scutalus]. obliquistriatus, Drymaeus, D a Costa, 1901: 238, p i . 24 f i g . 2 ( P e r u , S a n P a b l o ) [ H T B M N H 1907.11.21.41] [Bostryx]. olorinus, Bulimus, Duelos, 1833 : p l . 24 [ = albus S o w e r b y ] . pliculosa, Bulimulus turritella, A n c e y , 1901a: 92 ( [ B r a z i l ] M a t t o - G r o s s o ) . robertsi, Thaumastus, P i l s b r y , 1932: 390, p i . 27 figs. 3, 6 ( P e r u , Dept. S a n M a r t i n , R i o Jelashte, 4500 ft.) [ H T A N S P 150920a]. saltensis, Bulimulus, H o l m b e r g , 1909b: 91 ( [ A r g e n t i n a ] Salta, ad r i p a m R i o de los Horcones et R i o de las P i e d r a s ) . sowerbyi, Bulimus, P f e i f f e r , 1847a: 114 (Andes o f C o l o m b i a ) . spixii, Bulimus, P o t i e z & M i c h a u d , 1835 : p l . 15 figs. 13-14 (république A r g e n t i n e et Bolivienne) [ ? Bostryx]. stenogyroides, Bulimulus, Guppy, 1868: 431 ( D o m i n i c a ) . stilbe, Bulimulus, P i l s b r y , 1901: 145, p l . 25 f i g . 18 ( B r a z i l , State of S a o P a u l o ) [ H T A N S P 73434]. striatellus, Buliminus, Beck, 1837: 70 [indication]. striatus, Bulinus, K i n g i n Sowerby, 1833a: f i g . 56 (Santos, P e r u ) [?Bostryx]. tenebrosa, Thaumastus onca, P i l s b r y , 1926a: 7, p l . 2 figs. 9-10 ( B o l i v i a , Dept. Cochabamba) [ H T A N S P 138107] [?Plekocheilus]. turritella, Helix, d O r b i g n y , 1835: 13 (província Chiquitensi, republica B o l i v i a n a ) . turritellatus, Bulimulus, Beck, 1837: 67 ( N e w name f o r Helix turritella d O r b i g n y , 1835, not Férussac, 1821). ulloae, Bulimus, P h i l i p p i , 1869: 34 ( P e r u v i a i n pampa inter M a y o c et H u a n t a ) . umbilicatus, Thaumastus, M i l l e r , 1879: 122, p i . 12 f i g . 5, p i . 13 f i g . 1 ( [ E c u a d o r ] P r o v . L o j a , R i o Catamayo, 2000-3000 ft.) [?Scutalus], ventricosus, Mesembrinus, P a r a v i c i n i , 1894: 6 ( [ A r g e n t i n a ] província d i Salta, S. Rosa) [ ^.Drymaeus/ ? Bostryx]. woodwardi, Bulimus, P f e i f f e r , 1857b: 332 (Andes o f P e r u ) [ L T B M N H 1975334] [ ? Bostryx]. :
1 2
138
ZOOLOGISCHE VERHANDELINGEN 168 (1979)
Thaumastus
limneiformis
tenuis
W a r r e n (1926:
7)
a n d T.
l.
procerior
R u s s e l l (1926: 220) are o n l y m e n t i o n e d here. It is not clear whether they are correctly r e f e r r e d to the B u l i m u l i d a e .
VII.
PHYLOGENY AND ZOOGEOGRAPHY "Ideas deserving to be called O r i g i n a l ' i n the strictest sense probably do not e x i s t " Ghiselin, 1974: 9. PHYLOGENY
I n this section the methods outlined b y H e n n i g (1966) are used. C o m m e n t s o n these
methods
may
be
found in Ashlock
(1971,
1973,
^97A),
Colless
(1967, 1969a, 1969b, 1970), C r a c r a f t ( 1 9 7 4 ) , D a r l i n g t o n ( 1 9 7 0 ) , E n g e l m a n n & W i l e y ( 1 9 7 7 ) , F a r r i s et al. ( 1 9 7 0 ) , F a r r i s ( 1 9 7 6 ) , H e n n i g (1971, 1975), M a y r Peters
(1970,
(1974), Nelson 1972), Peters
(1971a, 1971b, 1972a, 1972b, 1973b, &
Gutmann
(1971,
1973), P l a t n i c k
1974, 1974),
(1976a,
1977), R e m a n e ( 1 9 7 1 ) , R o s e n ( 1 9 7 4 b ) , Schlee (1969, 1971), S o k a l ( 1 9 7 5 ) , V a n der S t e e n & B o o n t j e ( 1 9 7 3 ) . T h i s is the first time that H e n n i g i a n principle s are a p p l i e d to B u l i m u l i d a e . I have t r i e d to j u s t i f y the t r a n s i t i o n series (key-notion s o f these p r i n c i p l e s ) , w h i c h I recognize, as well as possible. I n most instances I have b e e n f o r c e d to make decisions o n account o f the f r e q u e n c y o f the v a r i o u s character states i n the B u l i m u l i n a e . I n some other cases, e.g. the presence o r absence o f a penis sheath, I h a d to take a n a r b i t r a r y decision. I hope that f u r t h e r research w i l l i m p r o v e m y transition series a n d their argumentation . The
following
character
transition series
have
been
recognized
(sum-
marized i n Table 4) : Radula. —
(1)
T h e central teeth o f the r a d u l a are n o r m a l l y t r i c u s p i d , not
o n l y i n B u l i m u l i d a e but i n m a n y other pulmonate families as well (cf. S o l e m , 1974:
170). T h e presence o f m o n o c u s p i d central teeth is generall y
hypothe-
sized to be a n a p o m o r p h o u s character, w h i c h presence i n d i f f e r e n t
genera
is best u n d e r s t o od as the result o f convergent evolution ( B r e u r e &
Gitten-
berger, i n p r e p a r a t i o n ) ; (3)
(2)
the same applies to m o n o c u s p i d lateral teeth.
T h e length axis o f the mesocones
of
(lateral a n d )
m a r g i n a l teeth
is
n o r m a l l y parallel to the length a x i s o f the basal plate. I n some g r o u p s , h o w ever, the length a x i s o f the mesocone ectocones)
( a n d n o r m a l l y also o f the e n d o - a n d
is shifted, i.e., t u r n e d ca. 20-30 degrees w i t h respect to the length
a x i s o f the basal plate. T h i s character state, w h i c h I consider a p o m o r p h o u s , is not correlated to the ecology
o f the species
d w e l l i n g a n d i n arboreal species. is most p r o b a b l y the
result o f
Its
presence
convergent
(it occurs both i n g r o u n d in different
evolution.
(4)
genus The
groups
transverse
present
(Laterals-) marginal teeth
Transverse rows
Supporting denticles
3.
4.
5.
Pulmonary veins
Glandular folds of spermoviduct
20.
21.
22.
Protoconch
Spermathecal appendix
19.
SHELL
Retractor muscle
Spermathecal duct
18.
Curved f o l d i n flagellum
Epiphallus-penis
17.
15/16.
Subepithelia l tissue i n d i s t a l penis
Penis lumen
12/13.
14.
Penis lumen
"Pseudo-sheath"
Glandular c e l l types i n penis epithelium
8.
9.
10/11.
Penis sheath
7.
GENITALIA
6.
PALLIAL ORGANS
straight
Lateral teeth
spermoviduct
sculptured
p a r a l l e l to length axis of spermoviduct
absent
as long as
d i s t a l l y inserted
single
gradual change
muscular fiber s
•divided', with pouches
undivided
one
absent
absent
weakly to moderately developed
length axis p a r a l l e l
bi/tricuspid
Central teeth
1. tricuspid
(1)
Plesiomorphous
2.
RADULA
Character
4
smooth
perpendicular to length axis of spermoviduct
present
reduced i n length
subdistally inserted
double
intrudes i n d i s t a l penis
glandular c e l l s
p a r a l l e l tubes
•simply' constricted i n median part
two or more
present
present
strongly developed, anter i o r l y ramified
absent
V- or W-shaped
(3)
do., i n d i s t a l + p r o x i m a l p e n i s
c i r c u l a r glana
'complex' c o n s t r i c t e d i n median p a r t
pericard t r a n s v e r s a l ly disposed
Apomorphous
length axis 'shifted'
monocuspid
monocuspid
(2)
S u m m a r y o f character t r a n s i t i o n series i n B u l i m u l i n a e
TABLE
>
r
1—1
w w c
ZOOLOGISCHE VERHANDELINGEN 168 (1979)
140
rows o f the r a d u l a are usually straight o r slightly bent i n the outermost teeth. I n some genera the transverse r o w s are V - o r W - s h a p e d a n d this is hypothesized to be a n a p o m o r p h o u s character. (5) I n most g e n e r a the central part o f the r a d u l a is p r o v i d e d w i t h a n i n t e r r o w i n t e r l o c k i n g system b y the presence o f ' s u p p o r t i n g denticles\ T h e absence o f this system is hypothesized to be apomorphous. Palliai
organs. —
n o r m a l l y weakly
(6)
The
m a i n p u l m o n a r y veins
to moderately well developed;
a n d side veins
are
veins parallel to the m a i n
p u l m o n a r y v e i n are absent. I n some genera, however, the veins are strongly developed a n d r a m i f i e d at the anterior e n d , where one o r two veins parallel to the m a i n p u l m o n a r y v e i n m a y be present ( i n part o f the genera, m o r e o v e r , the p e r i c a r d is transversall y d i s p o s e d ) ; this situation is considere d a p o m o r phous. Genitalia. —
(7)
The
f u n c t i o n o f the p r o x i m a l penis sheath, w h i c h is
present i n most genera, is not clear. It is not possible to decide a
priori
between the two possibilities: either the presence o r the absence o f a sheath is a n a p o m o r p h o u s character state. I have chosen f o r the presence o f the sheath to be a p o m o r p h o u s . [It m a y be noted that, i f the absence o f the sheath is hypothesized to be a p o m o r p h o u s , the resulting cladograms do not alter substantially]. (8) I n transverse sections o f the penis the outer layer is m a d e u p b y m u s c u l a r fibres. N o r m a l l y this layer is united, but i n one genus it is d i v i d e d into a n i n n e r a n d outer layer, w h i c h are unconnected o v e r the greater length o f the penis. T h e outer layer is here n a m e d 'pseudo-sheath' a n d its presence is considered a p o m o r p h o u s . (9) T h e epitheliu m b o r d e r i n g the penis l u m e n has a glandular function. U s u a l l y the epithelium is m a d e u p b y one cell type a n d the presence o f two o r m o r e g l a n d u l a r cell types is hypothesized to be a p o m o r p h o u s . (10) T h e shape o f the penis l u m e n i n l o n g i t u d i n al section leads to two transitio n series that, i n m y o p i n i o n , are independent. I n the first series the presence cylindrical
of
qua width
a 'simple', u n d i v i d e d l u m e n that is m o r e o r less is considered p r i m i t i v e . W h e n the l u m e n is ' c o n -
stricted* i n its m e d i a n part, w h i c h is hypothesized to be a p o m o r p h o u s , this m a y be either 'simple' (10) o r ' c o m p l e x' (11).
(12)
I n the second series the
l u m e n is ' d i v i d e d ' either into a b r o a d a n d n a r r o w p a r t o r into a 'central' l u m e n a n d 'side l u m i n a ' . T h e presence o f pouches is t h e n hypothesized to be plesiomorphous a n d parallel tubes are a p o m o r p h o u s ; (13) a specialized f o r m of these parallel tubes is the presence o f a c i r c u l a r g l a n d , where the tubes are reduced i n length a n d (nearly totally)
united. (14)
T h e subepithelial
tissue i n the distal part o f the penis m a y be made u p o f m u s c u l a r fibers, w h i c h is considered the p l e s i o m o r p h o us situation, o r b y large, r o u n d e d cells that p r o b a b l y have a g l a n d u l a r function; the presence o f these ( g l a n d u l a r ) cells is hypothesized to be a p o m o r p h o u s . (15) T h e distal part o f the penis g r a d u a l l y
141
BREURE, BULIMULINAE
changes into the epiphallus o r, hypothesized to be a p o m o r p h o u s , the epiphallus intrudes the distal part o f the penis;
(16)
w h e n the epiphallus not o n l y
intrudes into the distal part but also into the p r o x i m a l p a r t o f the penis this is hypothesize d to be relatively a p o m o r p h o u s . (17)
T h e flagellum has i n t e r n -
ally a c u r v e d , l o n g i t u d i n a l f o l d that has a f u n c t i o n i n the f o r m a t i o n o f the spermatophore ( B r e u r e & E s k e n s , 1978). I n some gener a this f o l d is doublec u r v e d a n d this situation is hypothesized to be a p o m o r p h o u s . (18) T h e penis retractor muscle is n o r m a l l y attached to the distal e n d o f the flagellum. I n some g r o u p s , however, the muscle is subdistally inserted at the flagellum o r at the t r a n s i t i on between epiphallus a n d flagelum; a p o m o r p h o u s character state. (19)
this is c o n s i d e r e d a n
T h e spermathecal duct is usually about
as l o n g as the spermoviduct . T h e r e d u c t i o n i n l e n g t h o f this duct is hypothe sized to be a p o m o r p h o u s a n d the presence o f this character state i n d i f f e r e n t g e n u s g r o u p s m a y be the result o f convergen t
evolution. (20)
The
sper-
mathecal duct is n o r m a l l y s u b c y l i n d r i c a l o r tapering. I n some genera, h o w ever, the p r o x i m a l part o f the duct is relatively thick a n d has a n a p p e n d i x ; the distal p a r t o f the duct is n a r r o w a n d originates below the distal e n d o f the p r o x i m a l part. T h e presence o f a spermathecal a p p e n d i x is hypothesized to be a p o m o r p h o u s . (21)
T h e g l a n d u l ar folds o f the s p e r m o v i d u c t are n o r m a l l y
p e r p e n d i c u l a r to the l e n g t h a x i s (Peters
&
G u t m a n n , 1971)
of
the duct. F o r reasons o f
parsimony
this is considere d a n a p o m o r p h o u s character
state. W h e n the g l a n d u l a r folds are a r r a n g e d parallel to the lengt h a x i s o f the s p e r m o v i d u c t this is considered p l e s i o m o r p h o u s . (22) T h e presence o f p r o t o conch sculpture is hypothesized to be p l e s i o m o r p h o u s ; a s m o o t h p r o t o c o n c h is considered a n a p o m o r p h o u s character state. T h e cladograms presented below are based o n the a b o v e - m e n t i o n e d t r a n sition series. A s these series o n l y c o n c e r n anatomical characters the genera f o r w h i c h n o anatomical data are available are left out o f these cladograms. T h e cladograms represent the least rejected hypotheses
of
phylogenetic
relationships a m o n g the ( s u b ) g e n e r a o f each g r o u p . W i t h the present data at h a n d it was impossible to fit the cladograms o f the d i f f e r e n t genus g r o u p s into one c l a d o g r a m f o r the whole s u b f a m i l y . T h e r e are, m o r e o v e r , groups of
w h i c h the m o n o p h y l y is not c o r r o b o r a t e d b y
some
synapomorphous
characters; these g r o u p s are treated as such o n account o f other characters, e.g.
the sculpture o f the p r o t o c o n c h ( a t r a n s i t i o n series o f the
different
sculptures is too hypothetical), o r out o f tradition . ZOOGEOGRAPHY T h e available zoogeographical (or, m o r e correctly perhaps : biogeographical ) theories have
recently been reviewe d b y
Cracraft
1976). T h e s e theories m a y be s u m m a r i z e d as follows:
(1975b;
see also
Ball,
BREURE, BULIMULINAE
räume" o f D e L a t t i n ; Müller
(1974:
143
f i g . 4 9) lists f o r the N e o t r o p i c s the
H y l a e a ( r a i n f o r e s t ) , savannah, steppe, desert a n d oreal b i o m e s ] ; the areas of these biomes are i n f l u e n c e d b y climate-oscillations a n d
vegetations-fluc-
tuations a n d so are the areas o f the species w h i c h f o r m part o f these biomes. U n d e r u n f a v o u r a b l e e n v i r o n m e n t a l conditions (e.g., a n a r i d p e r i o d f o r r a i n forest species) species m a y s u r v i v e i n a restricted area ( = dispersal centre ) ; x
i n these areas speciation ) m a y have t a k en place so that, e.g., populations o f 2
the same species w h i c h s u r v i v e d i n d i f f e r e n t areas p r o v e to d i f f e r at the subspecies level w h e n they m a k e contact u n d e r m o r e favourable conditions afterwards. 4.
H i s t o r i c a l biogeography, w i t h w h i c h I designate the theory o f
(1958, 1964, 1976;
also C r o i z a t , N e l s o n & R o s e n , 1974;
Croizat
cf. N e l s o n , 1973a).
T h e m a i n p r i n c i p l es o f this theory are (see C r o i z a t et al., 1974): a )
the
distribution of a g r o u p can be represented b y one o r m o r e tracks connecting the ranges o f all m e m b e r s o f that g r o u p ; b ) m a n y o v e r l a p p i n g i n d i v i d u a l tracks because
form of
a generalized track w h i c h estimates changing
geography,
has
become
a n ancestral biota
s u b d i v i d e d into
that,
descendant
biotas; c) overlap ( s y m p a t r y ) o f generalized tracks, o r a n y o f the components of
different
generalized tracks, reflects
geographical overlap o f
different
biotas due to dispersal. T h e three last m e n t i o n e d theories m a y be called analytical ( s e n s u B a l l ) . A
descriptive biogeographica l p u b l i c a t i o n o n the N e o t r o p i c s is C a b r e r a
&
W i l l i n k (1973). T h e evolutionary biogeographical theory is here repudiated because o f its non-analytical nature. T h e phylogenetic biogeographical theory has its merits because o f the p r i o r phylogenetic analysis c a r r i e d out o f the g r o u p i n question. A s B r u n d i n (1966) says, knowledge o f phylogenetic relationships is a prerequisite f o r the d i s cussion o n d i s t r i b u t i o n patterns, as the value o f biogeographical conclusions depends o n the value o f phylogenetic arguments. H o w d e n (1972: 130)
has
pointed out that " H e n n i g ' s methodology m e r e l y shows relationships, it does not e x p l a i n the dispersal m e c h a n i s m s " . T h e r e are two other points to c o m m e n t o n . T h e theory involves the concept o f centre o f o r i g i n i n the f o r m o f the 1) Müller (1973b: 3) states: " I do not assume at the outset that dispersal centres represent centres where faunas and f l o r a were preserved d u r i n g regressive phases", while the same author (1974: 157) defines : " D i s p e r s a l centres are areas i n w h i c h animals and plants survived unfavourable environmental conditions". T h e latter statement was expressed i n his earlier papers as w e l l and is used, therefore, to summarize the theory. 2) A c c o r d i n g to Müller (1973a: 232) there are two types of differentiation (speciation) : a) allopatric differentiation w i t h i n a continuous distribution area on account o f different selection forces ; b) differentiation by geographical isolation.
ZOOLOGISCHE VERHANDELINGEN 168 (1979)
144
' p r o g r e s s i o n rule': ancestral f o r m s r e m a i n at, o r near, the point o f o r i g i n a n d the d e r i v e d f o r m s migrate. A contrariwise o p i n i o n has bee n expressed,
e.g.,
b y D a r l i n g t o n (1963) a n d B a l l (1976: 420) correctly r e m a r k e d that " i t is d i f f i c u l t to decide a p r i o r i between these two p o s s i b i l i t i e s \ S e c o n d l y , reti ,
culate evolution (although possibly m o r e c o m m o n i n plants t h a n i n a n i m a l s ) m a y cause some problems, as S n e a th
(1975) has s h o w n , w h i c h cannot be
solved w i t h the present phylogenetic theory. T h e two theories that involv e the concept o f vicarianc e have been c o m p a r e d so f a r o n l y b y C r o i z a t (1976:
519
although his theory is 'analytical'
ff.).
It cannot be denied that
Müller,
( s e n s u B a l l ) , o f t e n allows h i m s e l f ' n a r -
rative' explanations (several examples i n C r o i z a t , o.e.). Y e t , the
differences
between the two theories are perhaps less than C r o i z a t supposes. B o t h accept vicariance as a p r i n c i p a l factor u n d e r l y i n g d i s t r i b u t i o n patterns ( M ü l l e r o n l y implicitly, C r o i z a t e x p l i c i t l y ) . F u r t h e r m o r e , as the dispersal centres o f M ü l l e r are a p r i o r i correlated to certain vegetational
f o r m a t i o ns these centres r e -
present i n some w a y the biotas o f C r o i z a t ) . A poin t also to be t a k e n into 3
account is the c o m m e n t b y Müller zoogeography
are nevertheless
(1974: 6) that "ecological a n d historical
not o p p o s e d to each other. A n y
argument
about w h i c h o f the two is m o r e i m p o r t a nt is, as de L a t t i n (1967) said, quite beside the p o i n t " . T h e m a i n difference s
between the two theories are that
Müller assigns relative importance to ecological factors w h i c h o c c u r r e d i n the P l i o - P l e i s t o c e n e ) , whereas 4
o c c u r r e d as e a r ly as the
C r o i z a t maintains that geological events w h i c h
T e r t i a r y are responsible
for "differential
form
m a k i n g " ) . It is not possible to decide a p r i o r i w h i c h o f these viewpoints is 5
correct a n d it is likely that b o t h are correct (e.g., ecological factors o f the Pleistocene to e x p l a i n the d i s t r i b u t i o n o f certain subspecies a n d
geological
events o f the T e r t i a r y e x p l a i n i n g the d i s t r i b u t i o n pattern s h o w n b y a certain genus).
T h e r e are, however,
some
severe
drawbacks
to Müller's
w h i c h make it d i f f i c u l t to accept it as a general theory, v i z . (1)
theory,
the t h e o r y
o n l y explains intracontinental d i s t r i b u t i o n ; (2) the t h e o ry h e a v i ly
depends
[ i n m y o p i n i o n to a f a r greater extent than does the theory o f C r o i z a t ] o n the t a x o n o m ie quality o f the g r o u p i n question, i.e., the availability o f
a
stable t a x o n o m y at the species level; f o r this reason M ü l l e r was o n l y able to analyse the d i s t r i b u t i o n o f vertebrate between
(sub)species;
( 3 ) the " r e l a t i o n s h i p s "
the d i f f e r e n t centres d e p e n d o n p o l y t y p i c a n d p o l y c e n t r i c
species
w h i c h are " c o r r e l a t ed w i t h the particular vegetational f o r m a t i o n s o r climatic 3) C r o i z a t (1958, I : 461) also speaks of "center of d i s p e r s a l " ; these centres are not correlated to vegetational formations but (more or less vaguely) to geological history. 4) Müller seems w e l l aware o f the importance of geological factors (1972: 106; 1973b: 153), but hardly mentions any. 5) A c c o r d i n g to C r o i z a t a succession of comparatively r a p i d climatic cycles may cause immediate extinction, but does not affect f o r m m a k i n g (e.g., C r o i z a t 1958, I : 29).
BREURE, BULIMULINAE
145
types to w h i c h the elements are a d a p t e d " (Müller , 1973b: 166). T h i s means that the d i s t r i b u t i o n o f t a x a that cannot be correlated to a certain vegetational f o r m a t i o n o r o f w h i c h the ecology is u n k n o w n , cannot be e x p l a i n e d . It also means that this theory is a s e l f - f u l f i l l i n g p r o p h e c y : i f species are correlated w i t h certain ecological factors t h e n the centres i n w h i c h these species o c c u r w i l l be related a c c o r d i n g to the same ecological factors (e.g., a r b o r e a l, o r e a l a n d n o n - f o r e s t centres; Müller, 1973b: 173, figs. 9 3 - 9 5 ) . F u r t h e r m o r e , (a) the correlatio n that M ü l l e r lies between d i s t r i b u t i o n a n d phylogenetic relation ships (o.e.: f i g . 86) is incorrect, as his "phylogenetic r e l a t i o n s h i p s " are not based o n a n analysis u s i n g the methods o f H e n n i g , but p r o b a b l y o n l y indicate m o r p h o l o g i c a l resemblance;
(b)
M u l l e r ' s theory is m e r e l y zoogeographical
(also " z o o g e o g r a p h i c a l " sensu C r o i z a t ) , rather t h a n biogeographical, i n that it does not e x p l a i n plant distributions; (c) to say that "the degree o f relationships between centres does not o n l y d e p e n d o n the ecological distinctiveness of a g r o u p . It equally depends o n the vagility, the course o f e v o l u t i on a n d the geographical isolation o f the f a u n a l elements", rationa l as it m a y seem, it is as 'narrative as the land-bridges o f most e v o l u t i o n a ry biogeographers. 7
F r o m the above d i s c u s s i on it m a y be c o n c l u d ed that I accept the v i c a r i a n ce theory advocated b y C r o i z a t . T h i s does not m e a n that I e n t i r e l y repudiate a l l other theories as such. T h e y p r o b a b l y each c o n t a i n several elements that are acceptable a n d applicable i n certain cases. Nevertheless some additiona l r e m a r k s need to be made. A s B a l l (1976: 421) has p o i n t e d out, C r o i z a t d i d not r e g a r d the phylogenetic relationships of the t a x a ; o f course he c o u l d h a r d l y d o so because he c o m p i l e d his data f r o m the w o r k s o f e v o l u t i o n a ry biologists. O n c e phylogenetic p r i n ciples are a p p l i e d " w e f i n d that the m e t h o d [of C r o i z a t ] is little d i f f e r e n t f r o m the multiple sister-grou p rule o f
H e n n i g " ( B a l l , I.e.).
Phylogenetic
principles have been a p p l i e d to the vicarianc e theory b y R o s e n (1974a, 1976), c o n v i n c i n g l y s h o w i n g the strength o f
the method. C r o i z a t , however,
rejects the phylogenetic principle s a n d writes
(1976: 8 1 5 ) :
still
"los caracteres
"diagnósticos" de u n t a x o n siempre f i g u r a n u n a c o m b i n a t i o n ; razón p o r l a cual n o h a y "espécie m a d r e " que se p a r t a e n dos "especies-hijas" "monofiléticas";
sino, al contrario , los caracteres
necesariamente
de las "espécies" que f o r m a n
parte dei g r u p o se c o m b i n a n e n sus descendientes e n medidas que escapan a toda definición p r e v i a " 6 ) (italics o f C r o i z a t ) . F i n a l l y , B a l l (o.e.) has g i v e n a n adequate d i s c u s s i o n o f the inductive v e r s u s deductive a p p r o a c h to biogeographica l hypotheses. 6) " T h e "diagnostic" characters o f a t a x o n always f o r m a combination ; and this is w h y there is not a "mother species" splitting into t w o "daughter species" i n a monophyletic w a y ; but, on the contrary, the characters o f the "species" f o r m i n g part o f the group, combine i n the descendants i n a w a y that cannot be defined a p r i o r i " . 10
146
ZOOLOGISCHE VERHANDELINGEN 168 (1979) BULIMULINAE
T h e p h y l o g e n y a n d zoogeography o f the B u l i m u l i n a e w i l l n o w be discussed, especially o f the genus g r o u p s . T h r e e m a i n d i s t r i b u t i o n patterns pattern (Plectostylus-growp) p r i n c i p a l l y A n d e a n (Auristic pattern (Simpulopsis-,
m a y be distinguished :
(a)
a n austral
; ( b ) a strictly S o u t h A m e r i c a n pattern that is a n d Scutalus-groups)
Bulimulus-
and
; (c) a N e o t r o p i c a l / N e a r c -
Cochlorina-groups).
F i g . 168a. Hypothesis o f phylogenetic relationship i n the Auris-group. V i c a r i a n t distribution indicated by hatched circles.
BREURE, BULIMULINAE Fig.
147
168 is a s u m m a r y o f alternate hypotheses o f phylogenetic r e l a t i o n -
ships i n the Auris-group.
T h e m o n o p h y l y o f the g r o u p is c o r r o b o r a t e d b y one
character ( 6 ) , but there are no s y n a p o m o r p h i e s to corroborate the m o n o p h y l y of
Thaumastus
a n d T. the
as a genus. Thaumastus
(Thaumastiella)
same
applies
(Scholvienia),
T.
(Paeniscutalus)
are separated o n account o f the shell m o r p h o l o g y ;
to Plekocheilus
(Eurytus)
a n d P.
(Eudolichotis).
hypothesis s h o w n i n fig . 168b is rejected because o f character ( 2 1 ) ;
F i g . 168b. Hypothesis o f phylogenetic relationship i n the Auris-group.
The more-
148
ZOOLOGISCHE V E R H A N D E L I N G E N
l68 (1979)
a v N n n w n n a 'asnaxa
ISO
Z O O L O G I S C H E V E R H A N D E L I N G E N l68
(l979)
over, the radula structure is the same in Auris and Thaumastus, differing from that in Plekocheilus
(Breure, 1978b). The vicariant distribution of the
three genera (fig. 169) is not in contradiction with the hypothesis of fig. 168b, nor with that of fig. 168a. Auris, which is probably the relatively oldest
Fig. 171. Distribution of the Bulimulus-group.
BREURE, BULIMULINAE
151
g r o u p , is restricted to the Coastal B r a z i l i a n S h i e l d [see H a r r i n g t o n ( 1 9 6 2 ) for
palaeogeographical
Brazilian
data].
Thaumastus
S h i e l d into the A n d e s ;
s.str.
ranges
the other subgenera
from
of
the
Coastal
Thaumastus
are
restricted to parts o f the A n d e s a n d their existence is p r o b a b l y correlated w i t h A n d e a n orogenesis. Plekocheilus tively
apomorphous,
(Eudolichotis)
are
s.str. a n d P.
restricted
to
the
(Aeropictus),
Andes;
P.
w h i c h are r e l a (Eurytus)
are m a i n l y f o u n d o n the G u i a n a S h i e l d [P.
and
(Eurytus)
P.
seems
to be intermediate: one species g r o u p i n the A m a z o n b a s i n a n d o n the eastern escarpments o f the A n d e s , another species g r o u p at h i g h e r altitudes i n the A n d e s ] . T h i s possibly indicates that the d i r e c t i o n o f dispersa l ( s e n s u P l a t n i c k , 1976b) is f r o m the G u i a n a S h i e l d to the A n d e s . T h e o c c u r r e n c e o f
subspecies
of
is not i n
Plekocheilus
(P.)
fulminans
(Nyst)
c o n t r a d i c t i o n to this hypothesis. Fig.
character
(9).
Fig.
T h e m o n o p h y l y o f the g r o u p is c o r r o b o r a t e d b y one 170b
p a r a l l e l i s m i n Naesiotus
fig.
[ C f . C r o i z a t , 1958, I: 298; C r o i z a t , 1976].
170 s u m m a r i z e s the alternative hypotheses o f phylogenetic relationship
i n the Bulimulus-growp.
partly)
o n the R o r a i m a - m a s s i f
a n d Bostryx
shows
a hypothesis
a n d Bulimulus
(partly:
that
is rejected
because
(character 12 a n d characters
characters
1 2 / 1 3 ) . T h e hypothesis
170C is also rejected because o f p a r t i a l convergence.
shown i n
T h e least
hypothesis is s h o w n i n fig. 170a. T h e m o n o p h y l y o f Rabdotus
and
of
15/16
rejected Naesiotus
c o u l d not be c o r r o b o r a t e d w i t h the data at h a n d . It is n o w hypothesize d that b o t h Bulimulus
a n d Naesiotus
are o f p r e - A n d e a n o r i g i n . I f one p r e f e r s a n
i n d i c a t i o n o f the geological p e r i o d ) , the L o w e r T e r t i a r y c o u l d be suggested. 7
F o r Naesiotus
this is c o r r o b o r a t e d b y its d i s t r i b u t i o n o n the C o a s t a l B r a z i l i a n
S h i e l d (see f i g . 171);
the A n d e a n d i s t r i b u t i o n o f the g e n u s seems to be i n
contradictio n to s u c h a hypothesis, but m a y be the result o f the requirements of
the
species:
all live i n the lowest
valleys a n d are correlated to a r i d vegetational t r i b u t i o n o f Bulimulus
( i n t e r a n d e a n)
f o r m a t i o n s . T h e disjunct d i s -
m a y be the result o f the u p h e a v a l o f the A n d e s a n d
the ecological r e q u i r e m e n t s o f the species c o n d i t i o n s ) . Bostryx,
part o f
ecological
( v i z. relatively h i g h
temperatur e
too, is p o s s i b l y o f p r e - A n d e a n o r i g i n (it is represented
b y some species g r o u p s i n the coastal areas o f P e r u a n d C h i l e [unless this is e x p l a i n e d as secondary d i s p e r s a l ] ) , but e v o l v ed m o r e i n accordance w i t h the orogenesis
of
the A n d e s
t r i b u t i o n o f Rabdotus
t h a n Bulimulus
a n d Naesiotus.
F i n a l l y , the
dis-
is not easily e x p l a i n e d . T h e v i c a r i a n t d i s t r i b u t i o n w i t h
the other genera corroborates the p h y l o g e n y s h o w n i n fig . 170a, but I d o not
7) Geologica l periods and times are only mentioned here to give an approximate idea on the absolute age o f a genus o r a subfamily. M y data are based o n H a r r i n g t o n (1962) and C r a c r a f t (1075a) ; further research w i l l undoubtedly alter these data to a certain degree.
ZOOLOGISCHE VERHANDELINGEN 168 (1979)
152
k n o w o f a geophysical event that m a y account f o r this d i s t r i b u t i o n pattern, n o r f o r the disjunct d i s t r i b u t i o n o f the genus i t s e l f . ) C r o i z a t (1976: 541)
has
8
suggested that a vicariance between B a j a C a l i f o r n i a a n d S o u t h A m e r i c a (i.e. P e r u ) m a y be e x p l a i n e d b y " h o r s t i a n d i s t r i b u t i o n " (see also C r o i z a t , I: 762-763, 797, a n d R o s e n , 1976:
1958,
figs. 18-19).
F i g . 172 is a s u m m a r y o f alternate hypotheses o f phylogenetic relationships i n the Scutalus-group.
I f relative importanc e is assigned to character ( 9 ) the
hypothesis as s h o w n i n fig. 172b emerges. I n this hypothesis character m a y o n l y be e x p l a i n e d b y convergent evolution between Scutalus S.
(Vermiculatus)
a n d S.
(Kuschelenia).
(14)
(Suniellus),
T h e least rejected hypothesis
is
F i g . 172. Hypotheses of phylogenetic relationship i n Scutalus. 8) T h e relationships of Berendtia and Spartocentrum w i t h Rabdotus have not been considered, a w a i t i ng the revision of Christensen (in press).
BREURE, BULIMULINAE
F i g . 173. D i s t r i b u t i o n of
Scutalus.
153
ZOOLOGISCHE VERHANDELINGEN 168 (1979)
154
presented i n fig. 172a.
It implies that character (14)
is relatively i m p o r t a n t
a n d that the o c c u r r e n c e o f character ( 9 ) is a case o f convergen t
evolution.
T h i s hypothesis, however, neatly shows the v i c a r i a n c e between Scutalus
s.str.
a n d the other subgenera (see f i g. 173).
priori
It is not possible to decide a
between the two possibilities that (1) Scutalus
s.str. is the oldest g r o u p , w h i c h
also l i v e d i n p r e - A n d e a n biota a n d that the three other subgenera arose w i t h the upheaval o f the A n d e s ; Scutalus
o r (2)
the g r o u p has a n A n d e a n o r i g i n a n d
s.str. arose secondary i n the coastal r e g i o n o f P e r u . T h e o c c u r r e n c e
o f two a p o m o r p h o u s character states i n Scutalus
s.str., v i z . the presence o f
two g l a n d u l ar cell types i n the penis epitheliu m a n d the m o n o c u s p i d teeth i n the central part o f the r a d u l a [correlated to the ecology:
species l i v i n g o n
r o c k - f a c e s ] , suggests that the second possibility is m o r e plausible. I n fig. 174 the alternative hypotheses o f phylogenetic relationships i n the Plectostylus-gronp
are s u m m a r i z e d . T h e m o n o p h y l y o f the g r o u p is c o r r o -
F i g . 174. Hypotheses o f phylogenetic relationships i n the Plectostylus-group.
iS5
BREURE, BULIMULINAE
F i g . 175. D i s t r i b u t i o n of Plectostylus
and
Discoleus.
156
ZOOLOGISCHE V E R H A N D E L I N G E N
l68 (l979)
I 0
§
1 s
hb
E
BREURE, BULIMULINAE
borated b y one character
157
( 3 ) , but the present data do not corroborate the
m o n o p h y l y o f Bothriembryon
s.str. T h e hypothesis
s h o w n i n fig.
174a
is
preferable, i n m y o p i n i o n , because it expresses the v i c a r i a n t d i s t r i b u t i o n o f Bothriembryon
( A u s t r a l i a n R e g i o n ) versus Discoleus
+ Plectostylus
(Neo-
tropical R e g i o n ) m o r e adequately. T h e austral d i s t r i b u t i o n pattern s h o w n b y this g r o u p (figs.
175 a n d 176)
is f o u n d i n other a n i m a l g r o u p s as well
(see
K e a s t , 1973) a n d m a y be the result o f continental d r i f t . A c c o r d i n g to C r a c r a f t
F i g . 177a. Hypothesis of phylogenetic relationships i n the Cochlorina-group.
BREURE, BULIMULINAE
159
borated; they are separated on account of the shell morphology. The same applies to Oxychona, Otostomus and Cochlorina. The relationships of Sphaeroconcha are not clear and the genus is only tentatively incorporated in this genus group. The genera Otostomus, Oxychona and Cochlorina are
Fig. 178. Distribution of the
Cochlorina-group.
BREURE, BULIMULINAE
161
character states in Drymaeus (Mesembrinus) suggests that dispersal was primarily directed from South America to Central America. Fig. 179 summarizes alternate hypotheses of phylogenetic relationships i n the Simpulopsis-group. The monophyly of the group is corroborated by two
Fig. 180. Distribution of the Simpulopsis-group. 11
162
ZOOLOGISCHE VERHANDELINGEN 168 (1979)
characters (15, 2 0 ) . I n fig . 179a relative importanc e is assigned to character (9)
a n d the occurence o f character
(3 )
m a y be the result o f
convergent
evolution. I f relative importance is assigned to character ( 3 ) the result as s h o w n i n f i g . 179b
emerges. T h e g e n u s Simpulopsis
shows a disjunct d i s -
t r i b u t i o n ( f i g . 180): M e x i c o / G u a t e m a l a ( N u c l e a r C e n t r a l A m e r i c a ) , n o r t h e r n A n d e s , coastal p a r t o f the G u i a n a S h i e l d , eastern B r a z i l ( C o a s t a l a n d C e n t r a l B r a z i l i a n S h i e l d ) . T h i s possibly indicates that Simpulopsis (see, e.g., C r o i z a t , 1976: t e r n s ) . Leiostracus
is a n o l d g r o u p
figs. 57, 70, 77, etc. f o r s i m i l a r d i s t r i b u t i o n pat-
is f o u n d i n G u i a n a / S u r i n a m ( G u i a n a S h i e l d ) a n d eastern
B r a z i l ( C o a s t a l B r a z i l i a n S h i e l d ) ; Rhinus Venezuela (Llanos Plain +
is f o u n d i n the latter r e g i o n a n d i n
M a r g a r i t a I s l a n d ) . T h e s e d i s t r i b u t i o n patterns
are not i n contradiction to the hypothesis s h o w n i n fig. 179a, but are m o r e i n accordance, i n m y o p i n i o n , w i t h the hypothesis o f fig . VIII.
179b.
INTRAFAMILIAR RELATIONSHIPS
I n the present study the f a m i l y B u l i m u l i d a e is c o n s i d e r ed s e n s u lato, v i z . comprising
five
subfamilies:
B u l i m u l i n a e , Placostylinae,
Odontostominae,
O r t h a l i c i n a e a n d A m p h i b u l i m i n a e . P r e v i o u s authors, e.g. Z i l c h ( i 9 6 0 ) , have g i v e n p a r t o f these subfamilies f a m i l i a r r a n k (all except Placostylinae that were
integrated
with
(Bothriembryontidae)
B u l i m u l i d a e ) . Iredale for
the
anatomical research ( P i l s b r y ,
(1937)
A u s t r a l i an members 1946b;
has of
erected the
a
family
Bulimulinae;
B r e u r e , 1978b) has s h o w n that this
classification is incorrect. T h e relationships o f the five subfamilies are n o w investigated, u s i n g o w n observations
( B r e u r e & S c h o u t e n , i n p r e p a r a t i o n ) a n d data f r o m literature
(Bulimulinae: K o n d o , 1948;
see
references.
Placostylinae:
R e n s c h & R e n s c h , 1935;
Clapp,
1923;
Climo,
1973;
S o l e m , 1959a; S t a r m i i h l n e r ,
1970;
T u r n e r & C l e n c h , 1972. O d o n t o s t o m i n a e : A r a u j o , 1963, 1973, 1975b; H e a t h , 1914;
H y l t o n Scott, 1951,
Amphibuliminae:
Breure,
1952,
1966, 1967c. O r t h a l i c i n a e : V a n M o l ,
1974c;
Ihering,
1886;
Leme,
1968;
1971.
Van Mol,
1971). W i t h the data at h a n d the relationships o f the subfamilies can o n l y tentatively be indicated a n d f u r t h e r research is r e q u i r e d . T h e character t r a n s i t i on series that have been recognized are s u m m a r i z e d in Table
5 a n d c o n c e r n the mandíbula,
[characters
that involve convergence
r a d u l a , genitalia a n d p r o t o c o n c h
(e.g., r a d u l a structure, penis
spermathecal duct) are left out o f the t r a n s i t i on s e r i e s ] : (1)
sheath,
the n u m b e r o f
plates o f the mandíbula is u s u a l l y relatively large ( 2 0 o r m o r e ) . A
reductio n
i n the n u m b e r o f plates is hypothesized to be a p o m o r p h o u s . ( 2 ) T h e surface o f the mandíbula m a y be smooth o r granulate. T h e latter character state is
M
Os 00
164
ZOOLOGISCHE VERHANDELINGEN 168 (1979)
considered a p o m o r p h o u s .
(3)
The
internal structure o f the mandíbula is
n o r m a l l y m o r e o r less a m o r p h o u s . I n one s u b f a m i l y the internal structure is c o l u m n a r ( B r e u r e & S c h o u t e n , i n p r e p a r a t i o n ) a n d this is hypothesized to be a p o m o r p h o u s . (4) T h e presence o f a relatively large genital a t r i u m ( i n some species w i t h the a p o m o r p h o u s . (5)
7> 8, 9 ) , the m o n o p h y l y o f the O d o n t o s t o m i n a e b y two (1, 14), the m o n o p h y l y o f the O r t h a l i c i n a e + A m p h i b u l i m i n a e b y two (11,
13), the m o n o p h y l y
o f the O r t h a l i c i n a e b y three (3, 4, 6 ) , the m o n o p h y l y o f the A m p h i b u l i m i n a e b y two (10, 12). N o r the m o n o p h y l y o f the O d o n t o s t o m i n a e + n o r the
monophyly
of
the
B u l i m u l i n a e is c o r r o b o r a t e d b y
Bulimulinae apomorphous
character states, w i t h the data at h a n d . Solem
(1959b:
305, 327)
suggested a relationship between the
western A u s t r a l i a n Bothriembryon Placostylus,
and N e w
H e b r i d i a n Diplomorpha
b a s i n g h i m s e l f o n anatomical data. T h e present data,
southand
however,
BREURE, BULIMULINAE
165
do not j u s t i f y s u c h a relationship. S o l e m (o.e.: 318, 327) also suggested that Placostylus
has a n o r t h e r n o r i g i n . N e w
data o n continental d r i f t a n d plate
tectonics m a k e a n o r t h e r n o r i g i n f o r this g r o u p u n l i k e l y ( c f . C r a c r a f t , 1975a). Instead, m y suggestion is, (although hypothetical too) that the
Placostylinae
reached their present d i s t r i b u t i o n area v i a W e s t A n t a r c t i c a a n d N e w Z e a l a n d a n d that Bothriembryon
reached A u s t r a l i a v i a E a s t Antárctica.
T h i s hypoth-
esis makes the d i s t r i b u t i o n o f " p r i m i t i v e " a n d "advanced'* Placostylus o.e.:
fig. 20) m o r e plausible a n d , m o r e o v e r , makes clear that
a n d Placostylus
(Solem,
Bothriembryon
cannot be closely related.
F i g . 181. D i s t r i b u t i o n o f B u l i m u l i d a e . An
interesting question that remain s is: h o w o l d are the s u b f a m i l i e s ?
answer
is
only
partially
possible
and
is
rather
speculative,
but
The
Cracraft
(1975a: fig. 4 a n d a d d e n d u m ) gives some u s e f u l data. B a s i n g m y s e l f o n his data, the o r i g i n o f the f a m i l y lies between 80-95 m.y. ago (as n o b u l i m u l i d s are present i n A f r i c a ) , i.e., i n the Cretaceous. T h e P l a c o s t y l i n a e are ca. 80 m.y. o l d , as N e w Z e a l a n d separated about that time f r o m A n t a r c t i c a . L a c k o f geophysical data, however, prevents speculation as regards the t i m e o f o r i g i n o f the other subfamilies. IX.
SUMMARY
T h e relatively large f a m i l y B u l i m u l i d a e is m a i n l y d i s t r i b u t e d i n the
Neo-
tropics. T h e f a m i l y m a y be d i v i d e d into five s u b f a m i l i e s : B u l i m u l i n a e ( S o u t h
M
—.
M
167
BREURE, BULIMULINAE
a n d C e n t r a l A m e r i c a , southern U n i t e d States, A u s t r a l i a ; 43
(sub)genera),
Odontostominae
Zilch,
(eastern
South America;
18
(sub)genera;
O r t h a l i c i n a e ( n o r t h e r n S o u t h A m e r i c a , C e n t r a l A m e r i c a ; 15
i960),
(sub)genera;
Z i l c h , o.e.), A m p h i b u l i m i n a e (eastern S o u t h A m e r i c a , W e s t Indies; 4 genera; B r e u r e , 1974c) a n d Placostylinae ( N e w Z e a l a n d , M e l a n e s i a ; 15
(sub)genera;
Z i l c h , o.e.). S o f a r the classification o f the genera was m a i n l y based o n the sculpture o f the p r o t o c o n c h ( P i l s b r y , 1896a). T h i s character is still u s e f u l f o r i d e n t i fication, but it cannot be used f o r a phylogenetic analysis, n o r c a n a n y other m o r p h o l o g i c a l character o f the shell. O n the c o n t r a r y, the r a d u l a shows
a
v a r i a t i o n that phylogenetically m a y be analized. T h e v a r i a t i o n m a i n l y c o n cerns the shape o f the teeth, the n u m b e r o f cusps, the presence o r absence of s u p p o r t i n g denticles a n d the shape o f the transverse rows. T h e palliai organs show v a r i a t i o n i n the p r o m i n e n c e o f the veins,
the
position o f the p e r i c a r d a n d the structure o f the adrectal ureter ( o p e n o r closed). F u r t h e r v a r i a t i o n i n the anatomy is f o u n d i n the genitalia, v i z . the s p e r m o viduct
(position o f
presence
o r absence
proximal
sheath;
epiphallus
glandular folds), of
spermathecal duct
a n a p p e n d i x ) , penis
shape o f
(presence
penis l u m e n ; n u m b e r o f
( i n some g r o u p s i n t r u d i n g the penis)
(relative
length;
o r absence
glandular
of
a
celltypes),
a n d flagellu m
(place o f
insertion o f the retractor m u s c l e ) . I n the systematical part a s y n o n y m y , description, the d i s t r i b u t i o n a n d a list o f
taxa
are presented
f o r each
(sub)genus
o f the B u l i m u l i n a e . T h e
n u m b e r o f ( s u b ) g e n e r i c taxa used so far (80) is n o w r e d u c e d to 43. A c c e p t i n g the methods o f H e n n i g the g e n e r a can be d i v i d e d into s i x g r o u p s . T h e present data do not allow these g r o u p s to be fitted into one phylogenetic scheme f o r the whole s u b f a m i l y . T h e relationships between the five
sub-
families also r e m a i n tentative. T h e c o n c l u s i on o f a n analysis o f the e x i s t i n g biogeographical theories is that, after p r i o r phylogenetic analysis, the vicariance theory o f C r o i z a t m a y be adopted. T h e
distribution of
the B u l i m u l i n a e is e x p l a i n e d , u s i n g this
theory, i n the light o f the geological h i s t o r y o f the southern continents. RESUMEN La
família B u l i m u l i d a e , que es relativamente grande, se halla p r i n c i p a l -
mente e n la r e g i o n neotropical. S e puede d i v i d i r l a f a m i l i a e n cinco s u b familias: B u l i m u l i n a e ( A m e r i c a del S u r y C e n t r a l , l a parte s u r de los E s t a d o s Unidos,
Australia;
43
(sub)géneros),
Odontostomine
(la
parte
este
de
A m e r i c a del S u r ; 18 ( s u b ) g é n e r o s ) , O r t h a l i c i n a e ( l a parte norte de A m é r i c a
170
ZOOLOGISCHE VERHANDELINGEN 168 (1979)
A R A U J O , J . L . B . , 1965. E s t u d o comparativo de duas espécies do género C o c h l o r i n a J a n , 1830 (Gastropoda, P u l m o n a t a, B u l i m u l i d a e ) . — R e v t a brasil . B i o l . , 2 5 : 191-198. , 1971. Sobre a m o r f o l o g i a de Simpulopsis citrino-vitre a ( M o r i c a n d , 1836) ( M o l l u s c a , Gastropoda, P u l m o n a t a ) . — A r q . M u s . nac. R i o de J . , 5 4 : 77~8o. » 1973· Superfamília Bulimulace a do B r a s i l . Odontostomidae : A n o s t o m a depressum L a m a r c k , 1822 ( M o l l u s c a , Gastropoda, P u l m o n a t a ) . — R e v t a brasil . B i o l . , 3 3 : 11-18. , 1975a. Superfamília B u l i m u l o i d ea do B r a s i l . A m p h i b u l i m i d a e : Simpulopsis ovata (Sowerby, 1822). — A r q . M u s . nac. R i o de J . , 55 : 15-20. , 1975b. Superfamília B u l i m u l o i d ea do B r a s i l . Odontostomidae: Confirmação d a validade de A n o s t o m a ringens (Linnaeus, 1758), c o m u m estudo morfológico complementar ( M o l l u s c a , Gastropoda, P u l m o n a t a ) . — A r q . M u s . nac. R i o de J . , 5 5 : 21-28. ARAUJO,
J . L.
B.
&
A.
S.
H.
BREURE,
1977.
Notes
on
Bulimulidae
E u t h y n e u r a ) , 7. A n a t o m y a n d histology o f Simpulopsis P f e i f f e r , 1856. — Z o o l . Meded. L e i d e n , 5 2 : 19-25.
(Gastropoda,
(Simpulopsis)
miersi
A R A U J O , J . L . B . , H . E . B . R E Z E N D E & P . A . DE F R A G A RODRIGUES, i960. S o b r e B u l i m u l u s
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