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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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7: 97-99, 1907. Descriptions of new species of D r y m a e u s f r o m P e r u , M e x i c o , etc. — P r o c . malac. Soc. L o n d . , 7: 304-305. D A L L , W . H . , 1885. Notes on some F l o r i d i a n land and fresh-water shells w i t h a revision o f the A u r i c u l a c e a of the eastern U n i t e d States. — P r o c . U . S . natn M u s . , 8:255-289. , 1892. O n some types new to the fauna of the Galapagos Islands. — N a u t i l u s , 5 : 97-99, 1893a. B u l i m u l u s proteus B r o d e r i p and its distribution. — N a u t i l u s , 7 : 26-27. , 1893b. P r e l i m i n a r y notice of new species o f land-shells f r o m the Galapagos Islands, collected by D r . G . B a u r . — N a u t i l u s , 7: 52-56. , 1893c. L a n d shells of the genus B u l i m u l u s i n L o w e r C a l i f o r n i a , w i t h descriptions of several new species. — P r o c . U . S . natn M u s . , 16: 639-647. , 1895a. N e w species of land shells f r o m the Galapagos Islands. — N a u t i l u s , 8 : 126-127. , 1895b. Synopsis of the subdivisions of H o l o s p i r a and some related genera. — N a u t i l u s , 9 : 50-51. , 1896a. Diagnoses of new mollusks f r o m the survey o f the M e x i c a n boundary. — P r o c . U . S . natn M u s . , 18: 1-6. , 1896b. Insular land shell faunas, especially as illustrated by the data obtained by D r . G . B a u r i n the Galapagos Islands. — P r o c . A c a d . nat. Sei. P h i l a d . , 1896: 395-460. , 1897. R e p o rt on the mollusks collected by the International B o u n d a r y Commission o f the U n i t e d States and M e x i c o , 1892-1894. — P r o c . U . S . natn M u s . , 19: 333-379. , 1900. A d d i t i o n s to the insular land-shell faunas o f the P a c i f i c coast, especially o f the Galapagos and Cocos Islands. — P r o c . A c a d . nat. S e i . P h i l a d . , 5 2 : 88-106. , 1905. R e p o r t o n land and freshwater shells collected i n the Bahamas i n 1904 by M r . O w e n B r y a n t and others. — Smiths, misc. C o l l . , 4 7 : 433-452. , 1909. Repor t on a collection of shells f r o m P e r u , w i t h a summar y o f the l i t t o r a l marine M o l l u s c a o f the P e r u v i a n zoological province. — P r o c . U . S . natn M u s . , 37 : 147-294. , 1910. O n some land shells collected by D r . H . B i n g h a m i n P e r u . — P r o c . U . S . natn Mus., 3 8 : 177-182. , 1912a. Report on landshells collected i n P e r u i n 1911 by the Y a l e expedition under P r o f e s s o r H i r a m B i n g h a m , w i t h descriptions of a new subgenus, a new species and new varieties. — Smiths, misc. C o l l . , 59 (14) : 1-12. 12