tot de

Bijdragen

A black, Proteus

Sket

B.

anguinus parkelj

&

from the karst

non-troglomorphic amphibian n.

64

Dierkunde,

SPB Academie

of

(1)

33-53

(1994)

The

Hague

Publishing bv,

Slovenia:

(Urodela: Proteidae)

ssp.

J.W. Arntzen

1Department of Biology,

Biotechnical

P.O. Box 486/492, 61000

Faculty,

Vecna pot 111,

University of Ljubljana, 2

Slovenia;

Ljubljana,

Binnen

1013 JA

Oranjestraat 10,

Amsterdam,

The Netherlands

Urodela, Proteus, taxonomy, morphology, allozyme

Keywords:

Abstract

tions

diffèrent

A

distinct

morphologically

from southeastern Slovenia

anguinus as

P.

parkelj

a.

cavernicolous

pigmentation, fully developed shorter that

bones

gives

the head

trunk with and

a

a

higher

shorter

a

fewer

and

tail.

the

to

from Stična

(DNei

dissimilar

cally

and

=

to

with

traits

An

in

a

dark

broader

and

musculature

jaw

proportionally longer shorter

extremities,

considered

are

allozyme analysis

pigmented

taxon to be

to

over

différenciation

tiennent connu

bien la

Both

populations in

geographically

Postojna (D Nei

but

The new taxon is P.

a.

parkelj

protégé

par

lignées

fort

Le

la loi,

P.

a.

et il est

parkelj

n.

1

History,

habitat, and

are

geneti-

The

observed

only

a

level

of

from

under strict

the uppermost

single species a small

area

is

name

Miocene

recognised. rare.

is threatened

discovery

Ehre

des

in the

Postojna

Jersinovic

by

by

remarquable de

comme

Proteus

sous-espèce

les yeux bien

Slovénie du sud-est

anguinusparkelj nominative

développés,

de

le

par

ssp.

la

le crâne à

de

ainsi

ces

caractères

pigmentation

(

=

DNei

dents,

que par

morphes. L’analyse à

de

0.23)

localité située

des

foncée

à la à

la

développée (ce qui

corps relativement

vertèbres,

plupart

to

Val-

in his

trea-

von

Crain".

caves

With its

in Central Slovenia in

Loewengreif

1797

first known

(Bela Krajina) Celui-ci

n.

se

pigmentation très os

plus

courts

est

distinfon-

places

caves

formally in

where it

occasionally after

was

found

specimens

(Fitzinger, specialised

washed

J.N. in

species

a

musculature donne à la

plus allongé

les pattes et la

monotypic

1768, obtained his material from such

location in Cerknica

near

the

ressemble

plus

comme

à 40 loci montre

Postojna

au

point

de

aspect

plus

supérieur

courtes.

étant

vue

La

plésio-

ce

ces

genus

surface

caves.

compasses the

Dinaric karst

almost

anguinus

entirely,

the lower reaches of the Isonzo-Soca River in in the northwest

to

the river

vina in the southeast.

Trebiänjica

Altogether

in

en-

from

Italy

Hercego-

almost 200 locali-

taxon

ties

are

known

(Sket,

1983;

Sket

&

Aljanëiü,

in

génétique

population troglomorphedépigmentée d’une

proximité (Stična). Cependant,

a

of

who

élargis,

et

que

out

Laurenti,

mandibules

un

et à nombre

queue

sont considérés

allozymes

des

tête

waters, but in the open

were

were

heavy rainfall.

described the

cave

The known distribution of Proteus inférieur

fortement

massif),

par

animal.

when

nombre

plus

rare;

menacé

pollution.

Un Protée

un

est

species

Herzogtums

1850) Proteus became the

the

la

ssp.

Proteus goes back

who mentioned the

(1689)

"Die

tise

most

cée,

décrit est

apparemment

genetic

and may be

legal protection,

of the

history

population

Résumé

gue de

appar-

distribution

The normal habitat of Proteus is

décrit

depuis

qu’elles

taxon nouvellement

restreint,

de

occiden-

distinctes

considère tout de même

espèce.

niveau

populations

industrielle.

pollution

cave

industrial

même

territoire

des

Le

0.49).

=

montre que les

40

genetically

turn

distant

more

lineages since

hitherto

only known

now

une

population géographique-

Nei

be

that western and southeastern Slovenian

separate

conservatively

The

0.49).

=

differentiation suggests

populations form

à

D

représentent

supérieur; on

d’un

que

vasor

from

génétique

tales et sud-orientales le Miocène

d’une

(Postojna;

troglomorphic neighbouringpopulation

0.23).

a

a

vertebrae,

states.

dark

new

white

a

of

Most of these

loci has

skull

a

described

anguinus

a.

voluminous

a

appearance,

number

character

similar

eyes,

teeth,

bulky

plesiomorphic shown

(Bela Krajina) is

It differs from P.

n.

ssp.

Proteus

salamander

génétiquement

distante

plus

ment

distribution

analysis, ecology,

deux

popula-

prep.). Proteus

is

highly

specialised

troglomorphic,

B.

34

I. Southern

Fig.

of white

require as

P.

a.

confirmation.

springs and

Vir

Slovenia with

and

caves.

Rupnica.

area

The

insert

highlights

Scale bar is 5 km.

Three vertical

of Dinaric

solid

bars

variation.

Proteus

Normally

weakly pigmented.

Karst

indicated

the

Bela

Krajina,

with

is

(from

However,

left to

right)

morphological

blind

and

it is able

the

at

to

syn-

thesise dark pigments and reared in the light it may become dark

gradually al., eye

degenerate 1973).

No

and

&

tempt

to

initial

by skin (Durand,

metamorphosis

failed

the

Schreiber, 1912). accepted by

subspecies

The only

serious

some

the

most

who described

name

Hypochthon

Of all those

level

; is anguinus

genus. The

(1850)

genus

as

taxa

(Mertens Proteus

only

&

seven

(see

at-

sister

spealso

records of P. dots

Planina

anginus zoisii.

at

Proof the

a.

Solid

of

and Planinska

Research

and

Arts).

and

unusual

Doblicica Slovenia

et

al.,

1986;

the

at

on

springs,

to

allow

Sciences

eyes

in

the

southeastern

Proteus is pop-

level

a

lot

(AljanCiC

1986; IsteniC,

1987;

sight-

some

M.

was

Zlokolica be-

2.5

1990. In the

April 21,

of

specimen

Miss

by

sufficient number of

obtained

springs

resulted in

popular

Bulog,

photographed

Doblicica,

the

caught

in

1986. As

Another black

low the Jelsevnik

a

18,

Istenic &

1993).

such

waters

with

developed

was

Slovenes, this finding

publicity, mostly

and

with

Crnomelj

near

October

hypogean

Academy

specimen

body proportions

of

Hodalic,

to

records

refer to reports that

Jama); S, Stična,

(Slovenian

spring, on

to documented

symbols

access

black Proteus

Črnomelj.

and

One black

among

1992

open

A

-

squares refer

documenting places

(Postojna

Karst

ular

Arntzen

parkelj;

Trieste, Ljubljana,

the cities

km

period

specimens (n

morphometric

north of

up

15)

=

to

was

and other investi-

gations.

H. zoisii has

Müller, 1940)

only living representative

likely

denticulated line.

ed

1941).

differentiation.

by Fitzinger

teus

et

later on in life the eyes

induce

Seliskar,

the

development,

retarded;

to

(Aljancic

elaborate upon the taxonomy of Proteus

cies under

been

black

obvious characters exist that suggest strong

intraspecific

was

is

even

become covered

Attempts

(Pehani

the

larval

1986). During development

or

a

black

D, Dobličica; J, Jelševnik; P, indicate

by

to documented

diamonds refer

neotenic and shows limited

strictly

most

the

anguinus populations;

Sket & J. W.

For the

purpose of the present paper

fer

to

to

unpigmented

densely pigmented and

Proteus

slightly

as

we

will

"black"

pigmented

re-

and

ones

as

"white".

Proteus is the

taxon to

genus Necturus from the New World

(Duellman

& 2 Material and methods

Trueb,

1986).

The first documented teus

in

nature

is that

by

finding of

a

"black" Pro-

members of the Institute for

Samples mainly

of

Proteus used

from the

or

following

mentioned

Slovenian

in

this

localities:

study originate (1)

the Na Trati

Bijdragen

de Dierkunde, 64

tot

at Jelsevnik

spring

usually

referred

springs,

both at

Planina,

at

Planina

that

Only specimens have for

been

forms part as

that

specimens

of the material

the

later transferred to 70%

was

or

were

maceration of the

prepared by

of KOH

and its skeleton then

dyed by

fixed in

alizarin

and

a

alizarin.

ethanol

and at

present

Biology

of the

Biotechnical

osteological

Some

from Planina tail

of

tips

cleared

and

some

taken from live

were

measured

Data

on

tron

Luknja

well

as

from

1992). The

phenotypic plasticity,

standard

clinical

31-32 kV

were

the

listed

in

Table

vertebrae, netic A

neither

tris

two

in

specimens

In

addition,

a

skinned

obtained

were

were

morphology

of the

head. elec-

by

gills was

were

For

a

specimens settings

most

Siemens

filter. In

were

collected

taken

40-41.5 kV

set at

obtaining

useful

with

"Selenos-4",

newly

using and

nor

by

a

specimens Siemens

a

1.6—2.5 mA.

of the

pictures

by X-ray photography,

a

tiny

nuclear

tail

mag-

of Necturus

were

used

specimens

for

from

of neotenic

adult

lections

of the

snout to the

width

tail

the

from

length

side of the

point the

Vir)

head

for the

tip

length

to the

length

head

of the

width

snouth

the hind

posterior leg (LiE);

side the

is

Jezero,

purpose.

tip

the Uni-

(Jelsevnik,

measurement

from the

from the

and

stored in the col-

four localities

subjected

total

the

alpes-

Slovenia

same

bases of the most anterior

where

between

were

variables:

tip (L);

connecting the

at

length trunk

and

in

Museum, Vienna, and

Ljubljana, originating from

following eight

the line

cleared

individuals, including specimens

Planina, Rupnica,

of

of Triturus

Alps

from Trnovicko

phenotype

of the Naturhistorisches of

versity

specimens

the Kamniske

Maglic Mts., Montenegro were

A series of 82

and its

provenance

morphological comparison

Two

and the backbone.

(Laurenti, 1768)

of unknown

sp.

tip

of the

of the snout to

gills (Lc);

head

largest (Ltc); pre-pedal

to the

of the

length

anterior anterior

of the

legs (LaP); and

the fore

anterior and the

the hind

material

both

the

the

(cf.

the

were

not

&

three

specimens

dissection

of

after

in

18th

to

20th

a

to

of

analysis

use

Pa

x

for

was

counts

partial

a

data of

the

of

some

reliability

of

the

the out

for each

character

ceased

our

pub-

L

&

values

was

therefore variables

purposes.

in

individual

character

by

Arntzen &

1 (Wolterstorff, 1923;

starts

(Durand

of the continuous

"Wolterstorff Index"

the

and

matures

morphometric

absolute

effect of variation

the

variance,

In

size.

analysis using (WI

=

100

Wallis, 1993))

x

was

Principal Component Analysis (PCA) including

A

Discriminant

ried

require used

we

Proteus

comparative

value

univariate

a

ln-transformed

as

set

estimated

Mio. Teeth

completely

not yet

posteriori comparisons

ty

data was

comparison with

when

year,

consequence,

reference

order to reduce

addition

a

By

of limited

taken as

LiE

neither

Instead

cor-

data. its

Delay, 1981).

in

animal be-

product-moment

a

versus

the

the

counting

attachment to the

this would

order to confirm

reproducing, growth has

are

disarticulated

proved highly

with

Because

since

of vertebrae

X-ray photographs.

characters

precious specimens.

specimens

its

in

1977),

of Ver

regression equation

was

end of the

the vertebrae count

routinely performed,

Dolivo-Dobrovolsky (1926)

in

exists in

and

is not resolved

of 0.70.

coefficient

and

(Myo)

X-ray photographs

directly

posterior

meristic

In fresh

the last vertebra before

ambiguity

Parsons,

(1)

The number

(2)

counted

towards the

both

Romer

complete, for from

or

Some

particular,

in

correlation

At

con-

measurements

the

determined from

was

specimens.

surprisingly,

lished

tip

mm

off to their

of myomeres

including

cartilaginous pelvic girdle

backbone

own

of

also taken:

were

of legs.

example)

an

number of vertebrae

Not

to the

soft tissues and in

precision

the number

pairs

pelvic girdle (ilium)

related

posterior leg

a

taken to the tenth of a

were

be noted that in

from the atlas up to and

cause

tip

diminished.

counted between

the

side of

the

to

Presented values are rounded

It should

alcohol-preserved material

(Ver)

of insertion

point

meristic measurements

Two

calculated.

resonance.

specimen

the and

preserved

be

may

not

taken

X-ray photographs were

photographs

succeed

X-ray image eyes

on

a

80" apparatus

"Polydoros We did not

serving as II,

Ljubljana.

quantitative aspects

equipment.

120 mA

at

below

glass plate

of

preserved specimens. Eyes and in

tracted

cleared

70%

their

callipers.

vernier

using

and

in

of

perpendicularly stretched leg (PaL, PpL),

Measurements

(Led).

stained

pigmentation

on

of the tail

Plate I for

preliminary observations pointed to

further taken into consideration.

by

Data

the structure of the eyes and skin

microscopy (Bulog,

of

the

finally tail length from

(see

and Planina

Vir, Luknja,

specimens

intact

some

studied in detail but since marked

as

solution

Department

cleared

on

on

specimens

preserved

Novo Mesto.

near

from

of

One skele-

glycerol,

at the

studied

osteological study.

an

were

in

were

For

2% solution

a weak

longest toe

conse-

thymol, following

are

alive

a

poor.

Other

Faculty, University

details

from

Proteus

kept

are

specimens

allow

to

six

with

captivity

As

from

posterior legs

integer values.

were sacrificed

in

carcass

glycerol

black

(1967). Eight

Humason

in

preserved

in

ethanol.

KOH and parts thereof have been cleared in

with

cave

Postojna-

is rather

glutaraldehyde and

specimens

Ljubljana

of the

analysis (see below).

of most

text

Rupnica

Jama

that died

or

histological studies

ton

and

"Planina") (Fig. 1).

damaged

were

taxonomie

quality

to

except for

preserved,

biochemical

quence the

which

(referred

system

Vir

the Planinska

"Sticna"), (3)

Postojna

near

cave

Krajina (in the

the

of apart, 25 km E.S.E.

1.7 km

as

to

35

1994

-

the Bela

"JelSevnik"), (2)

as

Sticna,

(together referred

in

Crnomelj

near

to

(1)

data.

A

Analysis

among

further

on a

was

performed

carried

was

priori grouped specimens

independent variable. using

localities

analysis

The statistical

the SYSTAT 5.1

software

out

on

using

with locali-

analyses

were

car-

package (Wilkinson,

1989).

and

Blood, liver, heart, stomach, trolateral

Triturus, animals.

side

see

(in Proteus) below)

or

Erythrocytes

quot

volume

were

of 40%

M

NADP, adjusted

aqueous

future

supernatant

to

pH

was

of muscle from the

removed

M

was

10~

7.0 with decanted

3

ven-

Mertensiella

blood

diluted

tissues

and

and

stored

brief

an

5

x

centrifuged. at

ali-

ground

M EDTA and

HCl)

by

with

were

and

sacrificed

freshly

from the

Other

Tris,

(in

from

supernatant

sucrose.

homogenising buffer (0.1

strip

dissected

were

centrifugation. The plasma

a

from the tail

-70

°C

in

10~

5

The for

electrophoresis.

Polyacrylamide slab gel electrophoresis

and

staining

of

plasma

36

Sket &

B.

Electrophoretic conditions

Table I.

Triturus. Starch buffers citrate

8.1

pH

buffer

Table

(1971:

protein systems, corresponding to (XIII);

electrode

pH 7.4,

F:

1:10;

at

diluted

6.0

pH

B: Tris-citrate

buffer is 0.22 M

Tris-EDTA-borate

pH

pH

Tris,

8.9

40

7.0

examined

loci,

(I);

0.10

Maleic

M

(Ayala

et

Tissues used

4.1).

H

are:

E.C.

Protein

L

heart,

=

=

No.

liver,

M

=

acid,

(1970). PAGE P

D:

refers to

plasma,

=

(V);

Lithiumhydroxide-tris-

M EDTA and 0.01

0.01

and S

M

MgCl 2

phosphatase

Adenosine

deaminase

stomach.

Buffer

Tissue extract

3.1.3.2

Acph-2

A

L

3.5.4.4

Ada

C

L

Alb

PAGE

P

C

L

Albumin

-

Alcohol

dehydrogenase

Catalase Esterase

1.1.1.1

Adh-1,

1.11.1.6

Cat

3.1.1.1

General

Glucose

dehydrogenase

Glucose

phosphate

Glutamate

isomerase

oxaloacetate

transaminase

Est-2

Est-l,

P

L

1.1.1.47

Gdh

C

L

5.3.1.9

Gpi

A

L

2.6.1.1

Got-1

G

L

C

L

1.1.1.8

Gly-I, Gly-2

1.1.1.42

Icd-1,

1.1.1.27

Ldh-1,

dehydrogenase

Leucine Malate

3.4.11

aminopeptidase dehydrogenase

Malic enzyme Mannose NADH

phosphate

isomerase

Ldh-2

Mdh-1,

1.1.1.40

Me Mpi-

1.6.99.2

1, Mpi-2

Nadhdh-1,

Nadhdh-2

L

Pgd

A

1.1.1.14

Sdh

A

L

dismutase

1.15.1.1

Sod-1,

E

L,M

PAGE

P

E

L

Xanthine

performed according

were

gels.

was

The enzymes

with

more

ples,

were

than

assigned

most

only

one

showed

were

Tyrosine

and

enzymes

while

most

was

assayed

Presumptive

numbers and

activity

GOT,

could

substrate

Glycyl-Glycine, PEP-2

Proteins

be

two

clearly

supplied

as

and buffer loci

and

sys-

alleles

letters, respectively,

zones

in

sequence

with

protein

for

resolved

substrate.

scored, although

Albumin teristics tion

in

To

assess

loci

the

tations

of

(He)

In order

to

and

populations

and its standard

tripeptide Leucyl-

migrating the

sam-

PEP-1

and

dipeptide Leucyl-

Three

plasma proteins

intermediate

the

of

we

error

values, Ncj

against

method

loci

calculated

based

the

of

on

standard

is that the

within

the

as

characfrac-

the Proteus counted the

unbiased

error

distance

into

Sokal,

we

(Nei,

differentiation

measure

esti-

expec-

1987). across

of Nei

which may, for low and

correlated

converted &

one

identified

Hardy-Weinberg

genetic

genetic

linearly

was

for

Arntzen, 1987).

outgroup taxa,

and

degree

were

phenotypic

genetic variability

(Sneath

this method

their

(Z3Nej ; Nei, 1972),

be

identified

the most concentrated

in &

and the

calculated

The matrix of D UPGMA

on

accompanying

the

assess

in

Proteus

degree

heterozygosity

digesting properties the

based

the

protein (GP-2)

not be

migrating fractions

Rafinski

polymorphic

mean

could

representation

plasma (cf.

individuals, populations,

mate for

These

unidentified

one

Transferrin,

and their

number

scored.

and

the

ob-

were

in

faster anodally

taxon. The

en-

the

when

were

corresponding electromorph(s)

of

activity

Xdh-2

Xdh-1,

forms. For two

consistently

cathodally

a

histochem-

Hopkinson,

observed

was

hori-

Got-1, respectively.

specificity the

starch in

standard

PEP-3, anodally migrating

resolved.

(1971). Enzyme

Harris &

anodally migrating

and

designated Acph-2 different

of

by

1970;

Table I.

one zone

from the

but

in

presented

systems, ACPH

Three were

Prasad,

few modifications. are

(electromorphs)

were

visualised

were

&

techniques (Shaw

tems used

to Maurer

performed using Connaught

Sod-2

Trf

-

1.1.1.204

dehydrogenase

electrophoresis

served,

L S

Pep-3

1.1.1.44

Transferrin

zyme

L

G

dehydrogenase

Superoxide

starting

L

C

L

Phosphogluconate dehydrogenase

when

C

A

2.7.5.1

1976)

L L

Pgm-1, Pgm-2

Phosphoglucomutase

zontal

F A

D

Pep-1, Pep-2,

ical

H,M

Mdh-2

3.4.11-13

proteins

L

A,B

Lap

Peptidase

Sorbitol

E

Icd-2

1.1.1.37

5.3.1.8

dehydrogenase

P,L

C

Isocitrate Lactate

L

D

PAGE

a-Glycerophosphate dehydrogenase dehydrogenase

D

G6pd-1, G6pd-2

GP-2

-

phosphate dehydrogenase

Adh-2

1.1.1.49

Protein

Glucose 6

pH

acrylamide gels according

=

Locus

gel

,

tris-citrate-borate

system

Acid

and

Proteus, Mertensiella,

genera 8.0

pH

black Proteus

A

G: Discontinuous

al., 1972);

muscle,

-

Arntzen

in the

C: Tris-citrate

Roman numerals refer to the buffer systems of Shaw & Prasad

(Poulik, 1957).

to Maurer

for 26

A: Tris-citrate

E: Tris-malate

(X);

buffer is electrode 8.2-8.7

are

J. W.

a

to time

1973).

phenogram

(Nei, 1987).

phenogram using A main

can

be

the

objection

interpreted in

Bijdragen

a

de Dierkunde, 64

tot

when rates of

phylogenetic sense only

satisfied

in

of

mity

evolutionary

measure

(Farris,

which

the

applied

is metric.

From the

1972).

one, in

of tree

ary

interpretation

terms

relaxed

This

Nei's

length,

assumed sister group of Proteus,

and

terbury siella used To

distinguish

data

show

advocated

of

generated

(Rogers,

R

evolution-

an

rooted.

As

Necturus,

the

compari-

from Can-

and two

Rize

near

in

Merten-

Turkey

the branches

tree

from

support

were

the

in

areas

for distance data.

by Lanyon (1985) for

analysis

(Swofford

Jelsevnik

locality

&

gills. is

cave

fauna

various size

to

3.1.2

Diagnosis.

sides,

was

the

obtain

that may

after

about the

us

the

the

as

The

all

at

only

as

prey

A dark

-

of P.

head with

a

3.1.3

legs,

data for white Proteus

ver-

of the

a

of the

violet

composition of

ly

drift nets of a

from

shells

gastropod

it is not

outflow,

(generally

generally

were

obtained

the

could

possible

to establish

one

body

brownish

hue. In

whitish

as

in

soot,

in

triangle

no

spring.

from seditheir

pale

with

a

dark

origin

with

body pale

surface

finely

An alcohol dark

the

we

taxonomie

assign

the black Proteus that will be named Pro-

status to

cells

a

bluish

in

black

near

Paratypes:

(J4)

from

All

Biology

J9,

which

spring,

of the

is

skin

(J5)

Doblice

in

in

and

one

Trati,

near

in

one

Crnomelj,

skele-

is ethanol

pre-

Department except

nr.

ZMA

Herp.

large

quantities

contains

more

numerous

than is found in white

pigment,

multicellular and

specimens

the

epidermis

small and

externally

present

in

of

dermis. The upper

Leydig

(Bulog,

1991,

eyes

are

are

is covered by

surrounded

by

a

distinct transparent

a

visible

as

black

white circle. The eye

9239.

or

similar

structures.

conjunctiva

but

The diameter of

the

conjunctiva

is

ca.

of

general

structure

epigean

amphibians

In

a

190

mm

long

bulb diameterwas

5% of the head its

bulb

with

a

similar

from to

1.3

(7%

in

retina.

Doblicica, mm

The

that

to

well-developed

specimen measured

is

length.

of

the

Lc)

for

Zoological Museum, Amsterdam,

under

prin-

and One

specimen (Dl)

Ljubljana, Slovenia,

a

years.

with dark brown

specimens,

contains

without lids

ethanol.

of the

seven

in ethanol.

disarticulated

Others:

the collection

of

the

and stored

from Na

female

(J 10), preserved

glycerol.

kept

a

(plate I), preserved

specimens.

has retained

coloration for

in the upper part of the

are

The

specimens (J1-J3, J6-J7, J9)

University

deposited

the Netherlands

six

mm

in

preserved

material

is

of 21

and stained

cleared

are

J8 is

nr.

Slovenia

locality,

same

Doblicica

served. of

Crnomelj,

embryo/larva

specimen

coll.

Holotype:

-

Jelsevnik

live

shade. Skin

pinky

preserved

branches.

dots that Material.

or

in

1992).

Sket & Arntzen.

anguinus parkelj

with

Ventral side of

preserved specimen (Dl)

dermis also

the black Proteus

discussion

is broad-

preocular region.

patterning.

granulated

brownish

glands Anticipating

snout

specimens

some

the

the

specimens

some

to

The

Description of

anguinus

black with

completely

very dark brown. The

only

or

almost

are

bordered black

pale

the

and

Jelsevnik

be collected

a.

below the

from

small

very

main

or

colour is

,

and

variability indices,

be taken from Table II. The dorsal and lateral

of the so-called

placed immediately

were

when

Holotype:

-

corresponding

mainly

ton

lateral

short tail.

The computer

cipal

one

short-

externally

convex

a

characters,

Results

teus

and

of the type series.

Description

Gills red

3.1

parkelj

and

with

and

angular

short

long trunk,

precisely.

3

red

with its red

pigmented

anguinus

morphometric

heavy rainfalls,

to Proteus

discharge continued

catches

were

empty

of the

ments

ending "j"

a

creatures

of the word

test

micro-computer

welling up

impression

an

serve

and mesh width

long

days).

animals Since

and

body

hypogean amphibian

subspecies

visible eyes, a

black

pronounced.

be

snouted

Limbs as

three

allegedly hypogean

a

Ljubl-

"parkeljni"

small

day,

with

Note that the

not

In

mythology.

sold. These

resemble the black

Selander, 1981).

visited

was

notified

"boiling-holes".To

holes

Nicholas

of the Devil's

one

"parkelj")

tongue are

can

BlOSYS-1

local inhabitants

the

of

(plural

parts The

around St.

is

"Parkelj"

-

Slovene Christian

for which which

uncertain, we applied the jack-knife

more

employed

programme

sion

the

on

strong

order is

branching as

Kingdom

from

1876)

in

outgroups.

as

the

(Waga,

we

parsimoni-

(l.aurenti, 1768)

United

Peterborough,

caucasica

genus

distance

metric,

D

available for

not

were

Triturus cristatus

seven

is not

to be

have

Etymology.

names

jana,

a

the most

of the

3.1.1

distance-Wagner

distance

trees

trees

appropriately preserved specimens

son, instead

the

selected. For

was

undirected

the

in

are

never

of unifor-

assumption

distance

undirected

37

almost

procedure requires

Since

resulting

ous

condition

Rogers' genetic

used

widely

is

rates

1972).

1994

-

evolutionary change

a

This strong and unreal

reality.

procedure

phyletic lines,

the

homogeneous across

(I)

and the diameter of its lens 0.2

preparations

for electron

mm

(glutaraldehyde

microscopy

were

made

by

38

B.

Table II. and

Morphometric

Vir. L

length; PpL WI

=

sented

total

=

=

data

length;

for Proteus

(mm)

Lc

=

head

posterior leg length;

Lcd

Wolterstorff Index. Values in

individually for

measured

all

length; =

tail

anguinus parkelj

Ltc

=

head

length; Myo

parentheses

are

width; =

n.

spp.

LaP

number

from

J. W.

and P.

Jelševnik

pre-pedal length;

LiE

of trunk myomeres; Ver

estimated from the

biochemically studied specimens

=

Sket &

of Led

regression

and for the type series from

trunk

versus

A

-

from

anguinus

a.

=

=

Jelševnik.

Arntzen

length;

number of neck L and

Ver

Planina, Rupnica, PaL

=

anterior

and trunk

versus

Statistical data

black Proteus

Myo.

include

Data the

leg

vertebrae; are

pre-

specimens

by Kranjec (1981).

Locality

Specimen

L

Lc

Ltc

LaP

LiE

PaL

Led

PpL

Myo

Ver

WI

31

(33)

10.8

number

JeUevnik Jl

227

27

16

31

130

14

13

64

J2

247

26

16

32

138

14

12

(69)

J3

240

25

17

34

140

16

13

67

J5

205

23

14

31

120

13

11

55

199

20

12

27

113

13

10

57

J8*

276

28

17

37

158

16

13

J9

232

24

16

30

135

14

7

7

7

7

.17

sample

size

mean

232.3

SD

24.7

26.0

7

15.4

2.69

1.81

31.7 3.15

7

133.4

14.3

14.6

-

10.1

-

35

11.4

34

10.8

30

(33)

11.5

78

30

34

10.1

13

65

33

35

10.4

7

6

12.1

1.25

32 -

5

64.3

1.22

4

31.2

8.19

7

34.5

1.30

10.7 0.57

0.58

minimum

199

20

12

27

113

13

10

55

30

34

10.1

maximum

276

28

17

37

158

16

13

78

33

35

11.5

PS

235

26

17

35

118

19

16

75

27

31

16.1

P6

255

30

17

37

129

19

20

82

28

31

14.7

54

54

54

54

54

54

54

54

53

54

29.0

15.3

36.0

17.5

72.6

25.5

30.8

17.1

Planina

sample

size

mean

54

223.2

SD

26.6

minimum

147

maximum

299

Rl R2

3.47 19

2.40

4.02

109.9

18.7

14.1

2.07

2.15

9.29

0.79

0.51

1.20

9

24

72

13

12

48

24

30

13.8

37

23

46

152

24

23

100

28

32

19.6

270

31

19

42

133

25

21

90

25

29

18.8

225

24

16

31

111

18

77

26

31

16.2

Rupnica

sample

7

size

mean

7

245.0

SD

40.6

1

7

27.7

17.5

4.82

3.37

36.6 6.45

7

15

7

118.4

7

20.4

21.1

7

17.2

3.40 16

2.53

minimum

188

23

14

29

88

maximum

294

34

22

45

144

219

24

15

32

108

19

14

14

14

19.1

16.7

21

7

7

25.4

11.7

15

25

7

83.9

29.9

0.54

17.3

0.69

1.64

67

25

29

15.7

102

26

31

19.7

76

27

31

11.1

14

14

Vir V7

sample

14

size

mean

SD

*

14

228.4

14

27.8

34.7

14

16.0

3.90

2.61

34.6 4.55

111.4 16.9

minimum

154

21

11

26

76

maximum

285

34

20

41

138

17

3.08 14

23

76.5

2.00

12.8

14

49

20

100

14

26.1

14

30.6

0.66

16.7

0.65

1.95

25

29

11.1

27

31

18.7

Holotype.

Bulog (1991, mm, of Lc

at

0.9

1992)).

mm

In

a

skinned

specimen

the eye bulb's diameter

was

of 205 3.9%

(ethanol preparation).

The head is slightly broader than the trunk and

approximately

11 %

of the

length

of the

body (L-

Led).

In its

sided,

the short

rounded

corners.

27%

Lc.

of

cushions

the head is

postocular part snout

The

Three

(that

we

shaped snout

pairs

of

as

a

parallel-

trapezoid

comprises

on

subdermal

tentatively identify

as

with

average

muscular the leva-

Bijdragen

de Dierkunde,

tot

64

(1)

39

1994

-

Fig.

3.

Rostral

ossification

Jelševnik

from

of the

of

part

and

the

ethmoidal

(specimen

body

trabecula

no.

The tail is

parallel.

than the trunk. The tail with

triangular

of the

body length

The skull is

(cf.

tioned

Cranial from

tale; f, frontale; pq,

and

mandibular

Jelševnik o,

operculare;

paraquadratum;

tores

q,

and

in

J4):

quadratum.

Scale

Proteus

a,

parietale;

p,

the head

depressores a

bulky

short, bushy gills of

no.

pm,

bar

anguinus

angulare; d,

5

den-

praemaxillare;

approximately The

trunk

is

are

mandibulae

appearance.

well

posteriores)

Three

developed

with

pairs a

of

length

average 58% of the

length

and

of the

mm.

built

similarly

differently.

of four in

placed

a

rounded

legs on

1926)

slightly

the

two

toes.

but

speci-

propor-

wider,

with

less than the

width

largest

equals

vertebrae. The ratio

pectoral

6%

vertebrae. The dentalia

pectoral

or

short

The poste-

that of white

to

that is

are

average

toes.

The neurocraniumis

wide arch,

of 2.3

lengths

The

form

running

slightly higher

or

broadly

tip.

legs

sides

shorter and have

rostro-occipital length

length

is

tip

fur-

com-

and flattened

length

Dolivo-Dobrovolsky,

occipital

length

averages

1.4

twice

as

long

anguilliform, body.

on

The flanks

to

are

the

rostro-

as

wide

rather

macerated skull

peculiar shape

and with convergent versus

The

other

with

that

its

we

In

ratio averages

of

less than

the

plaque

possess

rostral

protrude along

bones

are

The ethmoidal

tips (Fig. 3).

length

The frontalia slightly laria.

praemaxillaria

(Fig. 2).

only

mandibular width

largest

versus

1. The

in the

the width

20% of Lc.

cylindrical

anterior

mm.

mandibulae anteriores, levatores mandibulae

externi,

give

bones

(specimen

1

dorsal

and have three

slightly

are

legs

mens

2.

the

anguinus parkelj

with shallow

high

as

rounded

a

and slender. The anterior

parkelj

with

bar

running parallel

are

with the ventral and

laterally almost

Fig.

Scale

J4).

28% of the entire body

prising

a

cranii Proteus

between the myomeres. The tail is short,

rows

rior

in

plaque

has

a

tines curved the frontalia at

the

0.6

to

1.

praemaxil-

neurocranium

are

40

Sket &

B.

Table III. Number of teeth observed and in two outside

of P.

specimens

the main

and accessory

Maximal

row.

teeth.

in P.

taxa

In both

the

n.

ssp.

for white

numbers

from Luce and

anguinus (four specimens

a.

anguinus parkelj

from Jelševnik.

specimens

teeth

opercular

may

are

be

in

Figures

arranged

either in

one or

Vomer

in

Dolivo-Dobrovolsky, 1926)

to

denote the accessory

two to three

of teeth

black Proteus

A

-

for different combinations

Number

Praemaxillare

Arntzen

Stična, according

parentheses

given separately

J. W.

smaller teeth

of

standing teeth

principal (large)

rows.

on

Pterygo-

Dentale

Operculare

palatinum

P.

>* anguinus

a.

(minimum)

P.

P.

parkelj

a.

5

22

6

22

6

21

5

10 9

or

(2)

According

to

(2) 8

(4)

24

(8)

6

25

(8)

6

33 31

19

4

19

7

18

4

16

?

7 8

20

5

17

(6)

8

22

5

17

(4)

6

Dolivo-Dobrovolsky (1926).

the vertebrae

drifted larva of 21

was

in

did

not

legs

ones

only

are

are

Anterior

of

part

embryonic

bar

1

(specimen

larva

no.

of Proteus

J10)

at

operculare, teeth

are

The dentale bears 6 small main

auxiliary

to

so on

in

two or

three

rows.

19 teeth with another 4

teeth that

posterior legs is

of the

same

length

vertebrae get or

XXIV and

34

a

b

result

Cat

,

are

outside the

placed

as

The

the neural

XVII,

shape

of

a

sharp

posterior

crests

and

to

ones.

The caudal

smaller till number XXIII

no

spine

and

spines

owing

pectoral

then followed

tary vertebrae that have ber

35. The neck vertebrae are

the

gradually are

3.2

d

of the

in black

four alleles,

allozyme study,

Nadhdh-l

,

Comparison

a ,

and

Pgd

a

uniquely

were

,

specimens.

by

3

to

processes.

5 rudimen-

Up

to

num-

of the vertebrae is in the

caudally form

extended

irregular

the fact that

they

triangle.

trapezoidal are

with

related

3.2.1 General appearance,

gills.

to

grey and less

taxa

to

The number of vertebrae between the skull and the

specimen is pale

the dentale. On the

(Table III).

row

posterior

The eye di-

fewer teeth than in the

are

arranged

16

the

toes.

it is densely speckled

stage 21.

Cat

morph, particularly

short and three-toed, the

Ventrally

It

active feeder. Its

short stumps without

is 8% of Lc.

observed

white

an

J10)

no.

(Fig. 4).

anguinus

Briegleb

mm.

short and wide. There

21

the sides.

on

As

Scale

Jelševnik

high.

(specimen

have the appearance of

anterior

so

L

be

to

appear

mm

Briegleb's developmental stage

ameter

from

12

(8)

5

white. Dorsally

4.

12

(11) 23

or

5

A

parkelj

(2)

J4—right

ably short,

Fig.

23

or

J4—left

J5-right

*

8

or

J5—left

parkelj

a.

22

7

right

(maximum) P.

7

left

j* anguinus

a.

left

right

remark-

Proteus from other Dobliöica exhibit

sometimes

They

from

Differently

—

only

the

nearly

with pale

dark

or

pale

or

after

all

than Jelsevnik and

white

yellow

eyes, and

subspecies,

new

populations

a

become

pigmentation,

a

pinkish grey

long

skin,

patches.

exposure

to

light. In the stages val

hatching

21 of

in P.

is

a.

not

before

(corresponding

Briegleb)

of Lc,

immediately

the eye, with

evidently larger

to

a

an

or

after the lar-

advanced stage

diameter of 6

in P.

anguinus (corresponding

a.

to

parkelj

value is

8%;

8°7o

than mea-

Bijdragen

de Dierkunde, 64

tot

-

(1)

1994

41

has

non

been

3.2.2

to

276

mm,

in

our

sample

LiE,

than in

shape,

the

na

(a)

and

P.

a.

from Jelševnik

parkelj

(b).

and from

drawing

It

(Fig. 5).

behind the

et

the

of P.

embryo

al.

(1966)).

denser than in P. tation of adult P.

amphibians

a.

the

any

coloration

bright

Wayne King,

the

covered

might

be

skinned

skin,

shining

specimen

diameter

specimen. (Peters

morph

by

et

at

increase

sur-

vivid

species

are

without

Behler &

is small and without

morph

degenerated

through

in as

some

a

Lc

species

blackish

with

1985)

be

cases

It

concentration in

decrease of

gill

dot.

the

In

a

black

rudimentary

eyes

variable.

and ramification of the

oxygen

the eye

the diameter of the

highly

populations.

always

the eye bulb's

half that of

was

to

are

eyes

of white Proteus,

al., 1973; Sket,

or

most

more

in the genus Necturus,

most

although

As in other

able within white

varying

a

ing is

similar to that in Necturus. In

the

2°7o of

length

of

little

a

pigmen-

up

strikingly

morph

by

in

the

the head is

In the whites,

snout.

be

can

truncated

long,

a

with

pear-shaped

in

may the

gills

is

vari-

water

size. A similar

the

affects

phenome-

relative

due

triangle concave

posterior legs, strongly

head

length =

(productis smaller

0.94),

than in the white

morph

which

the

morph

than in the white value

corresponding in the black

morph

is

than in the white

Both

pairs

legs

49%.

Cornum-

the

legs

two toes

and with

a

at

are

the white

similarly

6

or

shorter in

skeleton

the

In

both

only

three

ones.

built with

the fore and hind

simplified

4; 5

morph. remarkably

are

than in

specimens

morphs and

of

length morph

the absolute

ber of trunk vertebrae and myomeres is

higher

Mak-

morph.

average 58% of L, relative trunk

in the black

respondingly,

limb, respectively,

(cf. Aljancic

&

Sket,

1964). The tail is Proteus.

longer

The tail

in white Proteus than in black

shape

tional variability but

quite from

constant

Planina,

height

be that

of the

length

with

correlation coefficient

on

larger

black

rudiment is expected The

differs from

in

1979).

superficially

white

only

(cf. Conant, 1975;

The eye in the black lids and

be

The intense

absence

of Proteus, group

pigmentation

seems to

pattern of coloration. Also sister

correlated

in the black

parkelj

a.

by

Briegleb (1962)

The dark

anguinus.

of both

flanks.

moment

parkelj

a.

in

of

shape

even

Vandel & Bouillon (1959)

photographs by

and Vandel

face

a

between

about

brought

data. In the black

The relative taken from

surements

(Lc, LaP,

weaker musculature behind the eyes, the whole

head has the from Plani-

L

to

found

The head differs than is

strongly

more

parallel-sided

anguinusanguinus

are

while relative Ltc

whites,

nearly equal.

morphometric

to a

of white

Relative Lc is smaller in the black Pro-

populations.

forms is

relative

differences

Led)

which falls

mm).

measurements

Lpa,

teus

147-299

(range

most

For

heads of Proteus

osteological

of the black Pro-

length

mm

Proteus

5. Skinned

and

morphometric

from 199

ranged

the

of medium size.

are

The total

—

(Conant,

populations,

within the range observed

teus

Fig.

morph

Univariate

comparisons.

Necturus

other Proteus

to

in the black

gills

for

supposed

1975). Compared

tail

as

tip,

mens

the

within

a

great interpopula-

contour

characters In

populations.

the tail with its

crest

usually bluntly

Rupnica

than the

body.

broadly

rounded. In the black

It does

not narrow

same

towards the

pointed.

and Vir the tail is

are

specimens

is of the

body, gradually lowering

which is

from

shows

some

In

speci-

usually higher

caudally

morph

and ends

the tail

tip

is

B.

42

6. Last caudal vertebrae in Proteus

Fig.

caudal

Fig.

of

vertebra

is indicated

by

an

anguinusparkelj

asterisk. Scale

from Jelševnik

bar is 5

mm.

shape

somewhere in between the recorded

ex-

a.

from Vir

anguinus

seen

here

from the

In the such

skull of the black

the

as

shorter

dorsally other

wider than in the white

visible intercalations

(dentalia,

the dentalia and The

morph.

in the white

the difference

praemaxillaria morph

angularia

morph

being

most

& Darevski with

1977)

is

was

differ-

as

formed

wider than in the

teeth

(Table III)

pronounced

Mioproteus on

its

less

markedly

some

are

than in the black

10—12 teeth

and the

shorter. Also

paraquadrata)

number of

tale. The fossil relative

Darevski,

are

flat bones

The mandibular arch such

ently shaped.

higher

and

parietalia, frontalia, or

bones

white

the

morph,

is

morph,

in the den-

caucasicus Estes vomer

(Estes

&

toothed.

cus

1993). tra

(b)

-

A

black Proteus

and Planina

The XVIIIth

(c).

perspective

same

the heads

as

found between Planina

and

(Rupnica

white

Sticna.

and

Vir),

Proteus

(cf. Herre,

The

Sticna

two

however,

are

also

from

populations

mostly indistin-

guishable. In

some

elongated, the black

white a

specimens

the

feature that has

morph.

neck

not

vertebrae

are

been observed in

Otherwise the trunk vertebrae

are

1935) In

alpestris.

vertebra is

in

gether

or

are

are

similar

although

specimens

of

the

are

specimens the

markedly

by

without

Alto-

till the

three

to

processes

from Vir the tail

inspected shorter.

black

In

the

speciwhite

from Planina the vertebrae are lower and

gradually diminishing

in size towards the a

specimen

has

vertebrae after the

spines (Fig. 6), another

ventral

between

specimens,

tail. One less

ver-

Triturus

and

spine

interspaces

followed

are

cen-

individual

gradually smaller

which

to

tail

substantially larger.

XXIVth and ones

caucasi-

Paleoproteus

regular

is variable. In black

shape

In the white

vertebrae

dorsal

and the

vertebrae get

rudimentary

mens,

centre

the

narrower

their

the caudal XXIIIrd

Proteus the

of the

than in

the

Schleich,

in all Proteus the

as

in neotenic and

vertebrae

subsequent

Estes &

the processes

developed

or

morph

Mioproteus

1977;

well

as

thinner,

are

process

five

are

populations

less well

in

stout as

particularly

are

in the black

even

Darevski,

In Necturus

and

tebrae

are not as

Estes &

(cf.

(Fig. 6). Some significant morphometric differences

and

similarly shaped, vertebrae

tremes.

by

P.

are

J. W. Arntzen

5.

a

are

(a) and

The vertebrae

Sket &

chain of

XXVIIIth

while there

specimen.

seven

are

only

tip

of the

small, spine-

that

still

bears

three of such in

Bijdragen

tot

de

64

Dierkunde,

(1)

1994

-

43

high component loadings interpreted

the univariate and

pendent white

and

PaL,

contrasting either

size is

analysis,

which is

de-

population

for

black

versus

The second axis is dominated by

PpL.

Component

which

signs,

short trunk and

a

axis,

size. In line with

not

discriminatory

not

populations.

LiE,

the first

on

representing general

as

have

loadings

that animals have

means

long extremities,

or

a

long

trunk and short extremities. Hence the second PC A axis

is

the

essentially

On the third axis with

with

Led,

separating

contrasting

specimens

overall

and

Rupnica

3.2.4 Protein

could be

symbols)

and P.

variate

number of trunk vertebrae

-

variate

(c):

nina,

parkelj (solid symbols). Upper

Wolterstorff Index

-

second

(PCA)

a.

and third

R

=

Rupnica,

of

axes

with centroids and

(WI, a

V

=

see

text);

(b):

lower

outline

Vir

(P.

polygons a.

for P

anguinus)

=

bi-

part

and

PlaJ

Data

Vir

(P.

a.

netic

—

and

multivariate

different in the black and white

(P< 0.001;

Table

II).

Similar

Triturus cristatus for which devised

morphometric

The Wolterstorff Index is

(Wolterstorff,

vertebrae (Ver) is

an

to

the

the WI

1923),

significantly of Proteus

morph

situation

was

in

originally

the number of trunk

equally good

or

out

of

popu-

of 21

speci-

23

different enzyme the

with the

(two loci and four

taxa

and

variability

exception of

Triturus, for

respectively).

the

of

variability

altogether

associated

populations

The Mpi-J

locus

40 pre-

estimates of ge-

presented

are

showed

no

in

variability

=

any of the

parkelj).

Bivariate

Rupnica

with 8

out

proteins

composition

and

loci

Table IV.

loci

11

better discrim-

variation The one

assayed populations,

showed

12.0

±

higher

for

the

4.0%,

Proteus

samples.

The

which is similar

=

(11.2

mean

1.3

±

to

3.3%)

±

Proteus

populations.

variability

combined Proteus

caucasica than H

for

while an addiacross

intra-populational

no

16 loci showed

of the Proteus

for M.

no

observed

was

remaining

ity ( He)

variation

In 12 loci, inter- but

samples.

analysis.

In

-

the outgroup

allelic

on

tional

3.2.3

morpho-

the

versus

morphometrically

are

with 7

plasma

Mertensiella

sumptive

over

Jelševnik

in

of black

uni-

Principal Component Analysis

convex

and

(Ver); details

(a):

part

to the middle

in

Rup-

the third

classified. The

consistently scored,

loci

some

analysis.

and three

systems

loci

espe-

classified.

(33°7o) wrongly

mens

that

statistically significant,

not

virtually indistinguishable,

anguinus anguinus (outline

helps

another,

versus

Planina

specimens (15%) wrongly

lations of

of Proteus

axis

separation

reveals

differentiation for

and Vir is

Morphometries

one

(Fig. 7c).

analysis

Discriminant metric

7.

(Fig. 7bc).

This

signs.

the second

Plotting excellent

provides

and white

Fig.

WI

dominate together

from

populations

Vir.

nica and

54

the

as

PpL

those from Planina and Jelsevnik from

cially

axis

all

same

Lc and

in

at

least

heterozygos-

populations

is

the value found

and

substantially

1.0% found for

T. crista-

e

inatory

feature than WI

1993) (Fig.

itself

(Arntzen

&

Wallis,

7 ab).

In the multivariate PCA all

tus.

Among Proteus populations

7.5% for the

eight

variables have

to

22.5

±

two

studied

6.7% for the

H

specimens

single

e

ranges

from

from Planina

individual from Vir.

B.

44

Table

IV.

Proteus, refers

ity

Allelic

composition

Mertensiella, and

to single individuals.

calculated

on

the

basis

for 40

gene loci

Triturus. Individuals

Missing of

data

are

Reference

no.

from two

indicated

by

a

and

measures of

populations of the hyphen.

Hardy-Weinberg expectation

Species

Locality

(upper part)

is the

are

corresponding

as

A

black Proteus

observed

in

the

standard

loci,

H

error

is the e

(in

average

percent

heterozygos-

values).

Mertensiella

Triturus

caucasica

cristatus

Rupnica,

Slovenia

Slovenia

P6

R1

R2

Vir, Slovenia

V7

genera

pooled. Allelic composition in parentheses

Proteus

Planina,

P5

genetic variability (lower part)

latter taxon

-

anguinus

Slovenia

J2

J. W. Arntzen

P is the number of polymorphic

and SE

Jelsevnik,

J1

Sket &

Azaklihoca

Köyü,

Rize, Turkey

ZMA 7277

Herp. (n

2)

=

Canterbury

and

Peterborough,

ZMA

Herp.

and 9200

(n

U.K.

9199 =

7)

Bijdragen

Fig.

To the left

8.

based

Nei’s

upon

correlation

rupted for

distance

line refers to

a

as

1994

-

for four

measured

To the

45

populations

40

over

gene

minimum

right (b):

of the tree is 0.62

Length

(1.75

with

loci.

taxa

the

jack-knife

included);

test

with “outline”

(white morph

estimated standard

length distance-Wagner tree

outgroup

branch that is not robust under

of Proteus

Boxes indicate

the

the basis

on

cophenetic

(see text). Outgroup

error

of

printed population name) The

(Nei, 1987).

cophenetic rooted

Rogers’ genetic distance,

correlation coefficient

taxa Mertensieila and

is 0.99.

Triturus

The inter-

are

included

comparison.

values of Z?

High

tively 0.56,

with Z)

Planina population

for

(0 Nei

from

Jelsevnik and

dard

error

and

the

of these

Nej ,

8a).

(Fig.

Using

with

the matrix of Z?

seen

from the root,

branches two

off

first,

Nei

(Fig. 8a).

supports In the mum

this

(Adh-2

d

distance-Wagner

the

clearly

not

in

a tree

from

populations

of

the results

tree

are

b ,

that

Jelsevnik.

the

seven b ,

Cat

b

A

Nadhdh-l ,

Sticna no

reference or

not

No

loci with

phenotype

ro-

most

Pgd

required

are

are

strictly

markedly

on

,

popu-

differ

0.14)

=

to

aut-

diagnostic

observed. The

(ö Nej

a

Proteus from

fixed alleles

were

and Vir

Rupnica

another

samples

these alleles

for the black

and ,

in the black

uniquely

Larger

lations from

a

Cat ,

whether

one

hence

resolution is found within the lat-

apomorphic.

from

separate

(Fig. 8b).

observed

out

charac-

and both

results and

Conflicting

Four alleles, were

Jelsevnik

—

both Ldh loci.

the

3.3

Ecology

and

reproduction of

the black Proteus

Rupnica

displays

Icd-2 ,

group

grouping together

of the

—

phylogenetic

3.3.1

Habitat:

present, P.

essentially

b

ter

the

Planina

jack-knife replicates

G6pd-P, G6pd-2

bust

potentially synapomorphic

which,

Jelsevnik and

distribution of alleles,

Alb ,

are

error

branching topology.

parsimony

Inspecting

by

Each of the

32% of

at

are

populations.

find

UPGMA-method

population

followed

remaining clustering

and Vir

the

results in

the

from

the stan-

cases

the standard

populations

the

population

averaging

taking

—

recorded

populations

In all

)

defining

ter states

from Planina

Substantially

were

black

white

estimates,

into account, the latter

separated

the

Vir, respectively.

is substantial and

0.24)

rela-

involving

the others.

0.23 and

between

comparisons

Rupnica

comparisons

versus

=

and Sod-2c

ob-

from 0.41

ranging

Nei

obtained for

lower values

were

Among Proteus populations

high values, were

8a)

among Proteus and the

comparisons

taxa.

outgroup

1.8; Fig.

(>

Nej

tained for all

£>

(1)

(a): UPGMA-dendrogram genetic

coefficient is 0.96.

the outgroup.

by

de Dierkunde, 64

tot

maxi-

two

same.

Krajina,

alleles

Mdh-2

d ,

a.

localities

abiotic

parkelj near

which

has

the

is in

parameters.

only

town

of

—

to

Crnomelj

the

in the Bela

the southeasternmost part

Slovenia. The Doblicica and Jelsevnik

discharge

Up

been documented for

into the river Doblicica.

springs

They

are

of

both

situated

46

in

close

proximity

below the

apart),

to

channels. At

On the

1991).

plain,

of

deep pool

25

ca.

black Proteus

autumn

the

on

depth

at a

of

level

Soon

after, but

there is

(called Jezero),

by

one

specimen group of

similar

of

(Aljancic

meters

the

(BS) inspected

us

were

the Doblicica

to

corresponded

response of the

At

seen.

and

springs

originates (HabiC The

by

of 11.2

perature of

a

100

to

holes

in

cm

eject

of

toring

only

When

black

specimens

collected meadow

in

at

the

Jelsevnik,

low

nearby epigean

large The

presence

a

as

surface

Physical nent

on

are

the

springs

were

standing

of the situation.

required

1990. In

10.0 and

water

the

they

a

by

and

hole

in

The

they

the

boiling springs fed

or

by

from

discharge

terrestrial animals.

partial origin

springs

surface

in

from

either

they

Close moni-

proper under-

of the permaet

autumn

sucked in

Upon

capture,

stomach

animals in its

hydrodynamically

cies.

Of

worms

these,

the

sp.

fluminensis cave

(1990) 1986—

are

night are

waters

in favour of

seen

to

be

the

have

(Sket,

open.

active

actively

1970).

In

mentioned for Doblicica is almost certainly

errone-

least

on a

registered

in

white

on

an

penetrated

that Proteus

captivity,

and

Sadlenot

forages

black

aggressive

illuminated

to

aquarium, Indeed,

at

Proteus

feeders

specimen during feeding

videotape.

earth-

These observations

attack

as

may,

at

sp.,

and the chilo-

has been shown

Kuester

pre-

prey spe-

aquatic gastropod

suggestions

in the

out

may

but

riana

of

Vitrea

gastropod

(Oligochaeta: Lumbricidae),

14.8 °C

value of

or

from the main

surface-dwelling

and bite their white counterparts.

extreme

stream,

food items

Surprisingly,

marginally

(the

drift

of the Proteus gave up their

consisted of

Lithobius

pod

some

contents.

dominantly

al.

of Doblicica and

outflow and associat-

unidentified

an

per-

springs (Table V).

deviate only

to

Some

-

from the

spring.

enter

temperature fluctuated between

expected

food.

ejected

the surface

from

originate

are

steady

no

fauna,

subterraneously,

Habic

and

Doblicica and Jelsevnik

were

background.

spring

biocoenoses

winter and

range

dis-

tem-

during upwelling

gain

11.3 °C and temperatures in are

from this

or

epigean aquatic organisms

the permanent

Jelsevnik,

to

animals have been

Since Na Trati has ed

probably

active,

some

measured in

especially

system,

heavy

found

accessory

and chemical parameters

17 occasions

summer

in

also the

was

was

boiling

been

rock.

some

When

some

stream

holes

most

be

Jezero

holes

boiling

stream.

afterwards,

soon

narrow

the drift of Na Trati suggests its a

The

from the mountain

of soil and

of

or

Doblicica,

zone.

amounts

can

Novem-

typical

year after

meadow

wet

well

as

a

occasionally

from

aquifers

karst

which

°C,

will be

manent

the karstified

the outflow of

the

few times

the

on

covering

on

southern

while the

(Na Trati).

active

are

straight

shallow local

a

have

around

crawling

holes

they

of the

10.3 °C,

periods,

stygobiont

rains.

water

the northern group of holes of

at

diameter

water

region

temperature of the

some

10.1

of the

Jelsevnik

at

to

water

the

al., 1990).

in

3.3.2 Associated 1

from where

areas

permanent spring and

charged

extrac-

connections between

remote

the situation

the range of

water

chemical properties suggest

1992. While the

1,

springs

The dissimilar .

hydrological complexity

illustrated ber

et

_1

scale

large

to

differing

and the

May 1987,

Jelsevnik

1

x

ra-

in Doblicica in the

and

PO

mg

springs

perhaps

mg~'

x

saturation. In

Doblicica

from

ranged

One black

m.

below

(Jamnice),

spring,

and

complex these

water

with Ca

content

near

0.01

to

black Proteus

active "boil-

temporarily

reach of 150

a

holes

caught

a

some

which

experiment,

two

photographed

boiling

was

are

permanent limnocrene spring

a

all within

was

from

rising

amphibians

no

associated groups of

ing holes",

in

O-phosphates

in the outflow (not in the

pumping

a

water

1986).

Jelsevnik,

series

level,

and its outflow. One

pool

caught

was

spring by diving,

two

sea

was

1

A

-

,

tios fluctuating. Oxygen of 1986

l"

x

water

flow

diameter. There

m

in the

after

spring)

lowered the

al.,

permanent spring,

a

deep pits

m

250 mg CaC0 3

to

tion and their

is

Arntzen

The total hardness of the

ous). 200

J. W.

set

meters.

Doblicica

main

subterranean

are

km

plains (Habic,

small rivers

some

2.5

of the Pol-

above

m

in karst

out

runs

surface while others several

karst

with deeply

altitude of 150

an

the conical karst

5-11

another (about

one

higher conical

Gora mountainridge,

janska

et

Sket &

B.

a

that

attack

violent

has been

Bijdragen

Table

V.

parkelj :

de

tot

drifted

Fauna

Jelševnik

J,

Na

hole); N2,

such as

mine,

from the

(I)

habitat

1994

-

of Proteus Na

Jezero; Ja, Jamnice; N1,

Trati

from sediments.

64

Dierkunde,

(small holes). of which

Taxa

Gastropods

the

origin

Nematoda, Oligochaeta,

47

anguinus Trati

collected

were

4 Discussion

(big 4.1

Taxonomic

and

Cyclopoida,

are

details

(for

see

Proteus

ex-

The black cluded

the black

of

status

is difficult to deter-

of Proteus

morph

text).

be

can

distinguished

morph by: (1) synthesis of pigment

from the white cd

Jelsevnik

light, (2) fully developed

in the absence of

>o

3 Q

Ja

NI

teeth, (4)

1847)

Belgrandiella sp. cf.

michleri

x

x

x

x

x

x

x

x

1932

Kuscer,

short

(6)

relative

x

cf.

Iglica

sp.

x

sp.

hauffeni (Brusina,

x

x

x

tion

cavernosa

Sadleriana

1978

Radoraan,

length

x

x

of the

x

sp.

of these

the

and

black

x

x

tion of separate

species

the

x

velkovrhi

Sket,

1960

x

sp.

Niphargus

sp.

x

Niphargus

sp.

x

x

shown

1960

in x

Petkovski,

1978

x

x

x

nature

So

impossible.

only

far

been documented

conditions. The observed

allopatric of

Z)

Triturus

1836

dis-

genetic

0.24 between the black

=

Proteus

population

from Sticna is similar

to

cristatus

are

that

superspecies, isolated

genetically

from

to

large

a

one

another

x

(larvae)

(Wallis

& Arntzen,

According

x

the species

black

x

to

1989; Arntzen,

the results

morph

populations

Chilopoda

of the

in

of Sticna than

to

the

prep.).

allozyme

study,

closer

is phylogenetically

population

the

to

of Pla-

x

nina. In view of them

Diplopoda

x

simi-

being morphologically

x

lar,

Diplura

the

large genetic

distance between Planina and

x

Sticna

populations

populations would fail

Reproduction.

embryonic

a.

almost

to test

x

x

P.

crossing experiments

x

fossarum Koch,

organised

been

1982).

the distances found for the various forms within the

sp.

vitellus and

in Proteus

have in

morphs

extent

an

Anoptichthys jor-

Nei

Cyclostomata (larva)

an

relative

interfertile (Culver,

will be

interfertility

tance

x

Cyclopoida

3.3.3

Examples

Astyanax fasciatus

x

(larva)

aquatic species

Terrestrial

genera.

differ-

recogni-

1936, that have recently

completely

and the white

Pisces

even

fish

the

to

x

parvulus (Sket, 1960)

Gammarus

have led

pose the greatest of technical difficulties and

both

Troglodiaptomussketi

be

to

Unfortunately,

x

for

Niphargus subtypicus Sket,

Hydra

occasions

many

or

cave

x

x

Niphargus

Surface

deserve attribu-

Strouhal,

spp.

1928

sp.

On

characinid

dani Hubbs & Innes,

Troglocaris

short

differ-

constant

morph

magnitude

Cuvier, 1819 and its

Proasellus

long trunk,

x

sp.

racovitzai

Monolistra

a

num-

tail, and (7)

evidently

white

taxa.

separate

of similar

include

Monolistra

high

legs.

the basis

to

ences

x

Crustacea Microcharon

of the

length

x

x

Sadleriana

a

x

ences

1885)

relative

from

shape stemming

musculature, (5)

x

On

Iglica

head

ber of trunk vertebrae and its associated

Belgrandiellafontinalis ( Schmidt,

Hauffenia

different head

a

well-developed

Gastropoda

Hauffenia

shape

(3)

eyes,

cranial bones and the number of

some

N2

stygophilic species

and

Stygobiont

the J

of

larva

-

probably

parkelj,

The

of

not

way) speaks as

guinus (Briegleb,

is

likely

finding

stage able

21 to

at

(still

be the

swollen

move

in favour of to

Jelsevnik of

case

by

around in

oviparity in P.

1962; Sket & Velkovrh,

a.

in

an-

1978).

we

would

that

to

indicate

identify

species separate priority

remarkable.

Rupnica

to

the

population

a

of

study

intermediate

pattern of clinal variation,

a

of Sticna

populations anguinus.

Proteus zoisii

is the type

logically indistinguishable lar

If

geographically

from P.

would go

of which

is

are

(Fitzinger,

locality.

and

as

a

Nomenclatorial

The

1850)

morpho-

biochemically

from Vir should be attributed

simito

the

48/I

B.

Sket &

Plate I.

a.

from left to the

J9, J8, J3, R1, live

(photo:

of P. tos:

a.

A.

and P5

specimen

Jelševnik in tail

Arntzen

-

A

Top: X-ray photographs of P.

anguinus,

left:

J. W.

dorsal B.

parkelj

Table

and P.

numbers

II; photo:

V.

a.

anguinus parkelj

and bottom: anterior

anguinus

are

Tabor); from

the insert shows the head in

middle

and P.

parkelj

right specimen

of Proteus

view,

Sket);

Hodalic).

(see

a.

black Proteus

in lateral view

de-

parts

(pho-

Bijdragen

of

name

Fitzinger,

P. zoisii

its

If such

black the

as

tail

a

morph

subspecies

P.

such

skeleton,

fol-

attributed

of

An

a

less

to

osteo-

equally

found in the popu-

as

(Fig. 6) might speak

taxonomie separation

a

would be

be

shape

given

schreibersii

parkelj.

z.

49

it has been

would

lations of Jelsevnik and Vir favour of

1994

-

revision

in the

logical synapomorphy segmented

(1)

Hypochthon

own:

1850).

the

lowed,

64

(although previously

taxon

same

a

de Dierkunde,

tot

of

the

in

of

of £>

with this

deserve

while the

of the black

ness

finding

of

white Proteus would absence of

firm

netic data we in favour of

a

the black A

dis-

genetic

distinctive-

in its

favour.

the

The

black and

sympatric

help proving

phylogenetic,

morph

priori incompatible

morphological

isolated

would

point.

In the

and ge-

ecological,

yet refute such taxonomie revisions

as

conservatively recognising

rather than

cies

not

morph speaks

genetically

Proteus

status.

Nei

view,

not

or

specific

0.24 is

=

of

species

two

whether

question

would also tance

My.

ago,

show

lineage

two

or

three

single

a

spe-

species.

and

Biogeographic

It is

the

present

but little doubt exists their

hypogean The

1990).

Proteus

genetic

Planina,

the Bela

and

Danube,

about the mutual isolation of

(cf. Gams, 1965; Novak,

data suggest

that

the

western

from Planina has become isoSticna and Jelsevnik

popula-

tions earlier than the Sticna and Jelsevnik

popula-

tions with In

eastern

regard

to

as

one

unit of Z) Nei

lineage separation (Maxson the

genus of

aquatic

reflecting

& Maxson,

has

14 My.

1979).

salamanders Triturus

been shown that this calibration fits with

dently basis

derived estimates of

a

consistent

put forward,

biogeography bian taxa

which

of other

this calibration dard

error

to

to

the

amphibian &

has

on

that

scenario could be

corresponds

(Oosterbroek

it

of

For

indepen-

speciation times;

biogeographic

been

to

the vicariance

and

non-amphi-

Arntzen, 1992). Applying

Proteus, while keeping the estimate of

genetic

stan-

distance in

for

Postojna Some

flew

some

9-5

line of argu-

populations

4.5-1.1

This

My.

the lower Pleis-

to

the

isopod

cavernicolus

There the

1951,

is,

same

however,

Jenko,

1959).

the distribu-

Asellus

in favour of

1932) speak

between

and the Gulf of Trieste

region

the

aquaticus

1925 and Proasellus istria-

connection

hydrographie

1952;

crustaceans

and

region

situation around

data, including

Racovitza,

(Stammer,

Sticna

approximately

today.

Melik,

(cf.

Miocene-

prekarstic

of the

in

do

they

as

Zoogeographie

tion of

nus

that the

streams

controversy concerning

some

an

the

(Sket,

ancient

Postojna

in

At

prep.).

the end of the Miocene, southern Slovenia became active which resulted in the presence of

and in

changes

lakes

(Prelogovic

Pliocene,

with

morphological

of

example

a

the

paleohydrology,

prekarstic

situation

relationships

(cf. Sket,

mental

karstification, to

conditions,

rather than

the black

the

a

hypogean

new,

the

not

through

is

onto

et

seen

Slovene

al.,

1986;

V.

or

hypogean

before

Pohar,

asso-

to a

that 1

pers.

rela-

habitat.

glaciations

document

territory

The

traits may be

several

to

such

question.

(cf. Sket, 1985)

available

evo-

adaptations.

undergone

open

colonization of the

evidence

Bowen

an

non-troglomorphic

The Pleistocene has

extended

has

remains

ciated to weak selection recent

habitat

differentiation

phenotypical

Proteus

change

retention of

(cf.

to

to

Proteus. Governed by environ-

have become obscured

locally

adaptive

no

relate

1970, 1986; Sket &

lutionary convergent, troglomorphic

tively

an-

dis-

the

1982).

With the

Why

be

to

in which

species

cave

phylogenetic

may

differentiation de-

between them. Proteus appears

present-day

a

in surface rivers and lakes before the

taxa

tribution and

Bole,

al.,

et

in the direction of the flow of

We suggest that Proteus may have formed

veloping other

with

depressions

became available

the "molecular clock"

amphibians

calibrated

each other.

same

Jelsevnik

independence

Krajina

direction

rivers.

Sticna,

the Sava and the

habitats

population

lated from the

of

regions

the

Sticna and

place

the upper Miocene

to

Following

generally agreed

series of

Jelsevnik drain towards

lineages

corresponds

first documented

took

tocene.

tectonically

evolutionary history

the

coincides with the upper Pliocene

1975)

At

the

mentation,

that

mean

lower Pliocene.

or

large

4.2

which

Pliocene surface

Recognition raise the

would

of Proteus

separation

Planina

population.

this

mind,

but

these

My.

ago

comm.).

B.

50

Intensive

karstification,

underground, With

changes. Proteus have

the

isolation

In

caves.

less,

or

Triturus

the

proceeding,

from Sticna and Jelsevnik may

isolated from

in

necessarily

streams

hydrographical

karstification

populations

become

surface

moving complex

caused

to

time span similar

a

within

stocks

some

into

developed

but

another,

one

the

distinct

clearly

Oosterbroek &

(Rafinski& Arntzen, 1987;

not

their

genus

below

the presence of

Proteus,

tiated eyes

seems

(1912),

the basis of

on

unique

P.

repeated

eyes and

develop

may

al observations of

ability

(Ehrenberg, conducted

Proteus,

on

that

pigmentation.

Occasion-

Proteus do suggest that

have

other

any

or

of it

only part a

white

P.

As in Proteus

thesis

found in

seem

parkelj, „,

duck-bill tribute

fishes be

typical

as

well

as

shape

and

seems

the

in

&

to

lost the

a.

to

a.

traits.

anguinus

1974),

of P.a.

has

compared

parkelj.

The

be associated with

of the

and

the

the present,

to

major

and

threats,

the

freely

skull

as

in

by Culver,

legal protection

Jelsevnik and DobliCica

only

localities where P.

with

certainty.

parkelj

a.

on

the

of

finding

(A. Hudoklin,

a

black Proteus near Mala

comm.),

pers.

located 7

southeast of Doblicica and within the has

karst, been

from

reported

south of

caves

the

of

specimens

ticular

the

by

town's

near

inhabitants

Vanin

the

it

can

plesioin head

narrowing

of

mandibular arch

head musculature.

A

in

been

now

In

recent

12 km

to

Fig.

on

the

in

(cf.

no

voucher

par-

in southfor

adopted

the

white

only

years

ac-

(formerly

Vas

of blacks?

Local

of white Pro-

finding

have become

specimens

does exist

hypogean

waters

Gora

its

of P.

springs

not

between Jelsevnik and Doblicica.

the northeast of

Poljanska

nels

is

be confirmed.

to

for

Jelsevnik

a

locality

at

(Klepec, 1981;

1).

If the

tence

only

"black"

of

data. The

Slovenska

populations

specimen

voucher

cf.

sampling

Kocevje (Gottschee)

reported

locality

a

in

has

White Proteus

in

over

have been obtained from here.

specimens

teus

morphological

supply.

water

all

of whites? The

occurrence

(1882) unfortunately

any

Slovenia

eastern

(F.

few active

speleobiological

populations

spring

karst

in any of the

or

natural

Graf

by

companied

the

unfavourable.

are

Black Proteus in mention

to

of low

region

Crnomelj. Unfortunately, Gora the

Poljanska

conditions

Lahinja

km

south of Doblicica

springs

Velkovrh, pers. comm.)

Unfortunately

(cf.

finding

been documented. No Proteus have

not

available and the reports will have

at-

the

on

the

found

of Proteus in the main spring of Jelsevnik. A report

the

an

are

has been

No reliable data exist

are

change a

carry.

cited

1944;

eyes,

occurring

to

affected

not

Since

head is

vertebrates

they

an

The

1982).

while

ability (see

plethodontid amphibians,

of P.

weakening

exposed

well-developed

Kühne,

neurocranial bones a

not

pigment-

pigmentation

troglobiont

apomorphic

morphic shape

the

unlike that

cavernicolous fish

The

of the P.

of

normally

plesiomorphic

Cooper

regarded

(Ehrenberg,

not

of Proteus is

completely

408).

with be

to

shape

1965;

some

have

to

along

considered

while

the atavism of melanin syn-

,

e.g. Vandel, 1965:

del,

skin

Windischdorf)

are

daylight.

others

in

the whole

al., 1986),

snout

perhaps

-

population

although specimens

is

et

cover

teeth

Astyanax fasciatus (Breder,

The

amphibian.

may

the

out

parkelj.

a.

The Jelsevnik ed

1878)

thoroughly

been carried

cave

on

but

1878),

(AljanCiC

triangle

1867 in: Knauer, found in

is retained indeed

not

newly developed pigmentation

retaining

1973),

(Durand,

pigment synthesis

in

far have

blind Pro-

1867 in: Knauer,

nor

so

light,

captive

experiments

Kammerer

parkelj.

a.

experiments

been

successfully

to

differen-

externally

claimed that under the influenceof

body

the

4.4 Distribution,

Within

the

number of

diminutionof the eye with the small eye bulb

Up

teus

the

may be associated with

snout

of the mandibular bones and

slight elongation

increase in

black Proteus

species

Character evolution

not

the

A

-

Arntzen,

1992).

4.3

J. W. Arntzen

elongation of

The a

Sket &

a.

the that

Habic

are

parkelj

of the

defined

may

et

al.,

1990).

of

As

karst

habitat,

be limited

hydrographical cover an area

high

of the

the exis-

to water

background

of

approximately has been

chanboth

55 km

2

suggested by

Bijdragen

P.

de Dierkunde, 64

tot

Habic

the

comm.),

(pers.

parkelj of

Bela

while

Krajina

1994

to

the

51

of P.

occurrence

may in fact be restricted

the

-

(1)

a.

the shallow karst

out

of

karst

deep

Kranjec (Koper) performed

with

the

mountains is

surrounding

inhabited

The

available

data do

contradict this

not

Indeed, the hypothesis

hypothesis.

data about the

(unconfirmed)

is

in line

sporadic

with

occurrence

of white Proteus in Jelsevnik and black Proteus in

Copepoda. A.

we

of

springs

Because

Lahinja.

wijk

(Amsterdam) provided

M. Garcia-Paris

thanks

are

hamlet

of

Proteus

does

The

their

find,

to

provide

not

from the

water

Jelsevnik is tributed doline

of

dump the

uphill

Na

absence

in any

Trati

area

holes

boiling

smelting plant

a

Poljanska

ing

debris

sandy

in

less

Gora,

grained

dumped

in

did

in

due to Silvo

captivity in

kept

close

tion The

serious threat

a

holes

boiling

similar but

of

rich

these

sands

pollution

Since its habitat is a

in

As

it appears

bution

be

to

P.

area,

taxon

Slovenian Red List

of

accessible be

with

has been

parkelj

a.

a.

pollu-

parkelj.

hydrologically

are

easily

cannot

a rare

Jame,

(Sket,

1992).

Slovenian law

a member

as

as

a

a

to

small distri-

placed

on

the

species

dinance

on

protection

of

of

protection

as

well

anguinus.

for any purpose.

collecting

The

Biol.

most

or

and

Gregori, a

new

their

1980)

act

the old

updating

ani-

Ministry

in its

one

is in

1986.

Crni moceril

iz Bele

12:

Vest.,

J.W. &G.P.

F.

to the

anguinus Laur.)

chez

J.R.

J.,

1972.

sophila

J.L. & F.

Naturk.

(Alfred

A.

The 'Wolterstorff Index'

in

Genetics,

American New

C.A.

Tracey,

70:

G.M.

super-

anguinus

superspe-

Vorgesch. Magdeburg, Special

variation

Knopf,

D.Q.,

de

Proteus

Symposium.

M.L.

in

the

Mouräo &

Drosophila

natural

Pérez-

willistoni Dro-

of

populations

113-139.

1979. The Audubon

Wayne King,

North

Protée

press.

Enzyme variability

willistoni.

to

le

superregener-

[Cas

taxonomy of the Crested Newt

Powell,

IV. Genie

guide

Nase

109-113.

Wallis, in

Mus.

group,

Behler,

Carniola],

and

reptiles

Society

field

1-743

amphibians:

York).

Richmond,

D.S.

V.

Fullerton,

Sibrava,

the

"perma-

nary

own

D.Q. tions

"Or-

glaciations

in

1986.

Velichko,

the northern

Bowen & G.M.

in the northern

Correlation

hemisphere.

Richmond

(eds.),

of

In: V.

Quater-

Sibrava,

Quaternary glacia-

hemisphere: 509-510 (Pergamon Press,

Oxford). S. &

J.

1980. Redke

Gregori,

in

ogrozene

zivalske

vrste

animal

endangered

(Brelih

&

in 1976 and

operational

accidentelle

R.J. Fulton & A.A.

recent

developmental stages"

has become

(Proteus

Wolterstorff Memorial

Issue

Ayala,

discourage

as

We

Mihevc,

1964. Primer akcidentaine

Sket,

mocerilu

cies. Abh. Ber.

v

species

the Slovenian

by

39-44.

and its value

Brelih, rare

conscientiously collecting

funded

Proteus from the White

black

régéneration

man,

performed.

of the category

animals", of P.

subspecies

was

P. Habic & A.

M. & B.

acije pri

Salas,

animal

endangered

under strict

cavernicolous

nently right

is

for

of the

about the local

Technology.

[The

28:

Aljancic,

Bowen, It

study

(T.

been noticed there.

not

directly

not

estimate

population

We consider the

Jamnice

has

and

Aljancic, M.,

day period

a ten

the survival of P.

to

family, inhabitants

and his

promptly informing us and

material.

outgroup

manuscript. Special

a

damag-

doline is

with

contact

comm.).

pers.

the

read the

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Medic

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