Annls Limnol
20 (1-2) 1984 : 129-139
Marthamea
beraudi (Navâs) and its European congeners
(Plecoptera : Perlidae)
P. Z w i c k '
The west palearctic species of Marthamea are reviewed. Adults are distinct from other European genera, but larvae
and eggs are very similar to those of genus Phasganophora. M. beraudi (Navâs) (Antilebanon Mts.) is re-instated as a species distinct from M. vitripennis (Burmeister). A lectotvpe of the tatter is designated. Its range includes south-west, central and south-east Europe. Records from Syria are unconfirmed, those from Israel are erroneous. The species has in the
past been confused with larvae of a Phasganophora species. M. selysii (Pictet) has brachypterous males and is known from
Spain north to Meuse and Moselle rivers. Keys to adults and larvae are presented, but larval M. selysii and M. vitripennis
can presently not be separated. The European Marthamea species are inhabitants of the potamon and are endangered ;
they are presently already extinct in parts of their former ranges.
Marthamea beraudi (Navâs) et ses congénères européens (Plecoptera : Perlidae).
L'auteur révise les espèces paléarctiques occidentales de Marthamea. Les adultes sont distincts de ceux des autres genres européens de Perlidae mais les larves et les œufs ressemblent beaucoup à ceux du genre Phasganophora. M. beraudi
(Navâs), de l'Anti-Liban, est une bonne espèce distincte de M. vitripennis (Burmeister). Un lectotype de cette dernière est
désigné. Son aire de répartition s'étend sur le sud-ouest, le centre et le sud-est de l'Europe. Sa présence en Syrie n'a pas
été confirmée et, pour Israël, il s'agissait d'une erreur de détermination. A l'état larvaire, elle a été confondue auparavant
avec une espèce de Phasganophora. Les mâles sont brachyptères chez M. selysii (Pictet), qui a été récoltée de l'Espagne
â la Meuse et la Moselle. L'auteur présente une clef pour les adultes et les larves mais, pour ces dernières, on ne peut
distinguer, à l'heure actuelle, M. selysii de M. vitripennis. Les espèces européennes de Marthamea font partie du potamon
et sont menacées. Elles ont déjà disparu dans certaines parties de leurs aires de répartition originelles.
In 1909, L. N a v â s c r e a t e d the subgenus Lerpa
(apparently o f Perla, although this w a s not clearly
stated) for a n e w species, beraudi, that he had received f r o m P è r e B é r a u d . The actual c o l l e c t i o n site of
the n e w species w a s not indicated but a c c o r d i n g to
the title of the paper the s p e c i m e n s c a m e from
« Siria ( B e y r o u t h ) » . B e i r u t w a s a Turkish p r o v i n c e
(vilajet) consisting of the c i t y of Beirut and the area
west of the J o r d a n valley to south of N a b l u s (E.
Debes Handatlas, 1895).
N a v â s ' figure shows m a l e genitalia not clearly different f r o m Perla (Marthamea) selysii (Pictet) and
vitripennis
( B u r m e i s t e r ) as shown by K l a p â l e k
1. Limnologische Flussstation des MPI fur Limnologie, Poslfach
260, D - 6407 Schlilz, Fédérai Republic of Germany.
(1907). N a v â s indeed said beraudi was s i m i l a r to
these t w o species without indicating expressively in
w h a t it differed, as w a s his habit. Klapâlek (1914)
c o n s i d e r e d Marthamea
g e n e r i c a l l y distinct f r o m
Perla. A t y p e species w a s not designated until 1923,
w h e n K l a p â l e k chose M. vitripennis
and i n c l u d e d
Lerpa N a v â s as a s y n o n y m of Marthamea K l a p â l e k .
In the s a m e paper, he a l s o translated an e x t r a c t of
the o r i g i n a l Latin d e s c r i p t i o n of beraudi i n t o German. H o w e v e r , the section o n m a l e genitalia w a s
replaced by what is c l e a r l y Klapâlek's interpretation of N a v â s ' figure : that they w e r e shaped exactly
as in M. vitripennis.
Klapâlek believed
vitripennis
and beraudi to be the same species but he d i d n o t
establish the synonymy f o r m a l l y . Navâs{1929) insisted on the specific distinctness of beraudi ( m e n t i o ning l a r g e r size, b r o a d e r w i n g s and, as a consequence o f both, richer v e n a t i o n ) but agreed w i t h the
transfer to Marthamea. As Marthamea beraudi t h e
Article available at http://www.limnology-journal.org
or http://dx.doi.org/10.1051/limn/1984005
130
P. ZWICK
(2)
s p e c i e s w a s listed b y Claassen (1940). l i l i e s (1955)
first d e s c r i b e d the distinctive penes of M. selysii and
M. vitripennis and indicated that he had seen a m a l e
of the latter from Syria. In 1966, he mentioned Israel i a n m a t e r i a l he had seen and a c c e p t e d the
s y n o n y m y between beraudi and vitripennis (that Klap â l e k had o n l y tentatively suggested) as d e f i n i t e l y
established.
Berlin, ZMB), Dr. B. Hauser (Muséum d'Histoire Naturelle,
Genève), Dr. A. Kaltenbach (Naturhistorisehes Museum
Wien, NMW), Dr. W. Kittel (University of Lodz), Prof. J.
Kugler (Tel Aviv University) and Dr. H.J. Schnur (Hebrew
(Unhersitv of Jerusalem), Dr. K. Kumanski and Dr. B. Russev (Zoological Museum Sofia, ZMS), Prof. Dr. G. Morge
(former Deutsches Entomologisches Institut, DEI, Eberswalde), and Dr. P. Ward (British Museum, Natural History,
London, BMNH).
D u r i n g study of s o m e P l e c o p t e r a f r o m the N e a r
East, I have seen a small number o f Marthamea spec i m e n s f r o m the h e a d w a t e r s of the Litani and Jordan rivers in the Antilebanon M t s . w h i c h are neither
selvsii n o r vitripennis. Although the t y p e of beraudi
is not a v a i l a b l e ( A u b e r t 1956), I have no hesitation
to assign these specimens to that species, which w a s
d e s c r i b e d f r o m the s a m e area. M. beraudi is reinstated as a distinct species and is c o m p a r e d to the
o t h e r t w o west palearctic species of
Marthamea
w h i c h are also r e d e s c r i b e d . N o n e o f the Asian spec i e s assigned to Marthamea is c o n s i d e r e d . W h e t h e r
a n y of them really b e l o n g to this genus is very quest i o n a b l e ( l i l i e s 1966). At least M. producta K l a p â l e k
has in the m e a n t i m e been transfered to genus Phasganophora
by R a u s e r (1968).
Genus Marthamea Klapâlek
Only a small fraction of the material used in this study
is in my own or the late J. lilies' collection at the Limnologische Flussstation Schlitz (LFS). Most was borrowed from
several collections. Grateful thanks for help or information are extended to Dr. N. Alouf (Lebanese University, Beirut), Dr. E.G. Burmeister (Zoologische Staatssammlung,
Munchen ; ZSM), Dr. A. Comte (Instituto Espahol de Entomologia, Madrid, IEE), Dr. J. Dlabola (National Museum,
Prague, NMP), Dr. M. Gonzalez del Tânago and Dr. D. Garcia de Jalon (Escuela Tecnica Superior de Ingenieros de
Montes, Madrid), Dr. K.K. Gunther (Zoologisches Museum
1907 Perla (Marthamea) Klapâlek, Ceskâ Akad. Cis. Fr. Jos.
I. (2) 16 (2) : 17. Type species (design. Klapâlek 1923) ;
Perla vitripennis Burmeister.
1909 Per/afZ.erpa;Navâs,Brotéria,Ser.Zool.8: 102.Type
species (by monotypy) : Lerpa beraudi Navâs.
A m o n g the genera of P e r l i d a e in the w e s t e r n
palearctic region, adult members of Marthamea can
be recognised by the f o l l o w i n g combination of characters : W i n g s clear, veins b r o w n except in costal
space w h e r e veins and membrane are y e l l o w i s h . N o
crossveins connecting anal veins in hindwing. M a l e
abdominal tergite 5 w i t h posterior extension ; centre o f tergites 6-8 variously adorned w i t h patches
of small spinules ; tergite 9 simple ; hemitergites 10
with d e e p anterior notch separating a l o w e r fingershaped process from the massive main part a b o v e
and behind it ; penis base transversely annulate,
faintly sclerotised ; eversible penis sac large to very
large, w i t h many fine spinules. Sternite 6 with setal
brush, a smaller one also on sternite 7. Females
without well projecting p r o p e r subgenital plate,
edge of sternite 8 m o r e or less sclerotised. T e r g i t e
10 rather long, m i d d l e almost tongue shaped.
Fig. 1. Head pattern of adult Marthamea ssp. ; a, M. vitripennis, cr from Kula Ljums ; b, M. selysii, left cr, right
9 paralectotypes of var. mosellae, from Alf ; c, M. beraudi, cr from Nahal Dan. Scale is 2 mm (two times 1
mm in b).
M A R T H A M E A BERAUDI A N D ITS E U R O P E A N C O N G E N E R S
(3)
Larvae.
Submentum simple. Occipital setal fringe clearly
d i v i d e d into different sections behind the eyes : the
long median section is f o r m e d by short club-shaped
setae standing in a v e r y regular line. At the sides,
it curves a little a n t e r i o r a d and meets the i r r e g u l a r
postocular fringe o f long setae at a distinct a n g l e .
At this meeting point, there is a single erect seta. N o
so-called s w i m m i n g hairs on cerci ; apical setae o f
distal cercus s e g m e n t s normal, short. A r m a t u r e o f
proventriculus consists of numerous small spinules,
no major sclerites.
131
Eggs.
Chorion c o m p l e t e l y smooth. T h e anchor has the
shape of a huge b i c o n c a v e disc, its short n a r r o w
stalk inserts on a small sclerotised nipple o f the
chorion.
Note that larvae and eggs of Marthamea are similar to
those of genus Phasganophora, of which several west
palearctic species exist. The only known European Phas
ganophora larva has in the past been confused with M. vitri
pennis, see there.
F o r a long time, species identification in Martha
mea relied e x c l u s i v e l y on colouration. C h a r a c t e r s
Fig. 2. Pattern on head and abdominal tergite 4 of larval Marthamea ssp. ; a, M. vitripennis from the Maritza ;
b, M. beraudi from Banyas ; c-e, M. selysii from Moselle, R. Lozoya and R. Jarama, respectively. Figures are
not to scale.
132
P. ZWICK
of penes d e s c r i b e d by lilies (1955) appear to have not
b e e n used b y subsequent authors. W h i l e structural
i d e n t i f i c a t i o n of m a l e s poses no p r o b l e m s , f e m a l e s
a r e best i d e n t i f i e d f r o m e g g s they contain. W h e n
eggs are not available, the head pattern must be used
as an auxiliary character. F e w larvae only have been
a v a i l a b l e . P r e s e n t l y , o n l y larval M. beraudi can be
identified with certainty.
Only distinctive specific characters are described below.
In the description of penes, the terms ventral and dorsal
apply to the position in resting condition, inside the ani
mal. The ventral side can be recognised by two small poorly
expressed brownish basal sclerites. Figures are so orien
tated that these are on the lower side.
Marthamea vitripennis (Burmeister)
1839 Perla vitripennis Burmeister, Handb. Entomol. 2 :
880.
1839 Perla bicolor Burmeister, Handb. Entomol. 2 : 880.
1852 Perla terminalis Walker, Cat. Neur. Ins. Brit. Mus.
1 : 155.
Material examined : 9 lectotype of P. vitripennis (here
designated): 261 \ivitripennis Burm. Pict.*/Halle, Erichson/'Marthamea vitripennis Burmeister : Pictet (Mus. Ber
lin). 9 lectotype, 9 paralectotype of Perla terminalis Wal
ker (des. Kimmins 1969) : East Indies/49. Perla terminalis
(BMNH). Spain : 1 penis on slide, Spain (coll. lilies) ; 1 9
with eggs, Sobradia (Zaragoza), 5. VI 1933, (DEI ; Navâs
del. vitripennis). France : I penis on slide, Toulouse, Aubert
(coll. lilies). Germany : 1 penis on slide, Main (coll. lilies) ;
1 cr, Dresden (Schiller ; coll. Klapâlek. NMP). Czechoslo
vakia : about 30 specimens including larval exuviae, Celakovice, May 1915, and Stvanice (coll. Klapâlek, N M P ) ; 2
era-,] 9 , Slowakei, Dimburg ( = Suchohrad) 5. VI. 1906
(F. Werner, N M W ) . Yugoslavia : 1 9 , Petrina planina (E of
Ohrid), 1 564 m (J. Thurner ; LFS) ; 2 o*cr, 1 9 , Uskub ( =
Skopje), 5. and 12. V I . 1918 ; 2 crcr, Orman, 14. VI. 1919 ;
1 o* without abdomen, Maced. centr. meridion., Drenovo
bei Kavadar, 200-800 m, 1.-9. VII. 1959 (F. Daniel) (all ZSM).
Albania : 3 crcr, Kula Ljums, 7.-14. VI. 1918 (at confluence
of Luma and White Drim, see Friese and Konigsmann 1962)
( N M W ) . Poland : 1 ç D e i m e (near Gdansk), 1.-15. I X . 1915
(W. Horn ; Ulmer del. vitripennis ; DEI). Bulgaria : 1 o \
Kresnensko Defile (Kressna Gorge of Struma r.), V I . 1935
(Dr. Taborsky, LFS) ; 1 9 , 4 . VII. 1932, 2 crcr, 27. VI. 1935,
2 crcr, 27. V I . 1935, Kresna-Defilé, Sali Aga (Tuleschkow ;
Rauser det. vitripennis) ; 1 9 , Bulgaria (Ananian ; all Mus.
Sofia) ; 1 larva, Maritza river at village Ljubimez, 29. V .
1966 (from B. Russev, Sofia). Several additional males and
females without adequate locality labels from the
museums in London, Geneva and Vienna have also been
examined.
B o t h sexes m a c r o p t e r o u s , w i n g s 12.5-21.0 m m
long. B o d y b r o w n w i t h yellow marks. Head pattern
as in Fig. l a , n o t e c l e a r y e l l o w p i g m e n t l a t e r a l l y
f r o m o c e l l i in front o f o c c i p i t a l suture. In f e m a l e s ,
(4)
y e l l o w is m o r e extended and the pattern less contrastive. R u g a e of p r o n o t u m , soft parts of thorax,
prescuta, tibiae (except n e a r base), l o w e r side of
femora, abdomen, and base of cerci yellowish to yel
lowish brown.
Male.
A distinctly bilobed spinulose extension of t e r g i t e
5 covers the membraneous anterior half of t e r g i t e
6 w h i c h has a well d e v e l o p e d entire antecosta. Ter
gite 7 s i m i l a r to 6 except a semicircular m e d i a n
b r o w n m a r k which carries many small spinules in
its centre. Antecosta of tergite 8 divided, triangular
median b r o w n mark with very few spinules. Depres
sed c e n t r e of tergite 9 sclerotised, r h o m b o i d , sim
ple. In side v i e w , p o s t e r i o r contour of h e m i t e r g i t e
10 strongly, almost rectangularly, bent ( F i g . 3).
Penis relatively small, w h e n e v e r t e d s h o r t e r than
w i d t h of a b d o m e n and shorter than last three tergites t o g e t h e r . Short ventral face of e v e r t e d sac
straight, longer dorsal side convex. A m o n g the
n u m e r o u s small spinules there is a g r o u p of slen
der spicules dorsally, at the highest point of the ever
ted sac ; l a r g e r triangular spinules f o r m a distinct
o b l i q u e subterminal ring.
Female.
Subgenital plate not distinctive. Dark m a r k s on
head less e x t e n d e d than in male, especially o c c i p u t
light. P a l e in front of o c c i p i t a l suture b e t w e e n ten
torial callus and p o s t e r i o r ocellus, dark b e t w e e n
p o s t e r i o r ocelli.
Eggs.
Ca. 0.38 m m long, greatest diameter near m i d d l e ,
oval. A n c h o r pole s o m e w h a t flattened, w i t h very
small apical nipple for anchor attachment (Fig. 6a).
La rva,
P a r a p r o c t s w i t h profusely branched gills. H e a d
pattern and abdominal m a r k i n g s as shown in Fig.
2a, s i m i l a r to M. selysii ; distinction is presently not
possible. Previous descriptions of p r e s u m e d larvae
of M. vitripennis w e r e based on misidentified mate
rial, see N o t e s b e l o w .
Notes.
T h e status of the vitripennis syntype is c o n f i r m e d
by o l d catalogue notes of the Museum in Berlin (Dr.
Gunther, in a letter). T h e specimen is without abdo
men, but the well expressed head pattern leaves no
doubt about its identity. R e d i s c o v e r y of this syntype
(5)
M A R T H A M E A B E R A U D I A N D ITS E U R O P E A N C O N G E N E R S
FIG.
MARTHANEA
10
Scale
FOR A , 0 , 5
IS
1 Tin
IN
SELYSII
HEMITERGITES
DORSAL
;
A,
A B D O NINAL
A N D LATERAL V I E W S ,
T I N FOR B , C
confirms the conspecificity of vitripennis
(females)
and bicolor (males ; lectotype designated by Z w i c k ,
1972). Pictet (1841) had seen syntypes of both but
was not sure of their identity. Schneider (1848) had
established the c o n s p e c i f i c i t y f r o m fresh m a t e r i a l
he had from Silesia, but o n l y f r o m the description,
without having seen syntypes.
For the requested suppression of an o l d e r unused
name by Fabricius for the present species and the
history of the use o f the name vitripennis see I C Z N
(1981). The female lecto- and p a r a l e c t o t y p e o f Perla
terminalis
W a l k e r (1852) have been c o m p a r e d .
S y n o n y m y w i t h M. vitripennis had been suggested
by K l a p â l e k (1923) and is here c o n f i r m e d . H o w e v e r ,
I doubt the correctness o f the locality label of the
terminalis-types
(East Indies). R e c o r d s f r o m the
N e a r East are u n c o n f i r m e d . T h e Syrian m a l e mentioned by lilies (1955) is not available. T h e Israelian
material r e c o r d e d by l i l i e s (1966) has been examined and is beraudi. T h e easternmost specimens of
vitripennis
I have seen are f r o m B u l g a r i a . I a m
; SCALE
TIP
OF
133
NALE
RESPECTIVELY
FOR D E T A I L S
; D,
I N D IS
:
B,
C,
EVERTED
0,02
.Tiale
PENIS.
tvïi.
c o n v i n c e d that records f r o m Romania ( K i s 1974),
Lithuania (Kazlauskas 1962), in fact most r e c o r d s
of adults from the m i d d l e and eastern part of central E u r o p e are c o r r e c t . This is also likely for
r e c o r d s from Paris (e.g., Pictet 1841). It is unfortunate that the accurate origin of a specimen f r o m the
M a i n river in c o l l . l i l i e s is not known. T h i s c o m e s
f r o m a suggested g a p b e t w e e n disjunct eastern and
w e s t e r n populations of M. vitripennis w h i c h w o u l d
have been separated by glacial events (lilies 1953).
H o w e v e r , in v i e w of the dramatic decline of the species practically e v e r y w h e r e in central E u r o p e I a m
w o n d e r i n g w h e t h e r the lack of records f r o m part o f
central E u r o p e may not be due to early p o l l u t i o n o f
the p o t a m o n o f rivers o f w h i c h M. vitripennis
was
an inhabitant. R e c o r d s f r o m France (e.g., Despax
1951) and Spain (Aubert 1957) require checking. M.
vitripennis occurs there but has at least s o m e t i m e s
been confused with M. selysii, see there.
S c h o e n e m u n d (1925) d e s c r i b e d larvae as those o f
M. vitripennis
w h i c h have a transverse d a r k b a n d
P. Z W I C K
134
across the head, in front of the M-Iine. An essentially
T-shaped p a t t e r n results, instead o f the Y-shaped
p a t t e r n present in true M. vitripennis. lilies (1955)
used s o m e of S c h o e n e m u n d ' s material from the
P l a n e f o r a s i m i l a r description. Although Schoenemund said the larva to be easy to keep in aquaria
(1925) and to be c o m m o n in the Plane (1927), he
a p p e a r s to h a v e n e v e r actually reared an adult. A
f e w s p e c i m e n s f r o m his c o l l e c t i o n s b e f o r e me
include m a t u r e f e m a l e larvae with eggs. T h e y have
been c o m p a r e d to adults and larvae (also including
e g g s ) of Phasganophora
senilis (Klapâlek) identified
and c o l l e c t e d by K i t t e l ( 1976) and o b t a i n e d through
his courtesy. T h e y are clearly the same species, and
not Marthamea vitripennis. A note on the true iden
tity of the species in question is under preparation.
M a r t h a m e a selysii (Pictet)
1841 Perla (Perla) selysii F.J. Pictet, Hist. nat. gén. part.
Inst. Névr. : 208, pl. 17, fig. 5.
1885 Perla selysii var. mosellae McLachlan, Ent. Mon. Mag.
(1885).
Fig.
3. M a r t h a m e a
vitripennis,
(6)
Material examined : Spain : 1 9, Perla selysii Env.
Madrid (specimen reported by E. Pictet 1865; Mus.
Geneva) ; 3 cr <y, Paracuellas, Jarama, VI. 1936 (D. Pelayez ;
Aubert det. 1954 : vitripennis) (coll. Inst. Espan. Entomol.,
Madrid); Talamanca, R. Jarama, I 9 , 28. V. 1983, 2 lar
vae, 5. V. 1983 ; Pied ras. R. Jarama, 3 larvae, 1. V. and 17.
VI. 1969 (all coll. G. del Tânago) ; 1 larva (mature 9 with
eggs), env. of Madrid, R. Lozoya, 1979 (coll. G. de Jalon) ;
1 9 , Andalusia (coll. Hiendlmayr, DEI). Belgium: 1 cr,
Dînant Loyon ; Belgium, coll. Camille van Voixem (coll.
McLachlan, BMNH). Germany : 1 O", 1 9 paralectotypes of
M. selysii var. mosellae, Alf, Moselle, 1894 (coll. McLach
lan, BMNH) ; 3 o-cr, 2 9 9 . 1 exuvia, Alf (le Roi, in coll.
lilies, Schlitz): 1 o-, 1 9 . 3 exuviae, 3 larvae, IX. 1958,
Mosel, km 21.8, km 37.8, km 65.7 (coll. lilies).
Female macropterous, w i n g s 18-21 m m long.
Males brachypterous, wings cover about half of the
abdomen and do not extend to the tips of the metafemora. Colour varies from more o r less uniform
dark b r o w n to yellowish b r o w n with a head pattern
as in Fig. l b , right side. In the last case, the dark
mark tends to be ill defined postero-laterally.
Male.
Spinulose extension of tergite 5 w i d e , shallowly
bilobed (Fig. 4). T e r g i t e 6 medially soft, except shi-
male g e n i t a l i a . Details and
s c a l e s a s in F i g . ^ .
M A R T H A M E A B E R A U D I A N D ITS E U R O P E A N C O N G E N E R S
(7)
ning longitudinal little folds in the anterior half.
S i m i l a r l o n g e r folds present anteriorly on tergite 7.
Behind these are t w o m e d i a l l y confluent patches of
spinules. Antecosta 8 d i v i d e d , vague indications of
longitudinal folds a n t e r i o r l y , a few spinules posteriorly on the soft c e n t r e of tergite. T e r g i t e 9 s i m i l a r
to M. vilripennis. In side v i e w , the posterior contour
line of h e m i t e r g i t e 10 f o r m s a regular, approximately s e m i c i r c u l a r line.
Penis large, w h e n everted as long as width of abdomen and as long as last four segments. E v e r t e d sac
long, w i d e , tubular, w i t h dorso-basal hump. A r m a ture consists of v e r y m a n y stout triangular spinules. V e n t r o m e d i a n spinules l a r g e r than others, but
no c l e a r pattern results f r o m these size differences.
Female.
Genitalia not distinctive (compare N o t e s !). In pale
specimens, the area b e t w e e n the ocelli is clearly darker than the rest, but this tends to be also infuscate,
instead of c l e a r y e l l o w , n o c o n t r a s t i v e pattern.
EggsCa. 0.38 m m long, s p h e r o i d , m o r e or less acuminate t o w a r d s small n i p p l e f o r m i n g a n c h o r attachment. Demarcation o r flattening of polar area hardly
recognisable. T h e c h o r i o n is distinctly thicker near
the greatest d i a m e t e r of the e g g than at the poles
(Fig. 6b, c).
Larva.
P a r a p r o c t s w i t h p r o f u s e l y b r a n c h e d gills. H e a d
pattern similar to M. vitripennis,
although sometimes m o r e extended. T h e abdominal pattern appears
to be quite variable, see Fig. 2 c-e. It should be noted
that the e x t e n s i v e l y p i g m e n t e d specimen from the
M o s e l l e is g e n e r a l l y q u i t e pale, perhaps faded in
alcohol. Distinction f r o m M. vitripennis
presently
not possible.
135
and ventral v i e w s presented in Despax (1951). l i l i e s
w a s right in treating the var. mosellae as identical
w i t h the nominate form.
M. selysii had been r e c o r d e d from the r i v e r s
Meuse (Dinant, Maastricht, Liège), Moselle (Alf, Wasserliesch, and other places) and the m i d d l e section
of the R h i n e (e.g., K l a p â l e k 1923, S c h œ n e m u n d
1925). R e c o r d s f r o m Spain by A . E . Pictet (1865) and
o t h e r s w e r e d o u b t e d by K l a p â l e k (1923), his v i e w
w a s shared b y lilies (1966). H o w e v e r , o c c u r e n c e in
Spain is n o w c o n f i r m e d . R e c o r d s f r o m H u n g a r y
(Pongracz 1914) are unconfirmed and p r o b a b l y erroneous. T h e R o m a n i a n m a l e illustrated by V a s i l i u
and Costea (1942) is c l e a r l y M. vitripennis. M. selysii a p p e a r s to be restricted to w e s t E u r o p e (Fig. 7).
T h e species is e x t r e m e l y endangered : s p e c i m e n s
f r o m the M o s e l l e listed above appear to b e the last
that have been taken in the northern part o f the spec i e s ' range. Spanish populations are a l s o s t r o n g l y
threatened by extinction ( M . Gonzalez del T â n a g o ,
in a letter).
l i l i e s (1953) b e l i e v e d that Ai. selysii w a s a t h e r m o philous inhabitant of Central European l o w l a n d
r i v e r s b e f o r e the Pleistocene and b e l o n g e d to the
« glaziale Mischfauna » of that area. N o r t h w a r d and
s o u t h w a r d p r o g r e s s i o n of g l a c i e r s would h a v e eliminated this fauna f r o m its o r i g i n a l area and w o u l d
h a v e pushed it east o r w e s t w a r d , w h e r e s o m e surv i v e d in eastern o r w e s t e r n réfugia, or in b o t h . Af.
selysii w o u l d be the o n l y e x a m p l e of a s p e c i e s surv i v i n g o n l y in a western refugium, the MeuseM o s e l l e r i v e r systems. H o w e v e r , as M. selysii in fact
o c c u r s a l s o in Spain, b e y o n d the P y r e n e e s and far
f r o m the influence of Central European g l a c i a t i o n ,
this assumption is not too likely.
M a r t h a m e a beraudi ( N a v â s ) , spec, propr.
Notes.
T h e f e m a l e t y p e of M. selysii f r o m near L i è g e w a s
no l o n g e r in P i c t e t ' s c o l l e c t i o n in 1865 (A.E. Pictet
1865) and has not b e e n r e t r i e v e d ( Z w i c k 1972). T h e
uniformly dark b o d y suggests that it was of the present species. l i l i e s ' (1955) illustration of the penis
is strongly schematic, existing slide preparations in
coll. lilies agree w i t h the present figure and description. C. B e r t h é l e m y has kindly d r a w n my attention
to the fact that the f i g u r e o f f e m a l e genitalia in the
same paper a p p e a r s t o be a c o m b i n a t i o n of dorsal
1909 Lerpa beraudi Navâs, Brotéria, Ser. Zool., 8 : 102, figs.
Material examined : headwaters of the Jordan river :
Banyas ( = Banias, spring sources of Nahal Banyas = N .
Hermon}:3 9 9 , 4 . VI. 1943 (Bytinski-Salz) ; 3 e r a , 16.V.
1968 ; 2 larvae, 31. VII. 1970 ; Nahal Hazbani (N. Senir) :
1 larva, 15. V. 1967 ; 1 9 , 1 larva, 16. V. 1968 ; 2 larvae,
16.1. 1970; 3 larvae, 31.1. 1970; 1 a, 9. V . 1972 (Kugler) ;
1 larva, VII. 1974 (Freidberg) ; Tel Dan (spring sources of
Nahal Dan) and Nahal Dan ; 1 9 , 7 . VII. 1954 ; 1 o \ 15, V.
1968 (Nitzan); 1 larva, 2. I. 1973 (Freidberg) (all in coll.
Kugler, Tel Aviv). Headwaters of the Litani river, riv. Jahfufah, 1 000-1 200 m, on railroad Beirut-Damaskus, 1 cr,
1 9 , 1 6 . V I . 1978 (N. Alouf). my coll.
136
P- ZWICK
B o t h sexes m a c r o p t e r o u s , w i n g s 16-26 m m long.
Y e l l o w i s h b r o w n species, only a small central m a r k
on the h e a d ( F i g . l c ) and some rugae of the pronotum b r o w n i s h . Outer edges of femora and tibiae also
brownish.
Male.
T e r g i t e 5 w i t h b r o a d entire o r indistinctly divided
spinulose extension. T e r g i t e 6 with variable, n a r r o w
(8)
o r extended spinule patch in soft pale centre of seg
ment. T e r g i t e 7 with b r o w n anterior patch and pos
t e r i o r spinules in centre. A b r o w n i s h central m a r k
on tergite 8 ; tergite 9 n a r r o w , n o r m a l . H e m i t e r g i tes 10 w i t h p o s t e r i o r contour line bent at a blunt
angle. Penis as large as in M. selysii, e v e r t e d sac
huge, distally widened, with many fine uniform spi
nules except a fairly distinct subterminal band of
l a r g e r spinules (Fig. 5).
Fig. 5. Marthamea beraudi, male genitalia. Details and scales as in Fig. 3.
(9)
M A R T H A M E A BERAL'DI A N D ITS E U R O P E A N C O N G E N E R S
Female
and
eggs.
Genitalia not distinctive. E g g oval, 0.47 m m long.
Lid conical, greatest d i a m e t e r a little displaced
towards anchor pole. A raised circular fold delimits
the relatively flat a n c h o r pole w h i c h has a flat w i d e
nipple for a n c h o r attachment (Fig. 6d).
Larva.
Paraprocts without gills. In specimens before me,
dark pigment is v e r y reduced (Fig. 2c) but informa
tion from N . A l o u f suggests that the abdomen may
be dark with a pair of pale patches on every segment.
137
Notes.
O r i g i n and affinities of M. beraudi are not easily
interpreted. Similarities with M. selysii ( l a r g e penis
with p o o r l y differentiated armature) as well as with
vitripennis
( m a c r o p t e r o u s males) are in p r i m i t i v e
traits p e r m i t t i n g no conclusion on close affinities.
P a r a p r o c t gills shared by M. selysii and M. vitripen
nis are another example of such symplesiomorphies.
Clearly d e r i v e d characters are exhibited by only one
of the three species {vitripennis : differentiated penis
a r m a t u r e ; selysii : m a l e brachypterism ; beraudi :
reduced paraproct gills). T h e phylogenetic relation
ships of the three west palearctic species of Martha
mea r e m a i n therefore unknown.
Fig. 6. Eggs of Marthamea ssp. ; a. M. vitripennis ; b,c, M. selysii from R. Lozoya and Moselle, respectively ; d,
M. beraudi. Scale is 0,25 mm.
Key
Males :
1 Macropterous specimens
2
— Brachypterous specimens ; posterior edge of hemitergite semicircularly curved. Tergite 7 with distinct
longitudinal folds in front. Penis very large, arma
ture rather uniform (Fig. 4d)
M. selysii
2 Head largely dark, compare Fig. la ; posterior edge
of hemitergite rectangularly bent. Tergite 5 distinctly
bilobed, 6 practically unarmed. Penis small, with
complex armature (Fig. 3d)
M. vitripennis
— Dark patch on head small (Fig. lc). Process of tergite
5 entire or almost so. Tergite 6 with distinct spinule
patch. Contour of hemitergite curved. Penis very
large, with poorly expressed subterminal ring of lar
ger spinules (Fig. 5d)
M. beraudi
Females with eggs :
1 Contrastive head pattern, area between ocelli and
eyes clear yellow ; eggs oval
2
— Uniformly dark head, or at least light areas between
ocel li and eyes somewhat infuscate, not clear yellow ;
eggs spheroid, anchor pole rather pointed (Fig. 6b,
c)
Ml. selysii
2 Large pale species (wings 19-26 mm), head pattern as
in Fig. lc. Egg with relatively narrow apex and fairly
wide flat anchor pole, large, 0.47 mm long (Fig. 6d)
M. beraudi
P
138
—
Small (wings 18-21 mm) species, dark head pattern
more extended, resembling Fig. la. Oval eggs 0.38 mm
long, widest near middle, both poles rather similar
(Fig. 6a)
M.
vitripennis
Larvae :
1 Gills on paraprocts present : M. vitripennis
and
M. selysii.
—
Paraprocts without gills
M.
beraudi
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1
CONGENERS
139
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