Marburger Entomologische Publikationen Band 2 Heft 3 pp. 217-258 20.10.1986

M a k a r o r y s a n.g. - a r i c w genus for t h e

Canaro-Madeiran Eurysa r i b a u t i LINDBERG-group

w i t h remarks on s p e c i a t i o n , d i s t r i b u t i o n and

phylogeriekic r e l a t i o n s h i p withiri t h i s taxon

(Homoptera Auchenorrhyncha Fulgoromorpha D e l -

phac idae )

w i t h

22 f i g u r e s

by

RE€NHAKD KEMANE and MANFRED ASCHE

Key-words:

Homoptera, Auchenorrhyncha, Fulgoromorpha, Delphacidae, Eurysa

Makaroysa, Madeurysa, M.ribauti, M.canteca, M.madeco, M.madalta, speciation, phylogeny, island-zoogeography; Western Palearctic

Region, Canary Iclands, Madeira.

..

Abs trac t

Closer examination of Euryca ribauti LD.-material (formerly

recorded as a single endemic species from 4 of the western Canary

Islands and from Madeira) from different islands of the Canaries

and from Madeira revealed that this taxon is a monophylum of its

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.

1

Marburger Ent. Publ. 2 ( 3 ) 1986 R E M A N E & ASCHE: Makarorysa 1

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own not closer related to Eurysa FIEB. s.str., thus it is estab-

lished as a separate genus: Makarorysa gen.nov., type-species:

Makarorysa canteca n.sp.. This genus contains 4 morphologically

clearly distinguishable taxa: two allopatrically distributed on

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the Canary Islands (comb.nov.) on La Palma and

La Gornera and M.canteca n.sp. on Tenerife and Gran Canaria) and

two sympatric species on the main island of Madeira (M.rnadalta

n.sp. and M.madeco n.sp.1. The Canarian and the Madeiran species

apparently form two sister-groups, the Madeiran group is

separated as a subgenus of its own: Madeurysa subgen. nov.,

subgenus-type-species: M.madeco n.sp.. Al1 4 species seern to

belong to the set of laurel forest taxa, though they mainly occur

in graslands, where dense forest has already disappeared.

Speciation has very probably occurred allopatrically in the two

Canarian taxa, less certainly so in the case of the Madeiran

taxa. With this we have the first case of evolutionary speciation

within the Delphacidae of the Mid-Atlantic Islands.

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Delphacidae - though in the majority of their species flightless specimens with reduced winps prevail - are amongst those higher

taxa of Auchenorrhyncha which have successfully colonized oceanic

islands and island-groups. They obviously possess an adequate

"spread potential" (LESTOPI, 19571, i.e. they are able not only to

reach islands, but also to establish populations on thern. Beyond this these colonisators have been able in some cases to evolve

subsequently into a sometimes rernarkably large nurnber of

"daughter-species" by adaptive radiation as well as by splitting

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Marburger Ent. Publ. 2í3) 1986 R E M A N E & A S C H E : Makarorysa

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up into island-taxa (e.g. on the Hawaiian Islands, see GIFFARD,

1922; ZIMMERMAN 1948). In contrast to some other groups of oceanic islands the

archipelagoes of the eastern Atlantic Ocean north of the tropic of cancer and south of the 40th parallel of latitude, i.e. the

Azores, Madeira, Selvagens, and the Canary Islands apparently are inhabited not only by a disharmonic set of Delphacid-species, but - - - - - - - - - by a rather poor one, too (about 20 species so far recorded up to

now altogether, that is no more than about 20 percent of the species number of the southwestern part of the Palearctic. Only very few of these species are endemics of only one of these

archipelagoes: Javesella azorica REM., 1974: Azores, "Calligy- pona" gudruna GYLLENSV., 1968: Canary Islands (but this "species"

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is, according to the descriptions and figures published by GYLLENSVARD, 1968, a mixture of Laodelphax striatellus

(FALL . 1 -males and macropterous "Calligyponall anthracina .......................

(HV.)-females - as the holotype is reported to be a fenale, it

will probably run out to be a synonym of the latter). Another cpecies - Eurysa ribauti LINDB., 1936, the one to be dealt with in this article - was first recorded as endemic on the

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Canary Islandc, but later recorded from Madeira, too (LIPiDBERG, 1961) - thus seeming to live on two of these archipelagoes. Al1 - other delphacid-species are widely distributed or at least

from island-groups outside of this range. No cases of known

speciation -_-------

by adaptive radiation o r by splittiig up into island taxa were so far known within the Delphacidae of these archipelagoes.

An examination of "Eurysa ribauti" specimenc from severa1 of the islandc of the Canarian archipelago and from Madeira, however,

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brought about two results:

1. monophyletic group, to which the genotype of Eurysa FIEB., Delphax lineata PERRIS, 1857, belongs (see REMANE & ASCHE, 1983 for further

details): The cynapomorphically specialized features of the male

"Eurysa" ribauti LINDE. is not a member of that ___------------ e----- ----------------

,

Marburger Ent. P u b l . 2 ( 3 ) 1986 R E M A N E & ASCHE: Makarorysa

genitalia (for instance the dorso-basa1 direction of the

peculiarly shaped dista1 part of the aedeagus) of this group are

not present. As indicated before (REMAME & ASCHE, 1.c. 1, within

the old I1genuslf Eurysa FIEB. many species have been united by

severa1 authors, which have in common not more than the

relatively simple characters of more o r less llstoutll body, a

toothless ("stiromine") posttibial spur and a broad head with a

rounded transition between frons and vertex with at least at that

place evanescent carinae. In other characters of probably higher

phylogenetic importance there exist considerable differences

between some of these "Eurysal'-species leaving doubts as to their monophyletic origin. This has already led some authors to

transfer some species into genera of their own (e.g. E.lurida

FIEB. to Eurysula VILB., E.maculiceps HORV. to Stiromoides VILB.

- see NAST, 1972) or to other already existing genera (e.g.

E.clypeata HORV. to Pastiroma DL. by ANUFRIEV, 1981). The

remaining set of Old- and New-World species still is rather

heterogeneous, it is under work in order to analyse the phylo-

genetic relations between these taxa as well as to others - the

recultc may then serve as a base for taxonomic decisionc concer-

ning the formation of supraspecific taxa.

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2. Comparison of specimens from different islands and different

hiotopes revealed the existence of morphological differences ln the genital structures of the males and - though less pronounced

- in size and colouration. Four morphological groups with

non-overlapping ranges of variation were found up to now: two

allopatrically distributed groups on the Canary islands, and two sympatric groups on Madeira. If we assurne the shape of these

morphological structures to be genetically controlled, their

different shapes night be explained to have been brought about by diverging evolution after an initial interruption of gene-flow

between these populations.

In

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case of sympatric occurrence of such groups we may postulate

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M a r b u r g e r E n t . P u b l . 2(3) 1986 R E M A N E & ASCHE: Makarorysa

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t he e x i s t e n c e of i s o l a t i n g mecnanisms: s p e c i a t i o n t o t h e l eve1 o f

t r u e " b i o l o g i c a l " s p e c i e s h a s apparen t ly taken place . No safe

d e c i s i o n concerning t h e degree o f s p e c i a t i o n can be made, o f

c o u r s e , i n case of a l l o p a t r i c d i s t r i b u t i o n of such groups, b u t i n

case of non-overlapping range of v a r i a t i o n of a t least one

character i n one s e x it h a s been proposed t o t reat such groups as

d i f f e r e n t s p e c i e s , t o o (REMANE, 1968).

A s t h e Eurysa r ibaut i- group i s no t a member o f Eurysa FIEB.

s . s t r . , t h e phylogenet ic r e l a t i o n s h i p of t h i s group t o o t h e r t a x a --_------------ ------

s t i l l included i n Eurysa FIEB. o r a l r eady e s t a b l i s h e d as supra-

s p e c i f i c t a x a of t h e i r own had t o be examined i n o rde r t o decide ------

t h e p h y l e t i c p o s i t i o n and t h e sys temat ic s t a t u s of t h i s group.

Besides of t h e g e n e r a l c h a r a c t e r s p r e s e n t i n oviduct - gland

b e a r i n g Delphacini (see ASCHE, 1985) and those aforementioned --------- characters which induced LINDBERG (1936) t o p lace it i n Eurysa, ------ t h e Eurysa r ibaut i- group h a s rnany c h a r a c t e r s i n common wi th some --_----------- o t h e r taxa grouped by WAGNER (1963) i n h i s "Stirominaetl :

1. Reduction t o absence o f d e n t a t i o n of t h e p o s t t i b i a l s p u r , 2. a r e l a t i v e l y long , rather c y l i n d r i c a l male g e n i t a l segment, 3.

Analtube a p i c a l l y n o t c losed by a complete s t r o n g l y c h i t i n i z e d

b r i d g e , thus v e n t r a l s i d e t h i n l y c h i t i n i z e d w i t h t h e except ion of - two t r a n s v e r s e basa1 armlike s t r i p e s almost touching each other

c e n t r a l l y , w i t h d i s t o v e n t r a l l y d i r e c t e d , rnostly s p i n e l i k e

appendages, lobe- l ike s e t o s e membraneous s t r u c t u r e s s i t u a t e d

b a s i v e n t r a l l y of t h e s p i n e l i k e appendages.

4. Aedeagus forrning more o r less a s imple tube wi th t e e t h

a r ranged i n l o n g i t u d i n a l rows, i ts v e n t r a l s i d e l a r g e l y membra-

neous, phallotrerna v e n t r o a p i c a l .

5. Sucpensorium a t t a c h e d medioventralay t o t h e base of t h e theca,

f u s e d d o r s a l l y t o a plate-lii te s t r u c t u r e a t t ached t o t h e c e n t e r

o f t h e basoven t ra l margin of t h e ana l tube .

6. Thinly c h i t i n i z e d reg ion of t h e g e n i t a l segment around t h e

base o? t h e s t y l e s heart- shaped by a v e n t r a l process o f a

M a r b u r g e r E n t . P u b l . 2(3) 1 9 8 6 R E M A N E 6 ASCHE: M a k a r o r y s a

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v e r t i c a l median phragma-rim.

A t t h e moment w e are no t y e t a b l e t o decide which o f t h e s e

p h e n e t i c similarities are not on ly homologous b u t i n a d d i t i o n

synapomorphic c h a r a c t e r s , and i n which sequence they were evolved

- a t least some o f them n i g h t have been developed more than once

or may be considered symplesiomorphic a t a c e r t a i n l e v e 1 of taxa

(and t h e r e f o r e no i n d i c a t o r s of c l o s e r e l a t i o n s h i p ) . Thus t h e

q u e s t i o n of the s is ter- group of t h e Eurysa r ibau t i- group cannot

be s e t t l ed here and now - t h e h i g h e s t number o f phene t i c

s imilar i t ies seems t o e x i s t t o t a x a placed i n He t rop i s FIEB.,

Delphacinus FIEB., and t o come Afr ican t a x a s t i l l placed i n

Eurysa FIEB..

A s t h e s e s p e c i e s and species- groups possess c h a r a c t e r s o f t h e i r

own which are t o be considered more apomorphic than t h e

r e s p e c t i v e c h a r a c t e r s i n t h e Eurysa r ibaut i- group and, on t h e

o t h e r hand, t h e t a x a of the Eurysa r ibaut i- group possess some

s p e c i a l c h a r a c t e r s , i n which they may be cons ide red more

apomorphic than a l 1 o t h e r t axa mentioned here (see below) the

l a t t e r seem t o form a mal1 monophylun of t h e i r own. T r a n s l a t i n g

t h i s s i t u a t i o n i n t o sys temat ica l arrangernent, it is thought more

adequate t o e s t a b l i s h a genus of i ts own f o r t h i s monophylum than

t o j o i n i t w i t h o t h e r taxa which might no t be i t s s i s t e r- g r o u p - t h e l a t t e r would i n e v i t a b l y l e a d t o wroqg zoogeographical or

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e c o l o g i c a l conclus ions concerning t h e ’ h i s t o r i c a l development of

d i s t r i b u t i o n as w e l l as of c h i f t s i n us ing e c o l o g i c a l n iches .

Makarorysa nov.gen.

C o n s t i t u t i v e characters ( i . e . characters assumed t o be synapo-

rnorphic (cornmon der ived) f o r t h i c s e t o f taxa) o f t h e s p e c i e s are

l o c a t e d i n t h e g e n i t a l region:

Male g e n i t a l i a : phragma-bridge wi th i ts median pa r t dorsocaudal ly

M a r b u r g e r E n t . P u b l . 2(3) 1986 R E M A N E & A S C H E : M a k a r o r y s a

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e l e v a t e d , formed l i k e a jugsocke t (see f ig . 2 b ) , w i t h a v e r t i c a l

median r i m weakeninp towards i t s v e n t r a l end. Dista1 end of t h e

s t y l e s twis ted a g a i n s t t h e b a s a l p a r t and formed l i k e a l a n c e t

( f i g . 3a-b). Dorsal row o f t e e t h o f t h e aedeagus s h i f t i n g b a s a l l y

t o t h e l e f t s i d e .

Female g e n i t a l i a : V a l v i f e r VI11 b a s a l l y o f t h e G I I X a t t h e inner

(median) margin wi th a d o r s a l l y i n f l a t e d area bear ing a small

s e m i c i r c u l a r exc i s ion (see f igs . 21, 22) (probably t h e "holds"

for t h e t i p s of t h e s t y l e s d u r i n g c o p u l a t i o n ) .

Addi t ional characters ( p r e s e n t i n i d e n t i c a l o r very similar way

s i n g l y o r combined i n o t h e r t a x a ) as far as no t a l ready mentioned

be fore : Wing dimorphism p r e s e n t , b u t brachyptery less developed

than e.g. i n Metropis FIEB. o r Delphacinus FIEB. : forewings w i t h

d i s t i n c t venat ion, c o r i o c l a v a l s u t u r e p r e s e n t , end of each wing

rounded, ending approximately on t h e s i x t h abdominal t e r g i t e .

Venation of the forewings wi th s c a t t e r e d b r i s t l e s a r i s i n g from

small granules . Colour dimorphism combined with wing dimorphism:

i n long-winged specimens thorax and abdomen l a r g e l y marked wi th

blackish-brown, wings clear, i n s h o r t winged specimens near ly no

d a r k markings on thorax (and head) and less on abdomen, b u t fore-

wings more o r less blackish-brown i n t h e males of t h r e e taxa

( i . e . a d d i t i o n a l sexua l colour-dimorphism i n al1 bu t one of t h e

t a x a ) . - Drumming organ of t h e male wi th long, t h i n S-2-apodomes

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n o t only reaching t h e dorsum, b u t ben t i n t h e i r d i s t a l p a r t s

medially - 6-gen i t a l segment s u b c y l i n d r i c a l , d o r s a l s i d e wi th a

r e l a t i v e l y narrow c h i t i n i z e d b r idge due t o a broadly v-shaped

basal and a more narrowly v- o r u-shaped d i s t a l i n c i s i o n

con ta in ing t h e ana l tuhe (see f i g . 2 c ) . Seen from behind more o r

less c i r c u l a r , margin wi thou t s p i n e s o r exc i s ions except for a

shal low, v-shaped i n c i s i o n on t h e v e n t r a l s i d e . Analtube

inc lud ing ana l s t y l e and surrounding p a r t s of t h e g e n i t a l segment

ye l lowish , t h e o t h e r p a r t s of t h e a n a l segment more o r less

in fusca ted t o black except f o r t h e narrowly w h i t i s h la tera l p a r t

M a r b u r g e r E n t . P u b l . 2 ( 3 ) 1 9 8 6 R E M A N E & A S C H E : Makaroryca

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of t h e p o s t e r i o r nargin . S t y l e s d ive rg ing , r e l a t i v e l y s h o r t , n o t

b i f u r c a t e d .

Aedeagus: s t r a i g h t , v e n t r a l s i d e excep t f o r t h e b a s a l p a r t o f t h e

theca t h i n l y c h i t i n i z e d , phal lo t rema ven t roap ica l , l a t e r a l l y

equipped from base t o t i p wi th d e n t a t i o n ( t e e t h o f d i f f e r e n t

s i z e , p a r t l y ar ranged i n l o n g i t u d i n a l rows).

Female g e n i t a l i a : two a t r ium- plates (d ivided by a narrow median

l i n e o f t h i n c h i t i n i z a t i o n ) s i t u a t e d cephal ly of t h e C V I I I .

Typus gener i s : Makarorysa c a n t e c a nov. spec.

As mentioned be fore , Makarorysa n.g. was found t o c o n s i s t o f f o u r

morphologically separab le taxa i n s t e a d o f only one. The t w o

Canarian t a x a have s o fa r been found on f o u r o f t h e f i v e t r u l y

ocean ic i s l a n d s : t h e one i n h a b i t s t h e c e n t r a l l y s i t u a t e d , r a t h e r

large i s l a n d s of Tener i fe and G r a n Canaria, t h e o t h e r t h e i s l a n d s o f L a Gomera and La Palma. These two taxa are r a t h e r similar t o

each o t h e r , t h e d i f f e r e n c e s between them are loca ted i n t h e male g e n i t a l i a , e s p e c i a l l y i n t h e shape of t h e processes of t h e anal-

tube and i n t h e basal p a r t of t h e aedeagus. Both Canarian t a x a

d i f f e r c l e a r l y i n severa1 c h a r a c t e r s from t h e two Madeiran t a x a

(see t a b l e 1). The two Madeiran t a x a apparent ly form a monophylum

o f t h e i r own, though t h e morphological d i f f e r e n c e s between them

are more developed than between t h e two Canarian taxa. I n o r d e r

t o express t h i s s i s ter -group r e l a t i ó n sys temat ica l ly a new

subgenus is e s t a b l i s h e d f o r t h e Madeiran Taxa :

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Madeurvsa n.subepn.

C o n s t i t u t i v e c h a r a c t e r s : theca of aedeagus on i ts v e n t r a l base

wi th a d i s t i n c t , a p i c a l l y d i r e c t e d process surmounting t h e base

of t h e membraneouc p a r t o f t h e v e n t r a l s i d e .

A l 1 o t h e r c h a r a c t e r s , i n which the Madeurpa-taxa ------ -- di f fe r from

t h o s e of Makarorysa s.str. are e i t h e r more plesiomorphic (e.g.

s imple form of the suspensorium) o r t h e i r evo lu t ionary s tate is

n o t t o be assessed w i t h c e r t a i n t y .

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Marburger Ent. Publ. 2 0 ) 1986 R E M A N E & A S C H E : Makarorysa

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Typus subgeneris: Makarorysa (Madeurysa) madeco nov. spec.

At least three of these four taxa - the two from Madeira and one

from the Canaries - have to be rlescrjbed as new - but which of the two Canarian taxa is Eurysa ribauti LINDBERG, 1936?

LINDBERG (1936) had described his species Eurysa ribauti after

three specimens (2dd, 19) collected by Frey and Stora on the

Canary Islandc: one specimen each on La Palma, Tenerife, and Gran

Canaria. LINDBERG 1s figures of the male genitalia show a rather

sketchy caudal and later in addition (LINDBERG, 1954) a lateral

asnect of the genital capsule, obviously no examination of the analtube, phragma or aedeagus was ever made by him. As usual

LINDBERG (1936) in his description mentioned no type locality,

but the collection-numbers of the type material only (in this

case he published the designation of two holotypes - "Holotypen, 6 (No 76641, 9 (No 7665)..." - but this rather unusual situation

seems to have been altered: a few years ago Dr.Meinander, from Helsinki Museum kindly informed me that the first mentioned male is kept as holotype). This male specimen was collected on La

Palma, the name ribauti LINDO. thuc has to be applied to the La

Palma -La Gomera-species, the one from Tenerife and Gran Canaria,

though already contained in LINDBERG' s original material, has be named and described.1)

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1)This could he confirmed by re-exaaination of the type material of %.ribauti Li!. kinely sent to us by Dr. A.Jansson, Helsinki. The holotype d is from La Palna, - the allotype 9 from Gran canaria concerns another species, and here ic additjonally inteprated as peratype into the tyne-series of !;.canteca n.sn..

Marburger Ent. P u b l . 2 í 3 ) 1986 REMANE E. A S C H E : Makarorysa

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1. Makarorysa (s.str.1 canteca nov.spec. (figs. 6-10, 21) Body size and proportions, colouration, markings and degree of sexual and wing dimorphism like in M.ribauti (LINDE., 1936). d

Genital segment with phragma-bridge and styles like in M.ribauti

(LD.), but appendages of the analtube with simple tip (not

bifurcate as in ribauti) (fig. 9 ) ; basa1 part of aedeagus (fig. loa-d) laterally less dentated, base of thinly chitinized ventral

region of aedeagus much broader (see fig. 10b). Female in al1 characters so far examined not differing from of M.ribauti (LD.). Holotype d : Canary Islands, Tenerife, Anaga-area,Igueste,

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that

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15.3.66. R.Remane, leg., in coll. Remane, Elarburg. Paratypes (numerous dd and 99) from above mentioned locality and date in addition from severa1 other localities, especially in Anaga- and Teno-area and from Gran Canaria (Reg. El Palmital),

Remane leg., in coll. Remane, Marburg.

Biology: Found especially on the northern sides of Tenerife and Gran Canaria from the lowlands above sea .levei up to about 1000 m

altitude. Like M.ribauti (LD.) apparently a species of the laurel forest region, but mainly found in marginal or open places, where

grasses grow, by this occurring in forest clearings, on meadow terraces, way sides of cultivated fields and similar biotopes. Probably no strict synchronisation of development, presumably more than one generation per year.

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1

1

Geographic Distribution: Endemic the oceanic islands Gran Canaria and Tenerife, replacing there

species of the Canary Islands, so far found only on

M a r b u r g e r E n t . Publ. 2(3) 1 9 8 6 R E M A N E & A S C H E : Makarorysa

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M.ribauti (LD.). ---------

2. Makarorysa (Madeurysa) madeco nov. spec. ( f i g s . 11-15, 22

Length dd (brachypterous): 2,6-2,7 mm, 99: 3,4 mm

Rela t ive ly la rge spec ies , sexual dimorphism i n t he colourat ion of t h e brachypterouc form no t developed (plesiomorphic s t a t e o r

secondary loss?). Forewings i n male greyish-stramineous as i n the

female, grey markings of t h e abdomen only i n some male specimens

more developed than i n t he females.

Geni ta l s t ruc tu re s of t h e male character ized by very broad,

depress appendages of t h e anal tube (see fig. 14a-b) and a near ly

symmetrical, d i s t a l l y notched, r a t h e r sho r t process of t he

ventrobasal p a r t of t h e aedeagus ( f ig . 15b).

Females: Srnall females perhaps no t dis t inguishable from females

of t h e following spec ies M.(M. Imadalta n.sp.

d Holotype: Madeira, Prazeres , ca 400 m , 8.4.67, R.Remane leg. ,

i n c o l l . Remane, Marburg.

Paratypes (dd and 99) from t h i s place and from several other

l o c a l i t i e s s i t u a t e d i n d i f f e r e n t regions and a l t i t u d e s of t h i s

i s land: Remane leg . , i n c o l l . Remane, Marburg.

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Biology : . ,

I n g rasses from coas t a l a reas up t o 1800 m (Pico Ruivo). Probably

severa1 generations per yea r .

Geographic d i s t r i bu t ion :

Endemic species of the Madeira main i s l and , l i v ing there

sympatric w i t h F.1. (M. madalta n.sp. --------------

M a r b u r g e r E n t . P u b l . 2(3) 1986 R E M A N E & ASCHE: M a k a r o r y s a

3 . Makarorysa (Madeurysa) madalta nov.spec. ( f i g s . 16-20)

Length: c‘ (macropterous): 3 , 3 mm; d f (brachyperous) : 2 , 4 mm, 99

(b rachyp te rous ) : 2,7-2,8 mm

Smal les t of a l1 f o u r s p e c i e s , sexual dimorphism i n t h e

c o l o u r a t i o n of t h e brachypterous form very pronounced: head,

thorax and t e g u l a e s t ramineous , wings shiny b lack i n t h e male,

wholly stramineous i n t h e female.

G e n i t a l s t r u c t u r e s of t h e male c h a r a c t e r i z e d by s l e n d e r , n o t

depresced appendages of t h e a n a l t u b e ( f i g . 19a-b) and subacute ,

aeyrnmetrically based long p rocess o f the ven t robaca l p a r t o f t h e

aedeagus ( f ig . 20a-d).

Females: i n p ropor t ions and c o l o u r a t i o n similar t o t h e preceeding

s p e c i e s b u t smaller - more material has t o be exarnined t o dec ide ,

whether t h e r e e x i s t cons tan t d i f f e r e n c e s t o M.(M.) madeco n.sp..

d Holotype: Madeira, east o f P ico de Areei ro , ca 1450 m, -------------

4.4.1967, R.Remane leg., i n c o l l . Remane, Marburg.

B io logy :

So far found only i n s p r i n g 1967 i n a grazed, meadowlike c l e a r i n g

surrounded by Erica-Vaccinicum-shrub on a mountain r i d g e east of

Pico Areei ro a t a n a l t i t u d e on 1450 rn. The specimens were found I.

i n t h e tussocks o f a Juncus s p e c i e s , but rnay have gone t h e r e f o r

s h e l t e r i n r e a l i t y f eed ing on grasses.. The cpec ies was n o t

p r e s e n t ( excep t perhaps i n t h e egg*stage) a t t h i s l o c a l i t y i n

J u l y 1977.

”

Geographic d i s t r i b u t i o n :

Endemic s p e c i e s of Wadeira main i s l a n d l i v i n g t h e r e sympatr ic

w i t h M.(M.) madeco n.sp. , rnaybe confined t o high a l t i t u d e

b io topes . __------------

- ~~ ~ ~~

M a r b u r g e r E n t . P u b l . 2(3) 1 9 8 6 R E M A N E & ASCHE: M a k a r o r y s a

- 229 -

The f o u r taxa c o n s t i t u t i n g t h e genus Makarorysa n.g. are p a r t of

t h e L a u r i s i l v a (laurel f o r e s t ) se t o f t a x a and - as many o t h e r

t a x a o f t h i s b iotop - probably r e l a t i v e l y o l d i n h a b i t a n t s of

t h e s e i s l a n d groups showing a r e l i c t a r y type o f d i s t r i b u t i o n . A s

mentioned, t h e ques t ion o f t h e i r n e a r e s t r e l a t i v e s cannot be

answered by now - maybe some of t h e African "Eurysa" w i l l be

c a n d i d a t e s f o r t h i s .

S p e c i a t i o n i n t h i s group h a s probably occurred by s p a t i a l

s e p a r a t i o n : very probably so i n t h e case o f t h e two a l l o p a t r i -

c a l l y d i s t r i b u t e d Canarian taxa (though it seems remarkable, t h a t

no rnorphological d i f f e r e n c e s were found between t h e popula t ions

o f each of these taxa l i v i n g on two d i f f e r e n t i s l a n d s : is g e n e t i c

exchange rnaintained between t h e popula t ionc o f La Palma and La

Gomera, bu t no t between t h e popu la t ions l i v i n g on T e n e r i f e and

t h e nearby L a Gomera? O r are t h e r e q u i t e r e c e n t e v e n t s o f coloni-

s a t i o n , i.e. La Gomera r e c e n t l y co lon ized by specimens from La

Palma? Why has no r e p r e s e n t a t i v e y e t been found on E l H ie r ro i n

s p i t e o f t h e presence of L a u r i s i l v a on t h a t i s l a n d ? ) .

Also i n t h e case o f t h e two s y n p a t r i c s p e c i e s on Madeira main

i s l a n d al lopatric s p e c i a t i o n seems p o s s i b l e : two co lon iz ing

e v e n t s separa ted by s u f f i c i e n t time f o r the first c o l o n i s a t o r ( o r

t h e s p e c i e s on t h e c o n t i n e n t ) t o d i f f e r e n t i a t e i n t o a s p e c i e s of

i ts own might have taken p lace . Sympatric s p e c i a t i o n by adapt ive

r a d i a t i o n t o d i f f e r e n t environments cannot be excluded, of

course .

Whether both Madeira and t h e Canary I s l a n d s have been colonized

independent ly from c o n t i n e n t a l r e g i o n s , o r whether one o f these

i s l a n d groups has been co lon ized by specimens stemming from the

o t h e r ("stepping-stone p r i n c i p l e " cannot be answered y e t .

F u r t h e r r e sea rch is needed t o f i n d o u t t h e autecology and

e thology of these spec ies . Crossbreeding experiments might help

t o e l u c i d a t e t h e r e l a t i o n s h i p s between t h e f o u r species.

----------

------

21

I O o (u

I

t

Tabe l le 1: Morphological d i f f e r e n c e s between t h e Canarian (Makarorysa s.str.) and Madeiran (subg. Madeurysa n.sg.)

taxa of Makarorysa n.g.

Characters assumed t o be synap0morphous:-

(e rac ters without underlining: evo lu t ionary state n o t y e t decided)

Charac te r s assumed to be symplesiomorphous ....

Makarorysa s.str. Madeurysa n. sg .

8 G e n i t a l characters Thinly c h i t i n i z e d area around aedeagal base

reaching l a t e r a l l y v e n t r a l only t o t h e dor-

sal margin of t h e diaphragm (see f i g .

d i s t a l p a r t of t h e s t y l i r e l a t i v e l y s h o r t

( f i g .

suspencorium between aedeagus and anal

tube completely fused and c o n i c a l l y produ-

ced caudad (see f i g .

D i s t a 1 part of aedeagus r e l a t i v e l y s l e n d e r

( f i g .

Basa1 part of t h e aedeagus l a t e r a l l y ex- p n d e d ( f i g .

Ventrobasal p a r t of t h e aedeagus b a s a l l y

of t h e membraneous pqr: pf,fce.yer$& s i d e n o t d i s t i n c t l y d i s t a l l y produced

( f i g .

.....................................

..................

..................................

9 Genit

Phallotrema a t t h e end of t h e v e n t r a l side

of t h e aedeagus , t i p of t h e aedeagus in

d o r s a l view pointed ( f i g .

1 c h a r a c t e r s Inner margin of VE V I 1 1 b a s a l l y of t h e ex-

c i s i o n without acu te proloiigation ( f i g .

Deiitation of po5t-

t i b i a 1 spur

Most specimens without rests of t e e t h

Thinly c h i t i n i z e d area around aedeagal base

reaching l a t e r a l l y c l e a r l y more v e n t r a l than

t h e dorsal margin of t h e diaphragm (see f i g .

d i s t a l p a r t oí? t h e s t y l i longer , m o r e s l ender

( f i g .

suspensorium beiween aedeagus and ana l tube

medial ly p a r t l y membraneous, n o t c o n i c a l l y pro-

duced (see f i g .

D i s t a 1 part of t h e aedeagus s t o u t and broad

a

...................................... .......................................... ......

( f i g .

B a s a 1 part of t h e aedeagus s t r a i g h t , no t lateral-

l y expanded ( f i g .

Ventrobasal part of t h e aedeagus b a s a l l y of the

membraneous part of t h e v e n t r a l side d i s t i n c t x y

d i s t a l l y produced ( f i g .

..........................................

..........

Phallotrema s i t u a t e d ven t roap ica l ly , t i p of

t h e aedeagus i n d o r s a l view d i c t i n c t l y not-

ched ( f i g .

Inner margin of Vf V I 1 1 basa l ly of the ex-

c i s i o n a c u t e l y prolouged ( f i g .

Small i r r e g u l a r t e e t h present ........................

Marburger Ent. P u b l . 2 ( 3 i 1986 R E R A K E 6 ASCHE: Makarorysa

- 231 -

Peferences

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rhynche 1. - Reichenbachia 19( 28) : 159-173

ASCHE,M., 1985: Zur Phylogenie der Delphacidae LEACH, 1815 (Homoptera Cicadina Fulgoromorpha) Teil 1: Text, Teil 2: Abbildungen. - Marburger Ent. Publ. 2(1): 910 pp.

GIFFARD,W.M., 1922: The distribution and island endemism of Hawaiian Delphacidae (Homoptera) with additional lists of their food plants. - Proc. Hawaiian Ent. Soc. 5: 103-118

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LINDBERG,H., 1936: Die Cicadinen der Kanarischen Inseln. - Soc. Sci. Fennica, Comment. Biol. XXIV(1): 1-19

LINDBERG,H., 1954: Hemiptera Insularum Canariensium. - Atlas Sci. Fenn. 1: 1-304

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NAST,J., 1972: Palearctic Auchenorrhyncha (Homoptera). An annotated check-list. - Warszawa (Polish Scientific Publishers), 550 pp.

REMANE,R., 1968: Erganzungen und kriticche Anmerkungen zu der Heterop- teren- und Cicadinen-fauna der Makaronesischen Inseln. Boc. Funchal, 16: 1-14

REMANE,R. e( ASCHE,!!. , 1983: Zur geperischen Stellung von Metropis for- ficula HORVATH, 1908 und einiger neuer verwandter Taxa aus der Südwestpalaarktis (Homoptera Auchenorrhyncha Fulgoromorpha Delphacidae): ein Formenkreis allopatrischer Taxa? - Marburger Ent. Publ. l(8): 57-84

WAGNER,W., 1963: Dynamicche Taxionornie, angewandt auf kiie Delphaciden

Mitteleuropas. - Mitt. Hamburg 2001. Mus. Inst. 60: 111- 180

ZIMliERMAN,E.c., 1948: Homoptera: Auchenorrhyncha, Insects of Hawaii, Vol. 4, University of Hawaii Press, Honolulu, 268 pp.

Figs. 1-2c' ( g c n j t a l s t r z c t w e r or' rzlesl k,ave beer, nade a f t e r s p e c i -

w n s of w!-.i.ich í;n? z.Sdc>rleri was vacerate:! i n 10% KOH, t h e n t r a n s f e r r e d

i n t 9 g l y c s r i n , res?. g l y c e r i n - g e l ~ . t i n ~ ,

F i g s . 21-22 ( g e n i t a l c t r u c t u r e s o? f e n a l e s ) have been t aken f r o n

specirnecs d r i e d by a i r .

F i g s . 1-5: i?akarorysa r i b a u t i ( L I N D B E R G ) (La Pa lma)

F i g . 1: d- g e n i t a i i a

------------------

a : ven t rocauda l view

b : l e f t l a te ra l view

Fig. 2: pygofer

a: caudal view

b: ventral view

c : dorsi: view

- 2 3 3 -

u*- .-."--