~-,------~------~~~ I~(~' cJ~ tt---- f;;x.-&~ 325

THE fllSTOfliONA THORELL {J~raneae, t\gelen~dae) WITH DESCRIPTION OF T\NO NEVV CAVE-DWELLING SPECIES by

Chrbt3 L. Dee!eman-Reinhold

Sparrenlaan 8 Ossendrecht (Pays-Bas)

The number of known species assigned to both Thorell and Roeweriana Kratochvil has in recent years been growing steadily. lvleanwhile, distinction between these genera is becoming vague, if existing at all. Thf genus Histopona was crea·:ed in 1869 for Tegenaria torpida (C.L. Koch), an epigean, agelenid spc('i(~s which, according to Thor·~ll (1 869) ·differs from other Tegenaria species by the slightly recurved posterior eye row and the distal ~ egment of the superior spinners longer than the basal segment. Later, T. /uxuriulls Kulczynski, T. conv,?nicns' KuJczynski, T. facta Kulczynski and T. sirlllata Kulczynski were added to Histopolla (Roewer, 19:)4; Lehtinen, 1967). Lehtinen also provided a diagno.;is of this genus. During the last decade, new species were assigned to Histopona by Brignoli (1976, 1977, J980) and Del­ tshev (1978). As the characters furnished by Thore!! do not always apply, many aut1Jor~ did not recogni­ ze the genus, among others Simon, Drensky, Bonnet, Wiehle and Loksa. Brignoli (1971 a) advocated the generic separation of H. torpida and related species from Tegenaria on the basis of the structure of the vulva: H. torpida is provided with an enlarged, membranous segment of the seminal duct, the «bursa copu!atrix», a conformation never found in any Tegenaria species and not unlike that found in the american genus Ageienopsis Giebel (Gering, 1953), but differing from t~Jat genus by the absence of " cOllpl:.ng cavity. I ilgree wil.h this point of view, the mOi-e since Hisfoporiil c8.tegorically differs from Tegenaria in some other respects too_ Rocl\-eria.'1a falls entireiy withiil the .limits of Hisfopolla. In the past, according to arachnological tradition, much weight has been accorded to eyelessness and many genera have been founded only' on this character. Thus arose genera like Typhloneta Ku1czynski 1914 for blind Trog/ohyphantes, Typhlonesticus Kulczynski 1914 for blind Nesticus, Adoxoneta Simon 1929 for a blind Centromerus, Typhforhode Kratochvil 1932 for blind Rhode etc. The genus Stalita Schibdte served for a long time as a dump for all sorts of unrelated bllnd Dysderidae. In general, the phylogenetic value of this character has heen grossly over-estimated. In /?,JclVeriana too, rnicrophthalmy is not valid as distinctive character. As phenotypic classification was gradually replaced by a phylogenetic system, it became clear that many anophthalmic and microphthalmic species are close relatives of forms with normal eyes. Loss of eyes in cavernicolous shollid be regarded as a minor, relatively easily acquired adaptation. One of the first to realize this was Kratochvil (1938). Noticing the discrepancy in phylogenetic characters between tegenarioides Kcyserling and the rest of the Hadites species, he created the subgenus Roeweriana for the latter. Later, also Brignoli (1971 b) emphasized that the vulval structure in Hadites regel/arioides is fundamentally different from that in Roe..veriana, anei that the latter is close. to ifisropona (l972). True, Roewer,'ana species are like Hisropona characterized by the peculiar type of epigyne with translucent, marsupium-like valves ancl ample bursae copulatrices, but they remained for a time distinguishahle from His{o;JOna not only by their troglobitic appearance ancI geographical distribu­ tion, but also by a set of minor gEnitalic differcr:ces ill the male. However, \vith the discovery oCrnore species in Europe belonging to this complex and being interme­ diate in appearance, with a mixture of Hisropona and Roeweriana traits, the morphological gap between Christa Dceleman-Rcinhold 326 the two genera was overbridged . If Roeweriana were to be maintained as a separate subgenus or genus, I hen for the sake of consistency another such taxon would have to be created for each of the species iso/ata n .sp., itatica Brignoli and pa/aeo/ithica Brignoli, whereas sinuata Kuk and vignai Brignoli would start to commute between them. In the present case, once again disappearance of functional eyes is of little value for c1assi fication. There is no reason to treat Histopona and Roeweriana as separate genera. Histopona s.l. as defined below is equipped with a set of non-pareil, synapornorphic characters.

GENUS HISTOPONA THORELL

Type species: Histopona torpida (C.L. Koch) ';yn. : Roeweriana Kratochvil 1938, type species R. bidens Absolon & Kratochvil 1932. Dil1xnosis. Spiders of medium size, resembling Tegenaria and Textrix. Head with straight front and parallel sides, sometimes slightly constricted behind the eye region. Width of head approxim:A~ely half the maximum width of carapace. Anterior median eyes, when fully developed, 2-3 times smaller than amerior laterals, the former may even be lacking; anterior row straight. Posterior eye row straight or sllghtly recurved, posterior eyes equal in siz.e to anterior laterals and about one d apart. In many of the southern species eyes reduced in size and unpigmented. Chelicerae with lateral condyle and 3-4 teeth in the anterior, 3-7 in the posterior row. Sternum pointed between coxae IV. Legs of medium length, trochanters notched, all segments except coxae and trochanters with spines; tar.s..i I-Ill spineless, tarsi IV with 1-2 prolateral spines. Legs TV IO-20 070 longer than legs I. Distal segmeIltOf"ftlperior spinners as long as or slightly shorter than basal segment, inferior spinners 8S long as basal segment of superior spinners, but stouter than these, l:olull13 small, p~in:~c!:-, /, Male palp with or without patellar apophysis and with one or more tibial apophyses. Cymbium distally constricted, elongate, embolus long and thread-like, supported by the conductor; radix sometimes w1th Cl ridge or apophysis, median apophysis lacking. Epigyne typically larg~ and bulging, a translucent, marsupium-like, srnoothvalve arising from the posterior margin, constricted in the middle or bipartite, ,:ovcring the entrance apertures of the vulva. The internal paired copulatory organ consisting of an entrance aperture, leading into the ample, plicate and coiled bursa copulatrix, which is connected I hrough a strongly sclerotized and ,coloured cluct into the spermathecae, the latter having a posterior position and opening through a short fertilisation duct into the ut.erus. Similar, but. smaller epigynal v,tlves are found in Hadites tegenarioides KeyserJing ancl in Malthonica dear/ali Brignoli, but. these spe,;ies differ fundamentally both in vulval st.ructure and in somatic characters. The males of both these species are unknown. The ant.erior median eyes in Histopona are much smaller than those in Tegellaria,. the median apophysis, always present in male palps of Tegena­ ria, is lacking in Histopona. Funhermore, Histopona can be distinguished in the field from Tegenaria by its vagary habits, its stouter less and its narrower head. The genus is also related to Textrix, but this latter genus has a much stronger recurved posterior eye row, longer distal segments on the superior spin­ ners, fewer cheliceralteeth and its genital organs are basically different. Many species, particularly those with southern ranges, are cave-dwelling and show various degrees of adaptation to the hypogean environment. They are usually larger than epigean species (exception: pa­ laeofithica) and paler in colour ; the vivid pattern on the abdomen of epigean species is lost and the anl1ulations on the femora disappear. The erect, tactile hairs on the sternum and underside of the coxae and fc:mora are longer and more dense than in the epigean species. These adaptations have been pushed furthest in group myops : these spiders have a troglobitic appearance, are uniform pale yellow and the eyes are reduced in size and pigm,~ntless. Loss of eyes is rare, but loss of the anterior medians seems to be a specific character in pa/aeo/it!!ica. gecords from caves in northern 'Yugoslavia concern the epigean species torpida and luxurians. In the cave-dwelling species, adult males are rare. The genus now comprises the following species: - group torpida .' H. (ofpida (C.L. Koch), H. /uxurians (Kulczynski), H. conveniens (Ku!czynski), H. sinuata (Kulczynski), H. vignai Brignoli, /-1. egonpretneri n .sp. - group myops : H. myops (Simon), H. bidens (Absolon & Kratochvil), H. dubia (Absolon & Kratochvil), H. krivosija­ na (Absolon & Kratochvil), H hauseri (Brignoli), H. tran(eevi Deltshev.

i am indcbted to P:·of. P.M. Brignoli f(,r lending mc types or par3ty;'cs of H. ilOlica, pa(aeoli/hica, vignai and sirinalii and for cornm,unicating nle his \"iews on tht' :a;xon,)rnic problems in ~his group. Ullkss orh(4rv..'ise st8[cd, the m~teri(-l1 entnlcnued below is in the coilection Dc-eleman. The genus Histopona 327

- group strinatii : H. strinatii (Brignoli), H. isolata n. sp. - group italiea : H. italiea Brignoli - group palaeolithiea : H. palaeolithiea (Brignoli)

Key for identification of the species groups 1 a : Epigynal valve very small, undivided group palaeolithiea 1 b : Epigynal valve larger, constricted in the middle or composed of two halves ...... 2 2 a : Epigynal valve constricted in the middle; bursa copulatrix membranous, rigid ducti twisted but not coiled. Male palp : patella with apophysis (except vignai) conductor supporting the distal part of the embolus, the latter originating distally from the radix group torpida 2 b : Epigynal valve consisting of two separate halves. Male palp : patella without apophysis, embolus originating proximally from the radix ...... 3 3 a : Bursa copulatrix with a median, unpaired and a lateral, paired section, the latter weakly sclero­ tized. Male palp : conductor small, basal part of embolus resting on apophysis of the radix of the same shape as the embolus amd partly fused with it ...... group italiea 3 b : Bursa copulatrix paired. Male palp with conductor, but no apophysis on the radix 4 4a : Bursa copulatrix rigid, ducti quasi helicoidally coiled, spermathecae situated near the middle . ••••••••••••••••••••••••••••••••••••••••••••••••I ••••••••••••••••••••••••• , • .• group myops 4 b : Bursa copulatrix membranous, epigyne posteriorly with chitinous patch in the middle; spermathe. cae widely separated ...... group strinatii

,, ,

.-...... - ..-

,, Map. l. Distribution of species of group torpida in Yugoslavia. t',

00000 southern limit of the range of H torpida ...... southern limit of the range of H. luxurians • H. conveniens,. 0 H. egonpretneri "

328 Christa Dee1eman-Reinhold

HISTOPONA TORPIDA (C.L. Koch) (fig. 3, map 1)

Age/ena torpida C.L.Koch 1834, in : Herrich-Schiiffer, Deutsch. Ins: 125. Tegenaria torpida C.L. Koch 1837, Uebers. Arachn. Syst. I: 13. Histopona torpida Thorell 1869, N. Act. Reg. Soc. Sci. Upsala 3 (7) : 133. Histopona debi/is Thorell 1875, Tijdschr. Ent. I: 95, <;> ?, loc. typo : Nice; Simon 1875, Arachn. de France 11 : 205. Tegenaria /aeta Kulczynski 1897, in : Chyzer & Kulczynski, Aran. Hung. II : 170, S?' fig., loco typo Rumanian Banat : Sasca Mon­ tana, Baile Herculane, New synonymy. Remark: Kulczynski's description of laeta agrees in all respects with torpida ,. «patellae inermis» is probably a lapsus. Its locus typicus lies within the range of H. torpida. Vulva: fig. 3 Distribution: ll1iddle cOI1~inental E~!_~pe, southern Europe _~xcept the mediterranean region.

2

1 I >------1 ! i I~

Figs. I, 2 : Histopona luxurians. 1) S?, Ulc g, Hercegovina, vulva, dorsal view; 2) id., right half of vulva with stuck embolus. Scale­ lines 0.5 mm. Fig. 3 : Histopona torpida, S?, Trnovo, northwestern Yugoslavm, left half ofYulva, dorsal view. Scale-lIne 0.5 mm. Figs. 4, 5, 6 : Histopona egonpretneri n. sp. 4) S?, paratype, vulva, dorsal view; 5) Id. epl6yne ; 6) [d., eye regIOn, dorsal view. Scale-line 0.5 mm. The genus Histopona 329

HISTOPONA LUXURIANS (Kulczynskil (figs. 1, 2, map 1)

Tegenaria luxurians Kulczynski 1897, in : Chyzer & Kulczynski, Aran. Hung. II : 172,d''(, figs., loco typo northwestern Croatia, Quarnero. Histopona luxurians Roewer 1954, Katalog der Araneae 2a : 63. Material examined: numerous6' and 9 from Croatia, Bosnia-Hercegovina and Montenegro. Distribution: Yugoslavia, Austria, Hungary, Bulgaria. This species is common in Yugoslavia above 600 m in forest litter. In Slovenia it is also frequently found in caves; in the caves of the Dinaric karst it is replaced by the closely related H. conveniens. In a? from Ulog, Hercegovina, a broken-off embolus was found stuck in a vulva (fig. 2). It shows that the embolus is introduced laterally, then follows the convolutions of the bursa, distorting them on the way; the tip has penetrated the rigid ductus till quite close to the spermatheca.

HISTOPONA CONVENIENS Kulczynski (map 1)

Tegenaria (Histopona) conveniens Kulcz:rnski 1914, Bull. Ac. Sci. Cracovie B 1914 : 383,0'( figs., loco typo Hercegovina, Bjelusica pecina near Zavala. Tegenaria laeta vaT. paganettii Nosek 1~'05, Verh. ZooI. Bot. Ges. Wien 55 : 216, loco typo Hercegovina, cave near Jasen. New synonymy. Material examined: numerouso ,9 and juvenile specimens from Croatia: Trebinje, Dolska peCina (= Eliashohle) ; region Mt. Biokovo : Zagvozd, Mu'iina peCina, Masina pecina and Celar jama, cave near Katu~ici and Vilenjaca spilja. B-Jsnia-Hercegovina : Zijemlje, cave near Padezine ; Divin : Danojlina peCina. Montenegro: Velimlje, cave Cista vIada ; Krstac : Kerovacka peCina near Nesiste ; Titograd, Du~ica pecina near Peuta. All th,~se localities are new. Distribution: southwestern Yugc·slavia. The species had only been recorded from caves around Popova polje in Hercegovina by Kulczynski (1914) and Kratochvil (1934). The record of H. conveniens from Bulgaria (Drensky & Jurinitsch 1917) from near a waterfall in the montane region of Mount Vitcosha near Bojane, may concern either H. tranteevi Deltshev or H. luxuria.'1s (Kulczynski). This latur species was recorded from Vitosha mountain by Deltshev (1967). H. conveniens is closely related to H. luxurians .. its genital organs differ only in details. H. conveniens is markedly larger and paler in colour and lacks the contrasting pattern on the abdomen of H. luxurians. The eyes are normal in size. The species has been found in caves only.

HISTOPONA EGONPRETNERI n. sp. (figs. 4-8, map 1)

Locus typicus : Yugoslavia, Cr,)atia : Mt. Velebit, National Park Paklenica, in cave Manita pee. The species is only known from its type locality. Material examined: 16' , holot~'pe, taken as subadult on 26.X.1978, maturity moult 7.X.1979, killed 6.xU979; 19, paratype, 14.VLl975 ; 1 », paratype, 4.V.1978 ; 19 ' paratype, 2l.IV.1976, all leg. P.R. + C.L. Deeleman from the type locality. Description :0'» : Carapace and chelicerae pale yellow-orange, sternum and ventral side of coxae white, legs ochre, abdomen creamy while. Eyes slightly reduced in size (fig. 6) and surrounded with thin, black lines, posterior row slightly recurved. Chelicerae with 4 teeth in the posterior row. Measurements in mm :d', holotype, carapace 3.55 long. 2.54 wide, abdomen 4.20 long. Legs: I: 3.84 ­ 1.20 - 3.84 - 3.85 - 2.16 = 14.89 ; II : 3.60 - 1.06 - 3.46 - 3.65 - 2.02 = 13.79 ; III : 3.36 - 1.06 - 3.12 ­ 3.84 - 1.92 = 13.30; IV: 4.22 - :.15 - 4.08 - 5.38 - 2.40 = 17.23. 9, paratype (2l.IV.1976), caraJace 3.36 long, 2.21 wide, abdomen 4.40 long. Legs: 3.41 - 1.20 - 3.46 - 3.41 - 2.16 = 13.64; II : 3.36· 1.06 - 3.02 - 3.31 - 1.92 = 12.67 ; III : 3.36 - 1.06 - 2.88 - 3.36 - 1.68 = 12.34; IV : 3.84 - 1.15 - 3.79 . 4.56 - 2.16 = 15.50; palp : 1.54 - 0.62 - 1.10 - 1.58 = 4.84. Male palp (fig. 7, 8) : femur with 3 dorsal spines, patella and tibia. each with one dorsal spine proximally, tibia also with a distal dorsal spine. Patellar apophysis as long as the whole segment; tibia distally with a complex protuber,mce bearing various apophyses. Cymbium and embolus both relatively short, conductor a massive sclerite, lacking the distal spoon-shaped projection found in torpida and luxurians, its broad distal plate supporting the embolus. Radix with proximal cup-shaped depression supporting the base of the embolus. 330 Christa Deeleman-Rcinhold

Epigyne and vulva (fig. 4, 5) : entrance aperture situated mesally, in the pouch formed by the valves; bursa membranous, with proximolateral part lightly sclerotized. Bursae debouching through a pair of rigid tubes into the strongly sclerotized and pigmented, elongate spermathecae. The species is distinguished from H. sinuata by the much larger size, the longer legs and by differences in the epigyne and vulva.

~ ~ '-" .. :.- ...... , . .:.~ " . " . I I

Figs. 7,8 : Histopona egonpretneri n.sp 7) (j, holotype, right palp, lateral view; 8) id., ventral view. Fig. 9 : Histopona krivo­ sijana, C(, Strmena peCina, vulva, dorsal view. Scale-line 0.5 mm.

The specimens were collected from a colony living in the entrance chamber of Manita pee (alt. 400 m) in the twilight zone. They were sitting on their webs between blocks and boulders. This species is dedicated to the memory of our much regretted old friend Egon Pretner, the modest but great Yugoslav biospeleologist, who died during the preparation of this paper. The genus Histopona 331

HrSTOPONA SINUATA (Kulczyriskil

Tegenaria sinuata Kulczynski 1897, in : Chyzer & Kulczynski, Aran. Hung. II : 171, 9, fig., loco typo Rumania, Closani (jud. Mehedinti), Pestere. Histopona sinuata Lehtinen 1967, Ann. Zoo!' Fennici 4 : 239 ; Brignoli 1980, Ann. Zoo!' Warszawa 35 (7) : 78,9, fig. «syntype IZ PAN Warszawa, Pestere». Material examined: none. Distribution : only known from the type locality.

HISTOPONA VIGNAI Brigholi (figs. 10, 11, map 3)

Histopona sinuata: Brignoli 1977, Rev. Suisse Zoo!. 84 (4) : 950Jd'9, figs. Greece, Mt. Olympus; Larissa, Mt. Ossa. Histopona vignai n. sp. Brignoli 1980, Ann. Zoo!. Warszawa 35 (7) : 78. loco typo 1\11. Ossa, wrroundings of Karitsa. Material examined: 1c3 , holotype, 1':( ,paratype, Mt. Ossa, ColI. Brignoli. Male palp : fig. 11. The embolus leaves the radix on the distolateral side, its coiling direction is inversed near the base in a bizarre twist. Distribution: Greece, Mt. Ossa, Mt. Olympus.

\

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~." .~.' 17· ..

Map. 2. Distribution of the species of group myops • H. myops,. 0 H. bidens,. ... H. dubiiJ ,. .. H. krivosijana,. 0 H. trallteevi,. • H. hal/seri 332 Christa Deeleman-Rcinhold

, I I /

Figs. 10, II : Histopona vignai. 1O)d', holotype, eye region, dorsal view; 11) id., left palp, ventral view. Scale-lines 0.5 mm. Fig. 12 : Histopona dubia, 9 Mocilska pecina, vulva, dorsal view. Scale-line 0.5 mm.

HISTOPONA MYOPS (Simon) (map 2)

Hadites myops Simon 1885, Ann. Soc. Ent. France 6 (5) : 212,9, loc. typo Greece, Mt. Ossa, grotte de Kokkino Vracho. Roeweriana myops: Brignoli 1972, Rev. Suisse Zool. 79 (2) : 865, 9, fig. grotte de Kokkino Vracho. Material examined: Greece, provo Macedonia, Pellis, Flamouria, speleon Gremos, 19 ' 25. VII. 73, leg. P.R. + c.L. Deeleman (new locality). This cave is about 60 km to the north of the type locality. The genital organs agree in all details with the good figures presented by Brifnoli of the lectotype. • ,j j .

t~ o·····.·.·\

",,',f .. , . ~..

il .~.

", .. ' .... . ti. ·~V~~ .... ~: . . .. ,.... ~ :...... •...... ;) ··· .. ·:··:·:·:···· .' .' .. . '. ", .... " ...... ,f~l;~&d Q~\<, c<{" .. o.~.>.:. d(l···. " ·'·-n . .v .... :.- . .;!? .:-. '. QD;;'(r' . .'.:.".: f -...... '- Wi .·.eN..•...,. ..\) ~ ....n' ,,'~."'"..' "(J'. ···vi.1>

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Map. 3. Distribution of H. vignai, H. italica and of the ,Ipecic' or group s'trinatii .':'., ~,' * H. vignai,. • H. italica ,. ..'fl. strhatti,. "'H. iso/alII

HISTOPONA I>UBIA (Absolon & Kratochvil) (fig. 12, map 2)

Hadites dubius Absolon & Kratochvil 1932, Acta Mus, Morav.29: 5:d;<; fig, caves in Dalmatia and southernHercego~ina. Hadites (Roeweriana) dubius : Kratochvil I')38, Acta Soc. Sc. Nat. Morav. II (I) 101 : 16 <19 figs loc. typo SIPUIl spllJa, Cavtat. The genus Histopona 333

Material examined: Sipun spilja, 9, Mocilska pecina,09' Vulva: fig. 12. The bursa is lightly sclerotized and rigid. Distribution: southwestern Yugoslavia. Localities known: Hercegovina, caves bordering Popovo polje ; caves near Grebci. Southern Croatia, Cavtat, Sipun spilja ; Osojnik, Moc'ilska pecina. Southern Montenegro, Orjen, Vucja peCina near Vrbanje.

HISTOPONA BIDENS (Absolon & Kratochvil) (map 2)

Hadites bidens Absolon & Kratochvil 1932, Acta Mus. Morav. 29 : 6,d' 9,fig., caves on the isle of Mljet. Hadites (Roeweriana) bidens : Kratochvil 1938, Acta Soc. Sc. Nat. Morav. 11 (1) \01 : 12,d' 9, figs., loc. typo Isle of Mljet, cave Ostasevica near Babino polje. Material examined: none. Distribution: Yugoslavia, caves on the isle of Mljet.

HISTOPONA KRIVOSIJANA (Kratochvil') (fig. 9. map 2) Hadites bidens krivosijanus KratochvillS'35, Acta Soc. Sc. Nat. Morav. 9 (12) 88: 21,9, southern Yugoslavia, Krivo~ije. Hadites (Roeweriana) krivosijanus : Kn,tochvil 1938, Acta Soc. Sc. Nat. Morav. II (1) 101 : 21,9 ' figs., loc. typo southern Montenegro, Dragaljsko polje, cave Goltbova peCina near Selakov do. Material examined: 19, Krivosije, Strmena pecina near Selo Han, Leg. J. Kratochvil VII. 1969. Differences with H. dubia and H. bidens are enumerated by Kratochvil (1938). H. krivosijana is smaller than dubia. Vulva (fig. 9) : the bursae are different in shape, and they touch in the middle over a much larger area. The epigynal valves are separated sharply in the middle by a V-shaped chitinized margin. Thed'is still unknown. Distribution: southern Montenerro, Mount Krivosije.

HISTOPONA TRANTEEVI Deltshev (map 2)

Histopona tranteevi Deltshev 1978, Acta Zoo!. Bulgarica \0 : 57,9, figs., loco typo Bulgaria, western Rhodopes, cave Zmeini borun near Mostovo. Material examined: none. Distribution: only known from the type locality.

HISTOPONA HAUSERI (Brignolil (map 2)

Roeweriana hauseri Brignoli 1972, Rev. Suisse Zoo!. 79 (2) : 863, d' 9 ' figs., loc. typo Greece, isle of Corfu, superior cave of Bodolekos. Material examined: 2 juvs., Corfu, Gravulitha cave S. of Spartilas, 28.IX.1976, leg. E. Gittenberger. This species has female genitalia that are very similar to those of the other species of group myops ,. in the d',the cymbium and conductOJ" are very elongate. The eyes are normal in size and pigmentation. Distribution: Greece, islands of Corfu and Zante, in caves.

HISTOPONA STRINATII (Brignolil (Fig. 17. map 3)

Roeweriana strinatii Brignoli 1976, Rev. Suisse Zoo!. 83 (3) : 567,d'9, figs., loc. typo Greece, Peloponnesos, Ton Limnon cave, 20 km S. of Kalavrita. Material examined: 16, holotype, 19, paratype, ColI. Brignoli. Epigyne : fig. 17. the spermathec:ae are comparatively very large. Distribution: only known from the type locality. 334 Christa Deeleman-Reinhold

HISTOPONA ISOlATA n. sp. (figs. 13-16, map 3)

Locus typicus: Greece, island of Crete, cave Aghia Sofia near Topolia. Only known from the type locality. Material examined: 1 cJ , holotype, collected alive as a subadult on 17.III.1981, maturity moult 1981, killed 6.VI.l981 ; 16<;?paratypes 17 + 19.III.l981, leg. P.R. Deeleman. i Description: cJ : Carapace and chelicerae pale orange, sternum and ventral side of coxae white, legs ochre, abdomen creamy white, its dorsal side with a faint pattern consisting of a longitudinal white band followed by 4-5 chevrons. Legs uniform yellow to pale ochre, Eyes : size and pigmentation normal. Anterior medians about one third in diameter of the laterals ; posterior eye row straight or very slightly recurved. Chelicerae with 3 teeth in the anterior, 4 in the posterior row.

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15 ...... ;;

I I Figs. 13-/6 : Histopona iso/ata n. sp. 13)d', holotype, right palp, venl r:i1 view; 14) id., lateral view; 15) 9, paratype, epigyne ; 16) id., vulva, dorsal view. Scale-lines 0.5 mn. The genus Histopona 335

Measurements in mm : d', ho1otype, carapace 2.90 long, 2.16 wide, abdomen 3.35 long, Legs: I: 3.60 ­ 1.06 - 3.84 - 4.05 - 2.65 = 15.20; II : 3.60 - 1.05 - 3.35 - 3.85 - 2.40 = 14.25 ; III : 3.35 - 0.95 - 2.78 ­ 3.96 - 1.92 = 12.96; IV : 4.37 - 0.98 - 3.98 - 5.40 - 2.40 = 17.13 9, paratype, carapace 3.65 long, 2.65 wide, abdomen 5.60 long. Legs: I: 4.18 - 1.20 - 4.08 - 4.13 ­ 2.50 = 16.90; II : 3.84 - 1.00 - 3.35 - 3.95 - 2.25 = 14.09 ; III : 3.95 - 1.20 - 3.35 - 4.08 - 2.02 = 14.60 ; IV: 4.82 - 1.44 - 4.65 - 5.76 - 2.35 = 19.02; pa1p : 1.58 - 0.62 - 0.95 - 1.95 = 5.07

17 .1

\ Y

Fig. 17 : Histopona strinatii, 9. paratypl:, epigyne. Scale-line 0.5 mm. Figs. 18, 19 : Histopona italica. 18) 9, paratype, vulva, dorsal view; 19) id., ventral view. Arrows indicate probable route of penetrating embolus. Scale-lines 0.:5 mm. Figs. 20, 21 : Histopona palaeolithica. 20) 9, holotype, vulva, dorsal view; 21) id., ventral, view. Arrows indicate probable route of penetrating embolus. Scale-lines 0.5 mm. 336 Christa Deeleman-Reinhold

Male palp (figs. 13, 14) : femur with 2 dorsal spines, patella with 2, tibia with 3. Patella without pro­ jection, tibia with two external apophyses. Conductor an elongate, cup-shaped plate borne on a stalk, supporting the middle portion of the embolus at rest. Radix without apophysis. Basal part of embolus thick and distal part ribbonlike. Epigyne and vulva (figs. 15, 16) : relatively small, median margin of valves attached to the body wall, at some distance from the posterior margin; entrance aperture situated mesally. Rigid tubes folded in a quasi-helicoidal loop, spermathecae widely separated. The species resembles strinatii, but is larger and there are many differences in the genital organs of d and? The spiders were found in the anterior part of the cave only, at a place where boulders are strain accross the loamy floor. They inhabited small webs underneath and between boulders. Twelve other caves in the same area were searched without result. Like so many cave , they seem to be very selective in habitat preference and are found only very locally, in colonies.

HISTOPONA ITALICA Brignoli (figs. 18. 19. map 31

Histopona italica Brignoli 1976, Quad. Mus. Speleol. V. Rivera 4 : 35,d'I(' figs., loco typo Latium, Colli Alqani. Material examined: 1d, 1 9, pm'atypes, ColI. Brignoli. Vulva: figs. 18, 19. The'bursa has a central, unpaired and a lateral, paired part, which is lateraly lightly sclerotized. Distribution: northern and central Italy in epigean habitats.

HISTOPONA PALAEOLITHICA (Brignolil (figs. 20. 211

Cicurina (? Chorizomma) palaeolithica Brignoli 1971, Fragm. Ent. 8 (2) : 128.1(, fig., loc. typo Liguria, Cave' delle Arene Candide. Histopona palaeolithica : Brignoli 1976. Quad. Mus. Speleol. V. Rivera 4 : 38 Material examined: 19, holotype, ColI. Brignoli. Vulva: figs. 20 and 21. Distribution: only known from the type locality.

Summary

A diagnosis of the genus Histopona Th,)rell is presented. In the past, ambiguous definitions of the genus have been the origin of some taxonomic confusion. All species r itherto included in the genus Roeweriana Kratochvil are here transferred to Histopona. Male and female of H. exonpretneri n. ,po (cave in Mt. Velebit, Croatia) and isolata n.sp. (cave on Crete) are described and pictured, Histopona laeta (Kulczynski) md H. debilis Thorell are synonymized with H. torpida and H. laeta var. paganeuii (Nosek) with H. conveniens Kulczynski. It is argued that differen~es between Roeweriana and HislOpona were based mainly on reduction of eye size and pigmentation in the former. According to modern views loss .of eye, in many species appears to be a minor, fairly easily accomplished adal'tation with little phylogenetic weight and this character alone cannot be accepted as a ground for the separation of genera. The genus Hislopona here conceived comprises at present 16 species; 11 inhabit hypogean habitats in limited ranges in southeastern Europe; several are known from one cave only. Hist{lpona torpida is a purely epigean species, distributed over most of Europe, the other epigean species occ lpy smaller, partly o\"t>rlapping ranges. A group of closely related species in caves in the southwestern Balkan Peninsula (group myops) is troglobitic ; no c)dess species are known at present.

Resume

Cet article presente une definition du genre Hislopona Thorell. Des diagnoses ambigues par plusieurs auteurs ont ete la cause de confusions taxonomiques dans la liueratt,re. Toutes les especc, du genre Roeweriana Kratochvil sont ici transmises a Histopona ; Ie male et la femelle de H. egonpretneri fl.Sp. (groue au Ml. Velehit, Croatie) et de H. isolata n.sp. (grotte Aghia Sophia pres de Topolia, Crete) sont decrits. Histopona {,]eta (Kulczynski) et H. debilis Thorell sont placees en synonymie avec H. torpida (C.L. Koch) et H. laela var. paganettii (Nosek) avec H. conveniens Kulczynski. Le genre Roeweriana a ete cree en pr,:mier lieu pour recevoir des especes a fone reduction oculaire et depigmentation. La reduction oculaire cependant s'est avere~ n'etre qu'une adaptation mineure. realisee facilement et frequemment ; il est donc inacceptable de maintenir des genres fond~s uniquement sur ce caractere dans une classification phylogenetique. Les Roeweriana ne sont rien d'autre que des Hislopona adal'tees au milieu hypoge. The genus Histopona 337

Le genre Histopona, concu ainsi comprend 16 especes, toutes europeennes. Les 11 especes hypogees presentent differents degres d'adaptation au milieu hypoge et sont toutes limitees a des aires reduites en Europe du Sud-Est. Les deux especes nouvelles ne sont a present connues que d'une seule grotte chacune. Seull'espece H. torpida, epigee, est connue de presque toute l'Europe moyenne et meridionale sauf de la region mediterranee. Les autres especes epigees colonisent des aires restreintes a la peripherie de celie de torpida et en partie superposees a celle-ci. Un groupe d'especes troglobies etroitement apparentees est cantonne a la Peninsule Balkanique Occidentale; aucune espece anophthalme n'est connue apresent.

References

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