ISOPODA: CIROLANIDAE L. BOTOSANEANU*, N. BRUCE** &J. NOTENBOOM*

The Cirolanidae is one of the two families of Isopoda Flabel- The Cirolanidae are a predominant marine family with lifera which include stygobiont elements (the other family about 300 species presently known (the validity of some ma­ being Sphaeromatidae). In the classification here adopted rine and some stygobiont species is open to question). In the the genera including — or comprising exclusively — tabellar part of this chapter, 45 species and subspecies are stygobionts are placed in the general frame of the numbered, but a few of them are doubtful Western- predominantly or exclusively marine (non-subterranean) Mediterranean Typhlocirolana species. Certainly more genera. This classification is as follows (genera represented stygobiont as well as marine species will be described, and it in groundwater printed in italics). is possible to say that almost 1/7 of all the Cirolanidae have found their way to the Stygal. If genera are considered, the proportion is even more impressive: from some 40 presently EURYDICINAE recognized genera, 17 are stygobiont or include also stygobi­ onts or stygophiles (only Annina and Anopsilana are presently Eurydice group: Eurydice. recognized as including both marine and subterranean spe­ Pseudaega group: Annina, Eurylana, Excirolana, Pon- cies; but this may prove to be true also for Cirolana: see notes togelos, Pseudaega, Pseudolana. for Anopsilana and Creaseriella); of course, this is partly a result Colopisthus group: Arubolana, Colopisthus, Metaciro- of the tendency to describe monotypic genera of stygobiont lana. Cirolanidae — a tendency which, in our opinion, is a justi­ fied one in this specific case. UNNAMED SUBFAMILY One species was excluded from the tabular part: Conilera stygia Packard, 1894, from Monterey, Nuevo Leon, Mexico: Conilera group: Conilera, Conilorpheus, Natatola- this is unanimously considered as extremely poorly na, Orphelana, Politolana. described and unrecognizable (will probably prove to be­ long in Speocirolana). Two species which are obviously non- CIROLANINAE stygobionts are nevertheless included in the tables, because they have clear relations to the subterranean habitats: Anni­ Bathynomus group: Bathynomus, Parabathynomus. na lacustris and Saharolana seurati (see "notes"). There are no Cirolana group: Anopsilana, Cirolana, Creaseriella, other problems with the delimitation of stygobiont and non- Hansenolana, Haptolana, Neociro- stygobiont Cirolanidae. lana, Saharolana. The subterranean Cirolanidae show a remarkable variety Gnatholana group: Cartetolana, Gnatholana. of body shapes, and the illustration of this chapter gives a Sphaeromides group: Antrolana, Bahalana, Cirolanides, good idea of this situation. Not less impressive are the differ­ Mexilana, Oncilorpheus, Speocirolana, ences in size (the figures which follow are maximal total Sphaeromides, Turcolana, Typhlocirolana. lengths in published accounts). Smaller than 5 mm are: Faucheria group: Faucheria, Skotobaena, Sphaerolana. Arubolana parvioculata (with 2.8-2.9 mm the smallest of all), A. aruboides, Saharolana, Turcolana, Faucheria; the group between Incertae sedis. 5 and 10 mm consists ofArubolana imula, most of Anopsilana, Bahalana cardiopus, most of Typhlocirolana; the following range Ceratolana between 10 and 15 mm: Annina, Anopsilana poissont, Hap­ tolana, Bahalana geracei, Mexilana, Speocirolana thermydronis, Sphaeromides polateni, and two N. African Typhlocirolana; be­ 'Institute of Taxonomic Zoology, University of Amsterdam, tween 15 and 24 mm: Creaseriella, Antrolana, Cirolanides, Plantage Middenlaan 64, 1018 DH Amsterdam, The Nether­ Speocirolana pubens, Sphaeromides bureschi bureschi and S. v lands. montenigrina, Skotobaena, and Sphaerolana; largest (24 mm "Australian Museum, 6-8 College Street, Sydney, N.S.W. 2000 Australia. more) are most of Sphaeromides and of Speocirolana, Speocffl' ISOPODA: CIROLANIDAE 413 bolivari being, with 33 mm, the giant of the group. The fe­ the subterranean realm (in the specific case of this group: males are often larger than the males. Almost all the stygobi- mainly a karstic aquatic environment) become available for onts are anophtalmous (Arubolana parvioculata is micro- colonization by marine immigrants? The peculiar aspects of oculate, as is the stygophile Saharolana) and devoid of pig­ the origin (time, place, possible ancestors...) of the different ment in the teguments; but it must be kept in mind that subterranean taxa are evoked in rather many publications, several true marine cirolanids are equally devoid of eyes a Pleistocene, Tertiary, Cretaceous, Jurassic or even and/or depigmented (examples: Cirolana lata, Natatolana Paleozoic origin being postulated. Possibly the most difficult californiensis, N. caeca, Metacirolana caeca...). case is that oi Antrolana lira, found in an area unexposed to Important characters used in the classification of the (sub­ marine waters since the early Permian; for this species (but terranean) cirolanids and in assessing the relationships in­ also, for instance, for Cirolamdes texensis, for some Western- side the family, are: the pleon segmentation (Bowman, mediterranean Typhlocirolana...) it was assumed, for different 1975), the morphology of the antennule- and antennal reasons, that they are possibly derived from a marine stock peduncles, the shape of the three first pairs of pereiopods, through the intermediacy of a freshwater epigean ancestor. the segmentation and setation of the pleopods, the insertion Rather little is known about the biology, autecology and of appendix masculina on the endopodite of pleopod II O", behaviour of these animals. A few useful references'. Nouris- and uropod characteristics. Extremely interesting is the fact son, M., 1956 (Typhlocirolana buxtoni); Por, F., 1962: Typh­ that 7 species belonging to five genera (in what seems to be locirolana reichi n.sp., un nouvel isopode Cirolanidae de la 3 different lineages) are capable of rolling into a ball when Depression de la Mer Morte. Crustaceana, 4: 247-252; disturbed, this implying of course more or less drastic Sket, B., 1964 (Sphaeromides virei virei); Delhez, F, 1966: modifications in the morphology of body and appendages; Recherches ecologiques sur un crustace troglobie Sphaero­ in Greaseriella, Turcolana and Faucheria, the volvation is mides raymondiDollfus. Ann. Speleol., XXI, 4: 838-844; Ber- described as perfect; it is very imperfect in Skotobaena mortoni trand, J.Y., 1974: Recherches sur lecologie de Faucheria and somewhat less imperfect in S. monodi; volvation is also faucheri. These doctorat 3C cycle, Univ. Paris VI, not print­ reported in the two species of Sphaerolana (no details availa­ ed; Collins, XL. & Holsiger, J.R. 1981: Population ecology ble); the phenomenon — absent in all non-subterranean of the troglobitic isopod crustacean Antrolana lira Bowman. Cirolanidae! — is considered by some authors as possible Proc. 8th Int. Congr. Speleol., 1: 129-132; Carpenter, J.H., defensive strategy, useful in the conditions of the subterra­ 1981 (Bahalana geracei). Most species are reported as normal­ nean realm. ly crawling on the substrata, never swimming or swimming In the geographical distribution it is possible to distin­ only when disturbed (Arubolana imula, Antrolana, Bahalana ger­ guish an important peri-ponto-mediterranean grouping acei, Speocirolana pelaezi, Sphaeromides raymondi, Typhlocirolana (with a western-mediterranean, a balkanic and a levantine reichi, Faucheria, Skotobaena mortoni, Sphaerolana), whereas sub-groupings), an even more important peri-caribbean swimming actively in open waters is the normal mode of grouping (Mexico having more species than any other part movement for some other species (Arubolana aruboides, Anop- of the world), and a small eastern-african grouping. Two silana crenaia, Speocirolana thermydronis). An absolute indiffer­ species cannot be included in any of these groupings: An- ence towards light or absence of light has been occasionally trolana lira (in the Appalachians) and Arubolana aruboides (on noted (Sphaeromides raymondi), and also the fact that the two Bermuda). Several clusters of species are clearly distin­ species of Sphaerolana are active mostly during the night. guished, this clustering having three different aspects con­ Although the mouthparts of most cirolanids seem to indi­ cerning affinities, distribution, inhabited biotopes. Many cate predatory or scavenging habits, there is no definite indi­ species are presently restricted in their distribution to locali­ cation supporting this (concerning Arubolana aruboides, ties very near to, or not far from marine or oceanic shores; Sphaeromides virei, Bahalana geracei, it is supposed that they out this is not valid for a series of other species; to name some could predate on living prey, and cannibalism was observed taxa found at 200 km or more from the nearest present ma­ in this last species); most species are more or less opportunis­ rine shores: Antrolana (ca. 200), Speocirolana thermydronis (> tic detritivorous (Arubolana imula: rotting wood; Bahalana ger­ 450, Sphaeromides bureschi (ca. 260), S. polateni (ca. 230), Typh­ acei: scavenging on a variety of animals; Cirolanides: organic locirolana fontis (ca. 750 at Fort Miribel), Sphaerolana ( > 450). material, like guano; Speocirolana guerrai: guano?; Sphaero­ The stygobiont and stygophile Cirolanidae being obvi­ mides raymondi: clay, animal remainders...). Concerning the ously derived from marine forms (this is unanimously reproduction, it is interesting to note that ovigerous females recognized), the distance separating their localities from or females with brood plates or pouches, were apparently Present shorelines immediately strikes the imagination in never found in the subterranean species {this was expressely ne context of the question of their origin. But this is certain- noted, for instance, for Antrolana, Bahalana, some Typh­ y not the essential aspect, far more important questions be- locirolana...); this phenomenon still awaits explanation (one ng: when, in the geological past, were the areas wherefrom published explanation being that ovigerous 99 are very ygobionts are known covered by marine waters? When did rare and secretive, rarely foraging in areas accessible to sam- 414 L. Botosaneanu, N. Bruce &J. Notenboom pling); evidence of ovoviviparity was found in (epigean)^n- KEY REFERENCES nina lacustris from the Comoro Islands. Three confirmed cases of co-existence of stygobiont species in the same locali­ Angelov, A., 1968: Sphaeromides polateni ein neuer Vertreter der ty are known: Speocirolana bolivari and S. pelaezi coexist in the Hohlenfauna Bulgariens. — Bull. Inst. Zool. et Musee (Sofia) XXVII: 195-213. Grutas de Quintero and in the Bee Cave (Tamaulipas) as Argano, R. & Pesce, G.L., 1980: A cirolanid from subterranean well as in the spring at La Laja (San Luis Potosi ); waters of Turkey. — Rev. Suisse Zool., 87, 2: 439-444. Speocirolana thermydronu, Sphaerolana affims, and Sphaerolana in- Bowman, Th. E., 1964: Antrolana lira a new genus and species of troglobitic cirolanid isopod from Madison Cave, Virginia terstitialis, are found together in some of their localities near Int. J. Speleol, 1(1 + 2): 229-236, pi. 50-57. Cuatro Cienegas; Typhlocirolana leptura coexists in some —, 1966: Haptolana trichostoma, a new genus and species of wells in Morocco with an unidentified species of the same ge­ troglobitic cirolanid isopod from Cuba. — Int. J. Speleol., II- 105-108, pi. 24-27. nus. —, 1975: A new genus and species of troglobitic cirolanid isopod In a large majority of cases, the stygobiont Cirolanidae from San Luis Potosi, Mexico. — Occ. papers Mus. Texas Tech. Univ., 27: 1-7. are clearly associated with karstic habitats, sometimes in­ —, 1981: Speocirolana pubens and S. endeca, new troglobitic iso­ habiting karstic waters near, or not far from marine shores pod crustaceans from Mexico. — Assoc. Mexican Cave Stud and more or less influenced by the vicinity of the sea, some­ Bull. 8: 13-23. times karstic waters which are far inland on the islands or — & Franz, R., 1982: Anopsilana crenata, a new troglobitic cirolanid isopod from Grand Cayman Island, Caribbean Sea continents, where all influence of the sea has been excluded. — Proc. Biol. Soc. Wash., 95 (3): 522-529. The richest habitats are the saturated zones in cave systems, — & Iliffe, T.M., 1983: Bermudalana aruboides, a new genus and and the karstic springs associated with them (underground species of troglobitic isopoda (Cirolanidae) from marine caves on Bermuda. — Proc. Biol. Soc. Wash., 96 (2): 291-300. streams are seldom inhabited, pools in the vadose zone of Carpenter, J.H., 1981: Bahalana geracei n.gen., n.sp., a troglobitic caves possibly never). Species definitely associated with in­ marine cirolanid isopod from Lighthouse Cave, San Salvador terstitial habitats are far more rare, and, unfortunately, in­ Island, Bahamas. — Bijdr. Dierk., 51 (2): 259-267. Contreras-Balderas, S. & Purata-Velarde, D.C., 1982: Speoci­ formation is sometimes scanty and misleading; Arubolana rolana guerrai sp.nov., Cirolanido troglobio anoptalmo de la parvioculata is very definitely associated with an interstitial Cueva de la Chorrera, Linares, Nuevo Leon, Mexico. — As­ habitat; such an association could also be made for Anop- soc. Mexican Cave Stud. Bull. 8: 1-12. Ferrara, F. & Lanza, B., 1978: Skotobaena monodi, espece nouvelle silana acanthura, Cirolanides (only in some of its localities!), de Cirolanide phreatobie de la Somalie. — Mon. Zool. Ital., Turcolana, some maghrebine Typhlocirolana, and the two N.S. Suppl. X, n°6: 105-112. levantine species of this genus. Herbst, G.N., 1982: New records of Typhlocirolana species (Crustacea: Isopoda) from Israel: ecological and biogeographi- An extremely vast gradient of water salinity is tolerated by cal significance. — Isr. J. Zool., 31: 47-53. the different subterranean cirolanids, from perfectly fresh Monod, Th., 1930: Contribution a l'etude des "Cirolanidae". — Ann. des Sc. Nat., Zool., 10l serie: 129-183. water to water having salinity near that of sea water. In fresh —, 1972: Contribution a l'etude de la Grotte de Sof Omar (Ethiopie water (upper limit of saline concentration: ca. 2 g/1) we may meridionale). N°.3 — Sur une espece nouvelle de Cirolanide mention: Anopsilana cubensis, A. radicicola, Haptolana, An- cavernicole, Skotobaena mortoni. — Ann. Speleol., 27, 1: 205-220. trolana, Speocirolana pelaezi, all the species of Sphaeromides, Tur­ —, 1976: Remarques sur quelques Cirolanides. — Bull. Mus. colana, Typhlocirolana moraguesi, Faucheria. In slightly brackish natn. Hist, nat, 3° serie, n°, 358: 133-161. water, often fresh to taste: Anopsilana acanthura, A. crenata, Notenboom, J., 1981: Some new hypogean Cirolanid isopod crusta­ Creasenella, Skotobaena monodi. Bahalana cardiopus was found in ceans from Haiti and Mayaguana (Bahamas). — Bijdr. Dierk., 51 (2): 313-331. strongly brackish water (9.68 g/1), and B. geracei in very —, 1984: Arubolana parvioculata n.sp. from the interstitial of an strong brackish to water with full marine salinity (21.26-35 intermittent river in Jamaica, with notes on A. imula g/1). Some species are known as tolerating more or less im­ Botosaneanu & Stock and A. aruboides (Bowman & Iliffe). Bijdr. Dierk, 54, 1: 51-65. portant variations of the salinity: Arubolana parvioculata Nourisson, M. 1956: Etude morphologique comparative et cri­ (4.85-26.5 g/1), Annina (from slightly brackish to sea water), tique des Typhlocirolana (Crustaces Isopodes Cirolanidae) du Arubolana imula (1.34-3.78 g/1, at different times), Typh­ Maroc et d'Algerie. — Bull. Soc. Sci. naturelles et physiques Maroc, XXXVI (2e trim.): 104-124. locirolana reichi 2.52-8.64 g/1). The two species of Sphaerolana Racovitza, E.G., 1912: Cirolanides (premiere serie). —Biospeolog- live in very strongly mineralized water. ica XXVII, Arch. Zool. exper. et gen., 5C serie, X: 203-329, pi- Concerning endangered habitats and species, special XV-XXVIII. Redell, J.R., 1981: A review of the cavernicole fauna of Mexico, mention has to be made of the catastrophic modifications Guatemala, and Belize. — Texas Mem. Mus. Bull. 27: 84-88. undergone these last years and decades by the habitats of Rioja, E., 1953: Estudios carcinologicos XXX. Observaciones Speocirolana thermydronis, Sphaerolana affinis and Sphaerolana in- sobre los Cirolanidos cavernicolas de Mexico. — Annls. Inst. Biol. (Mexico), XXIV, 1: 147-170. terstitialis, in the vicinity of Cuatro Cienegas de Carranza, — 1956: Estudios carcinologicos XXXV. Datos sobre algunos northern Mexico (one of the most extraordinary zones of the Isopodos cavernicolas de la Isla de Cuba. — Annls. Inst. Bio • globe for fresh water fauna, subject of systematic and savage (Mexico), XXVII: 437-462. Sket, B., 1964: Genus Sphaeromides Dollfus 1897 (Crust., Isopoda, destruction). Cirolanidae) in Jugoslawien. — Biol. Vestnik (Ljubljana), *• 153-168. ISOPODA: CIROLANIDAE

Strouhal, H., 1963: Sphaeromides bureschi eine neue Hohlenwas- serassel aus Bulgarien. — Bull. Inst. Zool. ct Musee (Sofia), XIII: 157-175.

FIGURES

1: Annina lacustns (from Gordon & Monod, 1968); 2: Arubolana imula (from Botosaneanu & Stock, 1979); 3-4: Anopsilana poissoni, lateral and dorsal (from Monod, 1976); 5: Creaseriella anops (from Creaser, 1936); 6: Haptolana trichostoma (from Bowman, 1966); 7: Antrolana lira (from Bowman, 1964); 8: Bahalana geracei (from Carpenter, 1981); 9: Cirolanides texensis (from Bowman, 1964); 10: Mexilana saluposi (from Bowman, 1975); 11: Speocirolana pubens (from Bowman, 1981); 12: Sphaeromides virei virei (from Sket, 1964); 13: Turcolana cariae (from Argano & Pesce, 1980); 14: Typhlocirolana moraguesi (from Racovitza, 1912); 15: Typhlocirolana steinitzi (from Strouhal, 1960); 16-17: Faucheriajaucheri, dorsal and in volvation, lateral (from Racovitza, 1912); 18: Skotobaenamonodi (from Ferrara& Lanza, 1978); 19-20: Skotobae- na mortoni, in volvation, lateral and frontal views (from Monod, 1972). L. Botosaneanu, N. Bruce &J. Notenboom ISOPODA: CIROLANIDAE 417 418 L. Botosaneanu, N. Bruce &J. Notenboom

Cirolanidae Harger, 1880

Eurydicinae Stebbing, 1905

Gr. Pseudaega

Annina Budde-Lund, 1908

lacustris IV 2: Shimoni, S. E. corner of Kenya D?,G?,a Budde-Lund, 1908 IV 6: Zanzibar (also Comoro islands) Also epigean!

Gr. Colopisthus

Arubolana Botosaneanu & Stock, 1979

imula VII 4: Mangel Cora Tunnel, Aruba D-I Botosaneanu & Stock, 1979

aruboides VIII 10: Church cave and Wonderland cave, Bermuda (Bowman & Iliffe, 1983)

parvioculata VII 9: dry bed of Rio Secco, Discovery Bay, Jamaica L-Rl Notenboom, 1984

Cirolaninae Harger, 1880

Gr. Cirolana

Anopsilana Paulian & Delamare Deboutteville, 1956

acanthura VII 8: well at Marigot, southern coast of Dept. de K (Notenbofjm, 1981) l'Ouest, Haiti

crenata VII 11: West Bay Cave, NW end of Grand Cayman Island G Bowman & Franz, 1982

cubensis VII 10-11: several freshwater caves in the Cuban provinces (Hay, 1903) Pinar del Rio, Habana, Matanzas, as well as on Isla D, E de Pinos

poissom IV 5: Grotte de Mitoho, South-Western Madagascar D; a Paulian & Delamare Deboutteville, 1956

radicicola VII 8: "Source Debarasse", large karstic spring flowing from T, A (Notenboom, 1981) cave near Jeremie, southern peninsula of Haiti

Creaseriella Rioja, 1953

10 anops VII 2: many caves (some of them true cenotes), states D, E;I? (Creaser, 1936) Quintana Roo & Yucatan of the Yucatan Peninsula. One well in Campeche, same peninsula ISOPODA: CIROLANIDAE 419

Haptolana Bowman, 1966 t 11 trichostoma VII 10: several freshwater caves, Sierra de Cubita, province D Bowman, 1966 Camaguey, Cuba

Saharolana Monod, 1930

12 seurati I 13-IV 1: epigean brook fed by springs at Kebili, near Chott Monod, 1930 Djerid, Tunisia

Gr. Sphaeromides

Antrolana Bowman, 1964

13 lira VIII 1: Madison Saltpetre Cave and Stegers Fissure, near D Bowman, 1964 Grottoes, Augusta Co., Virginia, USA

Bahalana Carpenter, 1981

14 cardiopus VII 12: Mount Misery Cave, Little Bay, Mayaguana Island, G? Notenboom, 1981 Bahamas

15 geracei VII 12: Dixon Hill Lighthouse Cave, San Salvador Island, G Carpenter, 1981 Bahamas

Cirolanides Benedict, 1896

16 texensis VIII 7a: caves and phreatic waters of south central Texas, D, I Benedict, 1896 from the vicinity of Del Rio in the west to San Marcos in the east (± southern limit of Edwards Plateau)

Mexilana Bowman, 1975

17 saluposi VII 1: Cueva del Huisache, 4 km NW Micos, San Luis Potosi A Bowman, 1975

Speocirolana Bolivar y Pieltain, 1950

18 bolivari VII 1: one cave in the Sierra de El Abra, one in the Sierra de D, T (Rioja, 1953) Guatemala (both Tamaulipas), and one spring at La Laja, San Luis Potosi

19 endeca VII 1: Sotano de las Calenturas, Yerbabuena, and Cueva del A Bowman, 1981 Tecolote, Los San Pedro (both NW of Ciudad Victoria, Tamaulipas)

20 guerrai VII 1: Cueva de la Chorrera, Linares, Nuevo Leon D Contreras-Balderas & Purata-Velarde, 1982

21 pelaezi VII 1: many caves in the states Puebla, San Luis Potosi, D; B (Bolivar y Pieltain, and Tamaulipas 1950)

22 pubens VII 1: Cueva de la Bonita, San Nicolas de los Montes, S. Luis Bowman, 1981 Potosi; Cueva del Ojo de Agua de Manantiales, 14 km of Ocampo, Tamaulipas 420 L. Botosaneanu, N. Bruce &J. Notenboom

23 thermydronis VII 1: Posos de la Becerra and one more poso, SW of Cuatro Cole & Minckley, 1966 Cienegas, Coahuila

Sphaeromides Dollfus, 1897

24 bureschi I 9a: 3 caves (near Iskretz, Cerowo, monastery Iwan Matnij) C, T Strouhal, 1963 and one karstic spring (by Opizwet), all in westernmost part of Stara Planina Mts., Bulgaria

25 bureschi serbica 1 9a: karstic spring Gradiste, ca. 10 km NW from Dimitrovgrad T; A? Pljakic, 1968 (in SE Serbia but quite near the Bulgarian border). Possibly also Peterlas Cave (same zone)

26 polateni I 9a: cave in Polaten, not far from Teteven, central part of C, T Angelov, 1968 Stara Planina Mts., Bulgaria

27 raymondi I 4: subterranean river of La Dragonniere, Vallon, Ardeche; D, C, T Dollfus, 1897 Grotte des Cent-Fonts, Grotte de Ressecs, and Grotte de TAvencas (Herault)

28 virei virei I 7b: 4 caves and 2 wells in Istria D, I, L3; (Brian, 1923) I 7c: one cave at Vrana, central Dalmatia, one cave at Ston, probably southern Dalmatia. All localities very near the coast also: C, T

29 virei I 7c: 2 caves (Milica Pecina and Kusaca), near Zegar, Middle A mediodalmatina Dalmatia. Rather far from coast Sket, 1964

30 virei I 7f: Obodska Pecina (karstic spring of Rijeka Crnojevica), T; certainly montenigrina near northern end of Scutari-lake, Jugoslavia also D-C Sket, 1957

Turcolana Argano & Pesce, 1980

31 cariae I 11: freshwater well between Ula and Koyecegiz, SW Anatolia Argano & Pesce, 1980

32 rhodica I 8e: springbrook Gadouras, 1.5 km. W from Apolona LI Botosaneanu, Boutin (Rhodos); interstitial water in peebles & Henry, 1985

Typhlocirolana Racovitza, 1905

33 buxtoni I 13: "Grotte du ravin de Bou Jacor", and artesian well at D, Kl Racovitza, 1912 Bredea, both near Bou Tlelis, wilaya Oran, Algeria

34 fontis I 13: several groups of springs near Biskra and in the wilayas T, S, a (Gurney, 1908) Oran, Sidi-Bel-Abbes, and Constantine (old provinces of Oran and Constantine); wells near Algiers. IV 1: wells at Hassi Chebaba (Fort Miribel) 135 km S from El Golea, and at Colomb-Bechar, both Algeria

35 gurney i I 13: "Grotte de la 4eme source du ravin de Misserghin", D Racovitza, 1912 wilaya Oran, Algeria (quoted also from "Grotte du ravin de Bou Jacor": see buxtoni)

36 leptura I 13: wells in Morocco (2 km E from Chichaoua, 10 km S and K Botosaneanu, 4 km N from Marrakech) Boutin & Henry, 1985 ISOPODA: CIROLANIDAE 421

37 moraguesi I 2: caves and wells on the islands Mallorca, Menorca, and D, I; K?K1? Racovitza, 1905 Dragonera. I 5c: "pozzi artesiani salmastri", Sicily

38 reichi I 12: two springs and one "water hole", Arava Valley, S (Z), a Por, 1962 S. of the Dead Sea; one well at Be'er Aharon, Negev; all in Israel

39 rifana I 13: "aguas freaticas", Monte Arruit, Melilla (Morocco) Margalef, 1958

40 steinitzi I 12: well ("forage") at Kfar Atah (Haifa, Israel) Strouhal, 1960

Gr. Faucheria

Faucheria Dollfus & Vire, 1905

41 faucheri I 4: wells in limestones and vertical caves reaching the water of C, I, D, T (Dollfus & Vire, 1900) underground river, near Sauve, Dept. Gard; Grotte et resurgence des Cent-Fonds (Herault). Also cave (s) in Dept. de TAude ? All in France

Skotobaena Ferrara & Monod, 1972

42 mortom IV 2: Cave of Sof Omar (on river Webbe, Prov. Bale, Ethiopia) C; probably Monod, 1972 also D

43 monodi IV 2: shallow wells at El Gambole, Southern Somalia a (probably I) Ferrara & Lanza, 1978

Sphaerolana Cole & Minckley, 1972

44 affinis VII 1: various "pozos" ("springfed wells") near Cuatro S (Z), a; D? Cole & Minckley, 1970 Cienegas de Carranza; flooded mine "about 30 m. below entrance of Cueva de la Boca, near Villa Santiago", Nuevo Leon

45 interstitialis VII 1: small springs and "pozos" near Cuatro Cienegas de S (Z), a Cole & Minckley, 1970 Carranza, Coahuila 422 L. Botosaneanu, N. Bruce &J. Notenboom

NOTES Margalef 1958). That fontis, buxtoni, and gurneyi represent only one species, is almost beyond doubt; but is this species 1: not stygobiont, but certainly stygophilous; this oculate distinct from momguesiP The status oirifana as species distinct and pigmented species was recorded from "Seewassertiim- from moraguesi is also rather uncertain. It seemed, neverthe­ peln" inside Zanzibar, from mangrove habitats, but also less, more reasonable to keep for the time being all these as from wells, and especially from the "partly subterranean independent taxa in the tabellar part, until a badly needed lakes" Machumwi Ndongo and Machumwi Kumbwa on revision of western-mediterranean Typhlocirolana is available Zanzibar (these being parts of a groundwater table accessi­ (only T. lulli Pujiula, 1911, described from Cuevas dels ble through caves with collapsed roof), and from the, appar­ Hams, Mallorca, and almost certainly synonymous with ently similar, cave Shimoni (SE Kenya). moraguesi, was excluded). It is possible that such a revision 2: in partly artificial gallery dug in coral limestones near will show that we are concerned here with an unique species sea-shore. (moraguesi) — perhaps showing some geographical variabili­ 3: described as Bermudalana; in large bodies of salt water ty. Mention should be also made of the following facts: a) two with characteristics of anchihaline habitats, deep inside populations of Typhlocirolana from S. Morocco are studied by caves ca. 250-420 m. from next marine shore. Nourisson, 1956, as Typhlocirolana sp.sp.; b) in recent years Anopsilana: only very slightly distinct from the marine abundant material still awaiting study, was colledted in Al­ Cirolana, and considered as possibly paraphyletic genus. geria, Morocco, and in Spain (the description of a n.sp. from Troglocirolana Rioja, 1956, and Haitilana Notenboom, 1981, peninsular Spain is presently in press: inf. L. Pretus). are considered as synonyms. An Anopsilana sp. was men­ 38: found also in the wet mud filling a "water hole" after tioned (as Haitilana sp.) from a well near Thomazeau, Dept. drought. de l'Ouest, Haiti. 42: in a small cave brooklet, sometimes dry; can leave the Creaseriella: apparently only slightly differing from the water to climb on wet mud; supposedly also living in "fentes marine Cirolana. noyees ou d'autres cavites ... humides". Locality remote 12: not stygobiont, but possibly stygophilous. This oculate from any marine coast. (very small eyes) and pigmented species is known from a 43: in wells dug in limestones, but reaching "une nappe habitat well described by Monod, 1930: a sluggish fresh­ aquifere tres superficielle", ca. 120 km from the Indian water streamlet fed by a nearby spring complex. Ocean coast. 14: in a "muddy hole filled with water (probably the water table)". 15: in cave pools with characteristics of anchihaline habitats, but in a cave ca. 1 km from the Ocean. ADDITIONAL NOTES 21: according to Bolivar y Pieltain, 1950, in the first disco­ vered locality, Cueva de los Sabinos, lives in "un pozo situa- Haptolana. A second species — ascribed with some do en la galeria superior de la caverna .... nunca pudo ser en- doubts by the authors to this genus — was described from contrada en las lagunetas de la galeria inferior de la caver­ Somalia (Messana, G. & Chelazzi, L., 1984: Haptolana na" (rather surprising situation). somala n.sp., a phreatobic cirolanid isopod (Crustacea) 23: in a complex and peculiar habitat: interstices in blocks from the Nogal Valley (northern Somalia). — Monitore of porous travertine along sides of natural wells (posos) fed Zool. ital., N.S. Suppl. XIX (9): 291-298). by the water of thermal springs. The localities are Gibaganle, Callis, and Eil, along the 27: Dr. R. Rouch (Moulis) has kindly provided some un­ Wadi Nogal; IV 2; apparently true phreatic water: K. published information concerning the distribution of this Typhlocirolana. With regard especially to the notes species. 33-35, 37, and 39: it is quite possible that we were too scepti­ 28: S. virei was initially placed (by Valle-as nomen nu­ cal concerning the distinctness and validity of the various dum-, then by Brian) in the genus Troglaega, now no longer Western-mediterranean species; see comments on this pro­ recognized. The specimens of virei virei from Istria and from blem in the publication mentioned below. Dalmatia are slightly different. An addition to the key bibliography: Botosaneanu, L., 33-35, 37, and 39: the systematic problems related with Boutin, C, Henry, J.P., 1985: Deux remarquables the western-mediterranean Typhlocirolana are complex and Cirolanides stygobies nouveaux du Maroc et de Rhode . difficult, and there is yet no definite solution for them (see, problematique des genres Typhlocirolana Racovitza, l"u for instance: Racovitza 1912, Monod 1930, Nourisson 1956, et Turcolana Arcano & Pesce, 1980. — Stygologia, 1 (2)-