INTRODUCTION TO IBERIAN GROUNDWATER AMPHIPODS

J. Notenboom Institute of Taxonomic Zoology, University of Amsterdam, Post Box 4766, 1009 AT Amsterdam (Present adress: National Institute for Public Health and Environmental Protection Post Box 1, NL-3720 BA Bilthoven, The Net- herlands).

Keywords: Iberia, Ground water, Amphipoda, zoogeography

ABSTRACT Zoogeographical data on Iberian ground water dwelling amphipods are given. The geographical ranges, habi- tats, and co-occurrences of the genera, are discussed. The Iberian peninsula is divided into zoogeographical dis- tricts based on these amphipods; some are correlated with geological units.

INTRODUCTION Cantabrian Mountains), Metacrangonyx (from the only), and Niphargus (N. ciliatus Intensive sampling in continental during clsmontanus Margalef, 1952, from Guipuzcoa). the period 1983-1986 revealed many data about GINET published a short paper in 1977, including composition and distribution of ground water in- data from Portugal as well (reproduced in Spanish habiting fauna (stygofauna), especially crusta- in 1980), in which he stressed the scarce knowl- ceans. This paper will deal with the amphipods of edge about the distribution in Spain of the genera this fauna mainly on the generic level. Data about Niphargus, Haploginglymus, Pseudoniphargus other groups of crustaceans collected simulta- and Crangonyx. However, he did not mention the neously have been published by HENRY & MAG- genera Bogidiella, Salentinella, and Hadzia. RUF- NIEZ (1987, 1988), CAMACHO(1987), and PESCE& FO (1953) already reported an unnamed represen- GALASSI(1988). A survey of localities sampled dur- tative of Bogidiella near Sitges (S of Barcelona), ing our own investigation and on which most of while B. hellenae was described by MATEUS& DE the information provided in this paper is based is LOURDESM ACIEL (1967) from the mouth of the ri- given in fig. 1. ver Douro (Portugal). The genus Salentinella was First, a review is given of what has been publi- recorded for the first time by MARGALEF (1970a) shed before 1983 about Iberian ground water dwe- from a cave near Alcoy (Alicante). In the same lling amphipods. Rather few authors paid atten- paper Margalef mentioned the presence of Pseu- tion to this fauna and their findings were based on doniphargus in southern Spain, in the central very scattered data. A first treatise on crustaceans Pyrenees and in the Catalan hills. Other records from Iberian continental waters was published by of Pseudoniphargus in northern Spain were given MARGALEF (1953). It included the subterranean by ORTIZ (1968), VILLOTA& GALAN (1970), MAR- fauna as well and served as the starting point of GALEF (1970b), GOMARINGUIRADO (1978), and Es- this review. Stygobiont amphipods recorded by COLA (1980). MARGALEF(op. cit.) are: Pseudoniphargus (as P. In 1972, MATEUS& MATEUS described Hadzia africanus Chevreux, 1901, from cave waters in the tavaresi from wells near the southern Portuguese Limnetica, 6: 165-176 (1990) coast (Algarve); STOCK (1977) established a new O Asociación Española de Limnología. Madrid, Spain genus, Metahadzia, for this species and also men- tioned some new localities. MATEUS& MATEUS Nacional de Ciencias Naturales. Their material (1978) provided up to date distributions of the ge- contained only Bogidiella (STOCK & NOTENBOOM, nera Bogidiella, Haploginglymus, Pseudoniphar- 1988). In this paper, bogidiellids from eastern and gus and Metahadzia in Portugal. In 1980a STOCK southern Spain are recorded as well. Our own in- described a second species of Haploginglymus vestigations have been concentrated on the Can- from a hyporheic habitat in the Cantabrian Moun- tabrian Mts. and the western part of the Pyrenees, tains (northern Spain) and published in the same the central-eastern part of Spain, and year a revision of the genus Pseudoniphargus, in (see fig. 1). In al1 areas visited it was attempted which five new species were recognized from the to sample a range of biotopes, such as the under- Iberian peninsula. In his revision STOCK (1980b), flow of rivers, springs, wells, and cave waters. also incorporated new material from northern and Data on the localities investigated in 1983 and southern Spain. Recently, KARAMAN(1986a,b) re- 1984 have been published by NOTENBOOM& MEI- ported on Niphargus and Haploginglymus based JERS (1985). Additional sampling has been execu- on the study of the older collections of Ginet and ted in summer 1985 by Peter van den Hurk and of Mateus, and established the existence of a third Remko Leys in the western part of Andalusia, and Haploginglymus species in central Iberia. A fourth by Ine Meijers and Jos Notenboom in southern form of Haploginglymus coexisting with Niphar- and the eastern part of Andalusia, and gus was recently discovered a spring south of the also in summer 1986 by Remko Leys and Katja Catalan Pyrenees (PRETUS & SABATER, in press.). Hogedoorn in Gallicia, and by Peter van den The area around Madrid has been investigated Hurk, Marc Koperdraat, and Jos Notenboom in during the last years by colleagues of the Museo Catalonia.

Figure 1.- Distribution of al1 groundwater samples taken during our campaigns of 1983-86. Squares indicate 100x100 km UTM- grid. Closed symbols indicate the situation of two or more samples taken within the same 10x10 km UTM-square. Distribución de las muestras de aguas subterráneas tomadas durante las campanas de muestre0 realizadas entre 1983 y 1986. La cuadrícula corresponde a las coordenadas UTM de 100x100 km. Los símbolos negros indican la situación de dos o más muestras tomadas dentro del mismo cuadrado de lOx 10 km. THE IBERIAN GROUNDWATER AMPHIPOD Niphargus region. Most likely the present distri- GENERA bution of Niphargus in Iberia is the result of dis- persa1 from the area north of the Pyrenees, Niphargus Schiodte, 1849 (fig. 2) through the low altitude corridors west and east This genus has an extended distribution; it stret- of the mountain ridge. This hypothesis is confir- ches from western Europe into the Near East (KA- med by the presence of trans-Pyrenean species, N. RAMAN & RUFFO,1986). In Spain it reaches its delamarei in the east, and N. ciliatus and N. lon- southwestern distribution limit and occurs in two gicaudatus in the west (MARGALEF,1952, 1970a,b; segregated areas on both edges of the Pyrenean KARAMAN, 1986b). mountain ridge only. In the eastern Catalan Pyre- Niphargus is a widely distributed stygobiont nees the genus is mentioned by GINET (1977) from amphipod genus with a high species diversity, in- La Mosquera cave near Beuda (prov. Gerona), cluding species with large ranges. Supposedly, one identified by KARAMAN(1986b) as N. delamarei of the reasons for its wide range is a rather high Ruffo, 1954. Our own investigations in Catalonia dispersa1 ability related with its capacity to colo- showed that Niphargus ranges from the Pyrenees nize not only the phreatic leve1 but also the un- into the (about 150 km saturated zone and saturated layers situated above south of Pyrenees). In the area west of the Pyre- the regional water table, whereby lower mountain nees (Basque Country and the northwestern part ridges can be crossed. In this context it is interest- of the ), the genus has been ing to mention some biotopes from where Ni- found frequently both in the past and during our phargus was recorded: in the Basque Country, for own investigations. This is the classical Iberian instance, in dripwater pools of a mine gallery, si-

Figure 2.- The distribution of Haploginglymus and Niphargus in the Iberian peninsula. Squares indicate 100x100 km UTM-gnd. Symbols with stack-bars indicate the situation of two or more localities of the genus within the same 10x10 km UTM-square. Distribución de Haploginglymus y Niphargus en la península Ibérica. La cuadrícula corresponde a las coordenadas UTM de 100~100 km. Los símbolos marcados con una línea indican la situación de dos o más localidades del género dentro del mismo cuadrado de coordenadas UTM de 10x10 km. tuated in non-carbonate rocks at 600 m altitude, stygobiont amphipod. It is uncertain if Haplo- without contact with the local groundwater table, ginglymus occurs in the central part of Iberia, sin- and in a spring from red sandstone, situated high ce few investigations have been done there. Until up a hill at 320 m altitude and with a pH of the now four species are recognized (KARAMAN, water of 4.0. This spring is being fed most likely 1986a; PRETU~& SABATER, in press), but since by a perched groundwater aquifer. In Catalonia more material awaits identification, it is to be ex- most Niphargus records are from wells and hypor- pected that more forms will be recognized. heic habitats, but it was found also in seeps of a The genus inhabits particular wells and under- railway tunnel. flows of rivers, but it has been found in springs as In Basque Country Niphargus has been found well. Only in the mid-eastern part of the peninsu- in caves, springs, and wells, frequently together la are cave-dwellers recorded, possibly because of with Pseudoniphargus, very few times with Salen- the absence of Pseudoniphargus in this region. tinella, and only once with Haploginglymus. In Haploginglymus is closey related to Niphargus Catalonia the only amphipod co-occurring with (cf. MATEUS& MATEUS,1958; BARNARD & BAR- Niphargus was Haploginglymus, in particular NARD, 1983; KARAMAN,1986a; KARAMAN& RUF- along the border of the range. FO, 1986). The observation that these two genera occur together only along the borders of their dis- Haploginglymus Mateus & Mateus, 1958 (fig. 2) tribution areas would suggest that there is some kind competition between them. The genus is fre- This genus, strictly endemic of the Iberian pe- quently found together with other amphipod ge- ninsula, is the most widely distributed Iberian nera.

Figure 3.- The distribution of Salentinella in the Iberian peninsula. See also legend of fig. 2. Distribución de Salentinella en la península Ibérica. Véase también la leyenda de la figura 2 Salentinella Ruffo, 1947 (fig. 3) species have restricted distributions often confined to a single hydrographic basin, including the up- The genus is confined to fresh or sligthly brac- permost parts of Atlantic drainage systems. kish habitats of the western peri-Mediterranean region (RUFFO, 1986). In Spain Salentinella occurs Bogidiella Hertzog, 1933 (fig. 4) in al1 regions prospected by us, not only close to the board, but also in inland Species of Bogidiella were found in coastal ground waters and even in the vicinity of the Bay ground waters, but also in continental ground wa- of . They are principally inhabitants of wells ters. A part of the material available was studied and underflows, although MARGALEF (1970a) re- by STOCK & NOTENBOOM (1988). They recognized corded them also from a cave. Frequently Salenti- five different species al1 endemic to Iberia belon- nella was found together with other stygobiont ging to the subgenera Bogidiella S.S.and Medigi- amphipods, especially with Haploginglymus. diella. Cases of sympatry within Bogidiella, and PLATVOET(1987) studied the Spanish material between Bogidiella and Medigidiella are found in and recognized nine different forms, belonging to Iberian inland water (STOCK & NOTENBOOM, op. eight different (sub)species and one incertae se- cit.). In spite of intensive investigations, Bogidie- dis. The most widely distributed species is S. an- lla has not been found in the northern part of the gelieri, whose distribution coincides with that of central plateau (Meseta), the Cantabrian Moun- the entire genus, and it occurs in various habitats tains, the western Pyrenees, and inland waters of near the Mediterranean coast, including insular Catalonia. habitats. In Spain it has been found up to some Above al1 Bogidiella species are inhabitants of 100 km from the coast. The remaining Spanish hyporheic habitats, although occasionally also

Figure 4.- The distnbution of Rhipidogammarus, Bogidiella and Metahadzia in the Iberian peninsula. See also legend of fig. 2. Distribución de Rhipidogammarus, Bogidiella y Metahadzia en la península Ibérica. Véase también la leyenda de la figura 2. found in wells. Frequently other stygobiont am- (1971) mentioned R. rhipidiophorus from an epi- phipods occured in the same habitats, viz., Pseudo- gean pool in a riverbed at Gerona. Our own in- niphargus, Haploginglymus, and Salentinella. vestigations yielded a new species, R. triumvir, from wells in Almería (NOTENBOOM, 1985, 1988a). Metahadzia Stock, 1977 (fig. 4) Sensonator Notenboom, 1986 (fig. 5) The genus Metahadzia has mainly a Mediterra- nean distribution. In the Iberian peninsula it was This monotypic endemic genus was discovered formerly known by a single species, M. tavaresi, in the southern part of the province of Valencia. from wells of the Algarve. Our own investigations It is a rather large amphipod with remarkable cha- revealed the existence of a second species, M. un- racteristics for a stygobiont, such as many calceoli cispina, in wells in the basin of the Quadalquivir on both antennae in both sexes, and a subfolia- river near Sevilla (NOTENBOOM, 1988b). Probably ceous, natatory third uropod. The phylogenetic the distribution in Spain of Metahadzia is limited position of the genus is uncertain (NOTENBOOM, to this river basin. The co-occurring amphipod 1986a). fauna existed of Salentinella. S. valentiensis was found in a phreatic cave lake (near Corbira de Alcira), but also in wells (near Rhipidogammarus Stock, 1971 (fig. 4) and ) and in an hyporheic habitat of the Río ; al1 these localities are situated A small number of localities with representati- within 30 km from the sea shore in a rather res- ves of this genus are known from continental tricted area. It has been found together with Ha- Spain, restricted to regions close to the Mediterra- ploginglymus, Salentinella, and Bogidiella. S. va- nean sea board in fresh to brackish waters. STOCK lentiensis is a bottom dweller but also free-swim-

Figure 5.- The distnbution of Crangonyx, Sensonator and Parapseudoniphargus in the Ibenan peninsula. See also legend of fig. 2. Distribución de Crangonyx, Sensonator y Parapseudoniphargus en la península Ibérica. Véase también la leyenda de la figura 2. ming, and is supposed to be very sensitive to wa- compact with deep coxa1 plates and relatively ter-borne pressure waves, whether produced by short posterior pereiopods and uropods, sugges- animal vibrations or other disturbances in the wa- ting special adaptation to interstitial life (NOTEN- ter. Probably it is a predator which has its most fa- BOOM, 1988~).Parapseudoniphargus is distributed vourable habitat in large waterfilled fissures of the in hyporheic habitats of tributaries and upper saturated karst. courses of the Guadalquivir river, and occured to- gether with Pseudoniphargus, Haploginglymus, Crangonyx Bate, 1859 (fig. 5) and Salentinella.

During our investigations species of this genus Pseudoniphargus Chevreux, 1901 (fig. 6) have not been discovered. Previously the genus was mentioned by GINET (1977) from two caves in The genus has a western Mediterranean- the north of Spain (Basque Country and Navarra). amphi-Atlantic distributional range and the Iberian In spite of intensive sampling, also in one of the peninsula occupies a central position in it. In Ibe- caves mentioned by Ginet (Cueva de Mairuelego- ria Pseudoniphargus has a disjunct distribution, retta), it was not found again. with three ranges: northern Spain, southern Spain, and Lusitania. Al1 these three ranges have Parapseudoniphargus Notenboom, 1988 (fig. 5) their own assemblages of endemic species. Stri- king is the absence of Pseudoniphargus in the mid- Our investigations revealed the existence of this eastern part of the peninsula. An ecological cause monotypic genus closely related to Pseudoniphar- of this pattern seems unlikely because many other gus and probably endemic to the Guadalquivir ri- stygobiont crustaceans such as Haploginglymus, ver basin. The body of this amphipod is rather Salentinella, Proasellus, Stenasellus, and Ibero-

Figure 6.- The distnbution of Pseudoniphargus in the Ibenan peninsula. See also legend of fig. 2. Distribución de Pseudoniphargus en la península Ibérica. Véase también la leyenda de la figura 2. bathynella are abundant in this mid-eastern re- morphological divergence is larger than in north- gion, especially in hyporheic habitats, while Pseu- ern Spain. Most of the southern species are allo- doniphargus in the other areas investigated often patric with quite point shaped distributions. In the co-occurred with these elements. Guadalquivir basin, however, the widely distribu- Karst waters provide important habitats for ted P. latipes co-occurred with the very localized Pseudoniphargus in northern Spain, in addition, species, P. illustris. Other amphipods frequently representatives are found in hyporheic waters, found together with Pseudoniphargus were Haplo- springs, and a few wells. Altogether 13 species ginglymus and Bogidiella, botb mainly in hypor- have been recognized in the area, most of them heic habitats, and Salentinella, only in wells (No- with rather small allopatric ranges, limited to a TENBOOM, 1987a). single drainage basin or a karst area. In the Bas- Lusitania harbours a few species from Portugal que Country Pseudoniphargus frequently co- and a single one from the northwesternmost part occurred with Niphargus. In hyporheic habitats, of Spain. The three species of Portugal are limi- in particularly of the area around the Picos de Eu- ted to the central-western part of the country (No- ropa, Haploginglymus was often found together TENBOOM, 1987b). with Pseudoniphargus (NOTENBOOM,1986b). The presence of Pseudoniphargus in the Catalan hills (MARGALEF, 1970b) remains enigmatic, our own investigations in that area in 1986 did not reveal Zoogeographical districts its presence. Subterranean waters of southern Spain appa- Based on the known distributional data of rently support an important diversity of Pseudo- stygobiont amphipods an attempt is made to divi- niphargus species. In the 15 species recognized de to Iberian peninsula into areas characterized

Figure 7.- Zoogeographic distncts of the Iberian peninsula based on presence or absence of stygobiont amphipods. Distritos zoogeográficos de la península Ibérica según la presencia o ausencia de anfípodos estigobiontes. by presence or absence of certain taxa. This ap- by the presence of various species of Pseudoni- proach is presented in fig. 7, but it needs empha- phargus, each with its own assemblage of endernic size that not al1 Iberian regions are equally tho- species. The eastern border of Cantabricum could roughly investigated (see fig. 1). not be established with certainty since it remains The stygobiont amphipod districts tend to be si- uncertain if Pseudoniphargus occurs in inland wa- tuated at the periphery of the Iberian Meseta. The ters of the region southeast of the Pyrenees. The Meseta district apparently has the lowest diversity border between Oriente and Meseta is the west- with Bogidiella and Haploginglymus only . The lat- ern limit of the range of Salentinella. A Vasco- ter genus is not particularly bound to any district, Pyreneo-Catalonicum district is distinguished due but an endemic for the entire peninsula. Regions to the penetration of Niphargus in Iberian ground with particular cases of endemism are the Guadal- waters. In northern Spain the ranges of the gene- quivir and Levante with the endemic genera, Pa- ra Pseudoniphargus, Niphargus and Salentinella rapseudoniphargus and Sensonator, respectively. are partially overlapping, the eastern zone of Can- The unnamed taxon close to Salentinella, mentio- tabricum is inhabited by Salentinella, and the north- ned by PLATVOET(1987), might be an endemic of ern part of that zone by Niphargus. Levante, too. Moreover, the Guadalquivir district In Iberia predominantly Mediterranean ele- has endemic species of Pseudoniphargus and Sa- ments like Metahadzia and Salentinella are found lentinella, and is the only Iberian region with re- in Atlan'tic drainage basins of Guadalquivir, presentatives of Metahadzia. Cantabricum, Lusi- Oriente and Cantabricum. The latter genus is re- tanicum, and Beticum are districts distinguished corded as well from the adjacent Aquitanian area

Figure 8.- Major geotectonic units of the Ibenan peninsula. 1) Hercynian basement; 2) deformed Mesozoid cover of the Pyrenean realm; 3) Mesozoic aulocogenic area; 4) Mesozoic externa1 units of the Betic realm; 5) intermediate Flysch units of the Gibraltar arc; 6) interna1 units (Mesozoic and Paleozoic) of the Betic-Rif realm; 7) undeformed Mesozoic cover; 8) continental and marine. Principales unidades geotectónicas de la península Ibérica. 1) Zócalo hercínico; 2) cubierta mesozoica deformada del dominio pi- renaico; 3) área ulocogénica mesozoica; 4) unidades mesozoicas externas del dominio bético; 5) unidades intermedias del Flysch del arco de Gibraltar; 6) unidades internas (Mosozoico y Paleozoico) del dominio Bético-Rif; 7) cubierta mesozoica indeformada; 8) continental y marino.

GOMARINGUIRADO , F., 1978. Biospeleología de «La Cuevo- NOTENBOOM, J., 1987a. Species of the genus Pseudoniphargus na». Cuadernos de Espeleología 3: 22. Santander. Chevreux, 1901 (Amphipoda) from the Betic Cordillera of HENRY, J.P. & G. MAGNIEZ, 1987. Isopodes aselloides stygo- southern Spain. Bijdr. Dierk. 57 (1): 87-150. bies d'Espagne recoltes par J. Notenboom et 1. Meijers. 1 NOTENBOOM, J., 1987b. Lusitanian species of the amphipod Le genre Synasellus et description de trois nouvelles espe- Pseudoniphargus Chevreux, 1901 with a key to al1 Iberian ces. Stygologia 3 (4): 331-344. species. Bijdr. Dierk. 57 (2): 191-206. HENRY, J.P. & G. MAGNIEZ, (1988). Isopodes aselloi'des stygo- NOTENBOOM, J., 1988a. Biogeographical observations on the bies d'Espagne récoltes par J. Notenboom et 1. Meijers. 11 genera of Iberian stygobiont Amphipoda. Crustaceana, - Le genre Bragasellus et de description de sept nouvelles es- Suppl. 13: 122-133. peces. Stygologia 4 (4). NOTENBOOM, J., 1988b. Metahadzia uncispina, a new amphi- KARAMAN, G.S., 1986a. Description of Haploginglymus ma- pod from phreatic groundwaters of the Guadalquivir nver teusi, new species of subterranean Gammandea from Ibe- basin of southern Spain. Bijdr. Dierk. 58 (1): 79-87. rian peninsula with remarks to other taxa of this genus (Fam. NOTENBOOM, J., 1988c. Parapseudoniphargus baetis, new ge- Niphargidae). Poljoprivreda I Summarstvo 32 (1): 75-90. nus, new species, a stygobiont amphipod crustacean from KARAMAN, G.S., 1986b. Discovery of Niphargus delamarei the Guadalquivir River Basin (southern Spain), with phylo- Ruffo 1954 in Spain, with first descnption of females (Gam- genetic implications. J. Crust. Biol. 8 (1): 110-121. maridea: Niphargidae). Poljoprivreda I Summarstvo 33 NOTENBOOM, J. & 1. MEIJERS, 1985. Investigaciones sobre la (213): 29-42. fauna de las aguas subterráneas de España: lista de estacio- KARAMAN, G.S. & S. RUFFO, 1986. Amphipoda: Niphargus- nes y primeros resultados. Versl. techn. Gegevem (Inst. group (Niphargidae sensu Bousfield, 1982). In: L. BOTOSA- taxon. zool., Univ. Amsterdam) 42: 1-93. NEANU (ed.), Stygofauna Mundi: 564-566, E. J. Brill, Lei- ORTIZ, E., 1968. Algunos crustáceos y miriápodos caverníco- den. las. ~uadernosde Espeleología 3: 85-87. Santander. MARGALEF, R., 1952. La vida en las aguas dulces de los alre- PESCE, G.L. & D. GALASSI (in press). Groundwater crusta- dedores del Santuario de Nuestra Señora de Aránzazu (Gui- ceans of Spain 11: Microparasellidae of Spain. Stygologia 4 púzcoa). Munibe 4: 73-108. San Sebastian. (4). MARGALEF, R., 1953. Los crustáceos de las aguas continenta- PLATVOET, D., 1987. The genus Salentinella Ruffo, 1947 (Crus- les Ibéricas. Biología de las aguas continentales, X: 1-243. tacea, Amphipoda) in Spain. Stygologia 3 (3): 217-240. Ministerio de Agricultura, Inst. Forestal de Inv. Exp. Ma- RUFFO,S., 1953. Anfipodi di acque interstiziali raccolti da1 Dr drid. C. Delamare Debouteville in Francia, Spagna e Algena. Vie MARGALEF, R., 1970a. Anfípodos recolectados en aguas sub- et Milieu 4 (4): 669-681. terráneas ibéricas. Speleon 17: 63-65. RUFFO, S., 1986. Amphipoda: Salentinellidae. In: L. BOTOSA- MARGALEF, R., 1970b. Anfípodos recolectados en aguas sub- NEANU (ed.), Stygofauna Mundi: 564-566. E. J. Brill, Lei- terráneas del País Vasco. Munibe. 22 (314): 169-174. San Se- den. bastián. STOCK, J.H., 1971. A revision of the Sarathrogammarus-group MATEUS, A. & M. DE LOURDESM ACIEL, 1967. Descnption (Crustacea, Amphipoda). Bijdr. Dierk. 41 (2): 94-129. d'une nouvelle espece de Bogidiella (Crustacea, Amphipo- STOCK, J.H., 1977. The taxonomy and zoogeography of the da) du psammon du Portugal et quelques notes sur son gen- hadziid Amphipoda. Stud. Fauna Curacao 55: 1-130. re. Publ. Imt. «Dr. A. Nobre» (Fac. Ciencias Porto) 59: STOCK, J.H., 1980a. Découverte, dans le nord-ouest de I'Es- 1-15. pagne, d'une deuxikme espece d'Haploginglymus, genre de MATEUS, A. & E. MATEUS,1958. Un nouveau genre et une Crustacés Amphipodes endémiques de la Peninsule Ibéri- nouvelle espece d'Amphipode troglobie du Portugal . Publ. que. Bull. zool. Mus. Univ. Amsterdam 7 (14): 141-147. Inst. «Dr. A. Nobre* (Fac. Ciencias Porto) 59: 1-15. STOCK, J.H., 1980b. Regression model evolution as exempli- MATEUS,A. & E. MATEUS, 1972. Une nouvelle espece d'Had- fied by the genus Pseudoniphargus (Amphipoda). Bijdr. zia (Crustacea, Amphipoda) du Portugal. Publ. Inst. .Dr. Dierk. 50 (1): 105-144. A. Nobre* (Fac. Ciencias Porto) 117: 1-30. STOCK, J.H., 1986b. Amphipoda: Gammarid grouping (Gam- MATEUS,A. & E. MATEUS, 1978. Amphipoda hypogés du Por- maridae s. str. sensu Bousfield) In: L. BOTOSANEANU(ed.), tugal. Publ. Inst. «Dr. A. Nobre» (Fac. Ciencias Porto) 142: Stygofauna Mundi: 504-513. E. J. Bnll, Leiden. 1-26. STOCK, J.H., 1986~.Amphipoda: Melitid grouping (Melitidae NOTENBOOM, J., 1985. Rhipidogammarus triumvir n. sp. sensu Bousfield, 1973, emend.). In: L. BOTOSANEANU(ed.), (Amphipoda, Gammaridae) from wells near Mojonera, Al- Stygofauna Mundi: 504-513. E. J. Brill, Leiden. meria. Stygologia 1 (3): 292-299. STOCK, J.H. & J. NOTENBOOM, 1988. Five new bogidiellid NOTENBOOM, J., 1986a. Sensonator valentiensis n.g., n.sp. Amphipoda from Spain - the first freshwater records in the (Amphipoda), from different biotopes in southern Valencia. Iberian Peninsula. Hydrobiologia 164: 75-95. Bijdr. Dierk. 56 (1): 60-74. VEGAS, R. & E. BANDA, 1982. Tectonic framework and Alpi- NOTENBOOM, J., 1986b. The species of the genus Pseudoni- ne evolution of the Iberian peninsula. Earth Evolution Scien- phargus Chevreux, 1901 (Amphipoda) from northern Spain. ces 4: 320-343. Bijdr. Dierk. 56 (1): 75-122. VILLOTA, J. & C. GALAN, 1970. Complejo Lerzo-Anudia II- Sabe Saia-Ko-Lezia. Munibe 22 (314): 175-182. San Sebas- tián.