The Echinogammarus berilloni-group, a number of

predominantly Iberian Amphipod species (Crustacea)

by

Sjouk Pinkster

Institute of Taxonomie Zoology, University of Amsterdam, The Netherlands

Abstract INTRODUCTION

The morphological diversity within the species Echino- the last 10 15 of staff- During or years a group gammarus berilloni (Catta, 1878), stressed by Spandl members and students the Instituut (1926), Margalef (1956) and Pinkster (1969) was a motive at voor

in of the to sample systematically drainage systems Taxonomische Zoölogie of the University of Am- Iberian peninsula and of great parts of France. It became sterdam, coached by the enthusiasm of Prof. Dr. clear that E. berilloni in reality was the collective name J. H. has been on both for of least 10 all of them with their Stock, working systemati- a group at species, cal and ecological problems of fresh and brackish own, restricted distribution area. Only one of these

species, the real E. berilloni, inhabits a much larger area, water As member of this amphipods. a young including the major part of France, the Channel islands, team, I collected some gammarids in the western Belgium, Luxembourg, the southern part of the Nether- Pyrenees in 1968. While studying this material it lands, and south-western Germany. In this study the became clear, that E. berilloni (Catta) was not characters of the E. berilloni-group are discussed and variable comparative descriptions are given of all species, along only a very species, as already noticed by

with a key for their identification. The variability and Spandl (1926) and Margalef (1956), but in reality

ecological preferences of each species are discussed. a collective name for a group of closely related An attempt has been made to relate the distribution species, each variable in itself. This resulted in the of this and of other in the pattern group species groups description of a new E. aquilifer Pinkster, Iberian peninsula (e.g. the E. simoni-group, members of species,

the G. and the and likewise it the start for pulex-group, genus Eulimnogammarus) to 1969, was a more

geological data. intensive study of the subject.

One of the fertile methods solve most to system-

Résumé atic when of problems, especially a group

variable is is La variabilité morphologique de l’espèce Echinogam- species involved, to sample syiemati- and in the inhabited marus berilloni (Catta), remarquée par Spandl (1926), caJly intensively area by the

Margalef (1956) et Pinkster (1969) a mené à un échantil- group in question. In doing so variability withinand lonnage systématique dans les systèmes d’eaux de la between various populations of one species, as well dans la France. Il Péninsule ibérique et presque toute a as differences become clear. E. berilloni était réalité le nom collectif interspecific may été prouvé que en

d’au moins 10 dont chacune a has been done, pour un groupe espèces, Consequently systematic sampling son aire de distribution limitée. Il n’y a qu’une seule de in the years 1969, 1970 and 1971, in all drainage le vrai E. aire beau- ces espèces, berilloni, qui occupe une of the Iberian systems peninsula (see map I) and la de coup plus étendue, comprenant plus grande partie in many parts of France (Pinkster et al., 1970; la France, les îles de la Manche, la Belgique, Luxem- Pinkster & Stock, bourg, la partie méridionale des Pays-Bas, et la partie 1971). Since sud-ouest de l’Allemagne. morphological characters are changing cette note les caractères des Echinogammarus du Dans during the development, and consequently, various berilloni discutés des com- groupe sont et descriptions stages of different species often are very much paratives de toutes les espèces sont données, ainsi qu’une alike (see Dennert et al., 1969), sexually mature clef de détermination. En outre la variabilité et les pré- be used In obtain férences écologiques de chaque espèce sont indiquées. specimens can only. order to

modèle de distribution du Enfin l’auteur propose un more information about the ecology of the dif- des de la Péninsule groupe et autres groupes d’espèces ferent members the of group, analyses of water le les membres du ibérique (p. ex. groupe E. simoni, samples were made at nearly every station for the G. ex et le Eulimnogammarus) par rap- groupe pul genre factors temperature, pH, chlorinity and calcium port aux données géologiques.

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ion concentration. Moreover, the width and depth tion to the authorities in their countries. The author is

much indebted to Prof. Dr. J. H. Stock of the of the streams inhabited, bottom type, vegetation, Institute of Taxonomie Zoology (Zoölogisch Museum), Amster- and degree of pollution were noted. dam (ZMA) for his kind assistance during part of the As a result of this systematic inventarisation, it fieldwork and for his critical remarks during all phases became clear that the name E. berilloni, as used of the work.

to in up 1969 (see Pinkster, 1969), reality is the

collective noun for a group of at least 10 closely THE CHARACTERS OF THE ECHINOGAM-

related, but distinct forms. Nine of these species MARUS BERILLONI-GROUP

are restricted to rather small areas in the northern

half of the Iberian of Stock and Pinkster Stock peninsula or the adjacent part (1968) & (1972) sum-

France. Only one species, E. berilloni s.str. has a marized the most important characters discrimi-

the much wider distribution, inhabiting the middle nating Echinogammarus pungens-group and

reaches of streams and rivers in France, the Chan- the E. simoni-group from other representatives

nel Islands, Belgium, Luxembourg, the southern within the genus Echinogammarus. In this paper

treated which part of the Netherlands and south-western Ger- species are are all closely related to,

or confused with, berilloni many. In Spain the species seems to be restricted Echinogammarus (Cat-

rather and defined the to a narrow zone along the north-west ta), are by following characters:

coast. (1) the eyes are large and elongate; (2) the hands

of the In spite of our intensive sampling, no members first gnathopods are much smaller than

of those of the all armed with the E. berilloni-group were found in the second; (3) legs are

of sometimes southern part of the Iberian peninsula. In this area groups setae, accompanied by some smaller they seem to be completely replaced by members spines; (4) the carpus of the second leg is of the African E. simoni-group (Pinkster & Stock, always much shorter than the propodus; (5) the

1972). segments of both metasome and urosome are

In this of armed with and often paper complete descriptions all mem- spines groups of, very long,

bers of for their calceoli the first the E. berilloni-group and a key setae; (6) are always absent; (7)

of the mandible is identification are given. The distribution of the segment palp always unarmed,

the third bears comb- various species is discussed and illustrated in maps, segment always a regular

and where possible, notes are given on the varia- like row of setae. These characters also occur in

and the of the different the females, although less pronounced than in the bility ecological range males. in E. calvus species. In a final chapter, the distribution of all Although some species (e.g.

and E. characters and less groups of amphipods inhabiting inland waters in echinosetosus) (3) (5) are

the Iberian and than in most other members of this peninsula its zoogeographical as- pronounced

and tend to become intermediate between pects are discussed. group

this group and the Echinogammarus pungens-

ACKNOWLEDGEMENTS group, I still believe, after weighing the various

characters against each other that the members of The author is indebted to Dr. R. Margalef of the Instituto the E. berilloni-group can be distinguished from de Investigaciones Pesqueras for the loan of type material the other within the Echino- of Echinogammarus klaptoczi ebusitanus; also for the species-groups genus

loan of of the material to Dr. R. W. Ingle British Mu- gammarus. seum (Natural History) (BMNH), London; to Dr. L. B. Considering the characters cited above, Echino-

Holthuis of the Rijksmuseum van Natuurlijke Historie & Ruffo, gammarus pacaudi (Hubault 1956) (RMNH), Leiden; to Dr. S. Ruffo of the Museo Civico but should be placed within this group of species, di Storia naturale (MCSN), Verona; and to Dr. J. Nies- the dorsal ornamentation of the metasome, very sen, of the Zoologisch Museum Utrecht (ZMU). To Mrs.

I. Pinkster-de Graaf and Miss A. M. C. Goedmakers, B. characteristic by its sharp, teeth-like projections, Sc., for their assistance during part of the fieldwork; to other makes this species so different from all Mr. R. Dijkema, B. Sc., for drawing the map with the that I species within the genus Echinogammarus drainage systems; to Dr. O. J. Simons, of the Geologisch it When to as a group. Instituut of the University of Amsterdam, for his advice prefer separate special of E. ebusitanus in geological problems. reading the description klaptoczi

The fieldwork was made of the whether this possible through grants Margalef, 1951, one hesitates as to Netherlands' Organisation for the Advancement of Pure form should be placed within the E. berilloni- Research (Z.W.O.) and of the University of Amsterdam. Examination of the material group or not. type Special thanks are also due to the Embassies of Spain the to this The taxon and Portugal in the Netherlands for their kind introduc- provided answer problem.

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ebusitanus cannot be included in the berilloni- urosome segments.

group because of the long, elongate carpus of the 5) The number of setae on the legs; in general

this number second leg, a character shared by the genus increases with age.

and the Echino Eulimnogammarus gammarus KEY TO THE SPECIES OF THE ECHINO- simoni-group. GAMMARUS BERILLONI-GROUP, BASED

THE VARIABILITY OF CHARACTERS ON ADULT MALES ONLY.

The idea of berilloni original Echinogammarus 1 a) Peduncle segments 4 and 5 and flagellum of A2

with curved (Catta) being an extremely variable species does provided long, slightly setae, forming a flag-like brush. Hand of both first and second not hold after the present study. Of course, each without medial gnathopods palmar spines . . . . of the 10 species in which the old „berilloni” is Echinogammarus meridionalis n. sp.

subdivided, is variable in some characters quite b) These characters not combined 2 but in others. and this is not Moreover, probably 2 a) Flagellum of A2 provided with long setae forming a

flag-like brush 3 the greatest difficulty in this group, the characters b) Flagellum of A2 armed with solitary setae only, in the may change course of the development and never forming a flag-like brush 4 growth. So the most characteristic features are 3 a) Dorsal surface of metasome and urosome armed found in the setation of the antennae, the the legs, with numerous hooked spines. Hand of first and

medial metasome, urosome and epimeral plates. However, second gnathopods armed with a palmar Live with vertical dark stripes the number of setae increase spine. specimens . . and length with age Echinogammarus zebrinus Pinkster & Stock. 1971 und become fully characteristic only when the b) Dorsal surface of metasome and urosome armed

animal is full-grown. are often Younger stages with small tufts of short setae only. Hand of second hard The holds for very to distinguish. same true gnathopod without medial palmar spine. No vertical

females of all species in the E. berilloni-group, not stripes Echinogammarus aquilifer Pinkster, 1969 only because of the absence of discriminating 4 a) Setae on the peduncle segments of Al and A2 longer setae, but likewise because of the characteristic than the diameter of these segments

on the hands of the spines gnathopods. Echinogammarus feminatus n. sp.

all Therefore, discussions concerning the specific b) Setae on the peduncle segments of Al and A2

shorter than, or as long as, the diameter of these characters and their variability are valid for adult segments 5 males only. Metasome and covered with clusters of 5 a) urosome

Stable characters are the of the the shape eyes, long, curved setae with small spines in between setation of the second antenna, the absence of them. Bpimeres with long setae along the ventral

and well the lateral calceoli, the structure of the mandible palp, the posterior margins, as as on surface presence or absence of medial palmar spines on Echinogammarus berilloni (Catta. 1878) the hands of the gnathopods, the morphology of b) Metasome armed with small spines only, urosome

the the of the inner ramus of the third with few tufts of setae. with setae legs, shape a Epimeres along

and the absence the ventral and .... 6 uropod, presence or of groups of posterior margins only

set of setae. 6 a) PI to P7 with many groups very long setae on the dorsal surface of the telson, on the

In P5 to P7 these setae are always much longer metasome and urosome segments (the number and which than the diameter of the segments on they are of these setae is less length stable, however). Urosome with of implanted. segments some groups

Furthermore can be observed in 7 variability very long setae

The number of b) on and P7 short, in P5 to P7 always 1) segments in the flagellum of the Setae PI

shorter than the diameter of the segments on which antennae. they are implanted. Urosome segments with short 2) The shape of the epimeral plates of the pleon; setae 8

the of the corners can angle ventroposterior 7 a) The setae implanted along the external margin of also vary largely (see Roux, 1967; Stock, 1968; the exopod of Ur3, are many times longer than the

the hand Pinkster, 1970). diameter of the exopod. The setae on of the gnathopods are very long 3) The number of spines participating in the Echinogammarus longisetosus n. sp. of the telson. In armature some populations b) The setae implanted along the external margin of

subbasal spines are in others present, they are the exopod of Ur3, are about as long as the

in diameter of the The setae on the hand of absent. Even a single specimen differences exopod.

the gnathopods are short can be found between the contralateral telson

Echinogammarus tarragonensis n. sp. lobes (see also Stock, 1968; Pinkster, 1970). with the 8 a) Epimeres 2 and 3 very long setae along The of 4) number on the metasome and spines lower margin; their posterior margin and lateral

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surface unarmed. Telson lobes elongate, more than

1.5 times as wide, their dorsal as long surface never

armed with setae

Echinogammarus calvus (Margalef, 1956) new rank.

b) Epimeres 2 and 3 with small groups of short setae

both along the ventral and posterior margins and

often on the lateral surface. Telson lobes short, less

than 1.5 times as long as wide, their dorsal surface

with of short ... 9 one or more groups setae

9 a) A large species, adult male 18 mm. Hand of second

gnathopod large, with a swollen lower portion. Fla-

of with than gellum Al very long, normally more

35 segments. Setae on legs, telson, and dorsal sur-

face metasome of and urosome short, almost spini-

form. Setae along the lower margin of the third

segment of the mandib'e palp implanted in a comb-

like with way some longer setae in between them;

inner surface of the second segment without setae.

echinosetosus Echinogammarus n. sp

small b) A species, adult male 10—11 mm. Hand of Fig. 1. Echinogammarus berilloni (Catta, 1878), habitus second gnathopod rather small. Flagellum of Al of a male from the neotype locality (scale I). short, always less than 20 segments. Second segment

of mandible palp with some setae on the inner

third with comb-like setae ed about surface; segment regular 20 mm. In general appearance, the

the lower in towards along margin, increasing length species makes a slender impression (fig. 1). The the distal end lateral lobes of the head are truncated. The eyes

Echinogammarus margalefi n. sp. are reniform, more than twice as long as wide.

the first Both and second antennae are long, very DESCRIPTIVE PART slender without characteristic setation. Their

have 32 and 18 22 All descriptions, unless the contrary is explicitly flagelli to 40, to segments, res-

stated, apply to the adult male only. pectively. has unarmed The mandible palp an first seg-

berilloni Echinogammarus (Catta, 1878). ment; the inferior margin of the third segment is

Figs. 1—3, 4 G-H armed with a row of spinules, subequal in length.

first The gnathopod has a piriform hand with a

Principal references. — A very oblique palm (fig. 2A). strong medial Gammarus berilloni Catta, 1878 : 68, fig. 1; Chevreux, palmar spine, from the 1896 : 29, figs. 1—3; Margalef, 1953 : 193, fig. 235. clearly separated palmar

Gammarus (Echinogammarus) berilloni; Schellenberg, angle spines, is always present. The hand of the

1942 : 36—37, 19; & 1953 : 301. fig. Stephensen Hynes, second gnathopod is stronger than the first (fig. Echinogammarus berilloni; Chevreux & Fage, 1925 : 259, 2B). Its oblique palm is also provided with a fig. 269; Boecker, 1926 : 5—8; Vandel, 1926 ; 35—39; strong medial palmar spine. The fifth to seventh Spandl, 1926 : 128—132, figs. 1 —4; Le Roux-Legeux,

are characteristic. In these the 1927 : 34—43; Legeux, 1927 : 67—68; Pacaud, 1944 : legs very legs,

36—44; Margalef, 1944 : 202 —206, figs. 7 —9; Pacaud, of anterior margin basis, merus and carpus are 1952 : 101—103; Holthuis, 1956 : 83—95, fig. 8; Hoff- armed with densely implanted tufts of short setae, 1963 96 1964 mann, : —98, figs. 29—37; Vincent, : 809; the number of them increasing with age. The Vincent, 1967 : 2248; Pinkster, 1969 : 137 —150; Marga- of basis and as well as lef, 1970: 169; Pinkster & Stock, 1971: 37—51; Vincent, posterior margin merus,

1971: 1 —131, fig. A; Gras & Maasen, 1971: 54—55. the lateral surface of the basis, is set with groups

of in and num- very long setae, increasing length

Material examined. — ber from P5 to P7 (see fig. 2C). More than 500 samples from the entire distribution area Uropod 1 extending as far as, or slightly farther (see Map II). than, uropod 2. The inner and outer rami of

Description. — uropods 1 and 2 are almost equal in length. They

of this armed with variable of A redescription species was already given are a number spines (see by the present author in 1969. Since E. berilloni figs. 4G, H).

is considered the ancestor of this 3 has short species-group, Uropod a very endopodal segment. the most important characters will be summarized. The exopod is armed with clusters of setae (some-

Male: A maximum observ- times large species, length intermixed with 1 or 2 spines) on the inner

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Fig. 2. Echinogammarus berilloni (Catta, 1878), ￿, from hand of second leg (III); C, seventh leg (II); D, telson

the neotype locality. A, hand of first leg (scale III); B, (III); E, pleosome and urosome (II). Downloaded from Brill.com10/08/2021 03:18:23PM via free access 6 S. PINKSTER - THE ECHINOGAMMARUS BERILLONI-GROUP

and outer margin. The second segment of the species has never been found in upper reaches of

is and is the terminal exopod reduced as long as streams. Although there is no indication for a

spines (see fig. 3H). strong interspecific competition, the species is The telson is little shorter than the a peduncle always replaced by other members of the family

of 3; each lobe bears a lateral of 1 in uropod group Gammaridae the upper reaches, viz. E. aquilifer

spine (rarely lacking) and 2 to 3 setae, and an in the western parts of the Pyrenees, E. calvus in

apical group of 1 to 3 spines and many long setae; the spring zone of the Rio Ebro, Gammarus

its dorsal armed surface is never (fig. 2D). duebeni in the western part of Brittany (Bretagne),

The metasome segments, urosome segments and and members of the G. pulex-group in the rest of

the epimeral plates are armed with clusters of Europe.

often sometimes with long, very curved setae, In the Nive, a stream that originates on the top

small spines in between them. Almost invisible of Mont Mondarrain (France, dépt. Basses-

underneath these setae, the meta- and urosome Pyrénées) the type locality of E. berilloni

bear segments a dorsal hump (fig. 2E). only E. aquilifer was found in the narrow fast

running upper stretch of the river; as soon as the

Female: Smaller than the male (maximal length stream berilloni gets wider, E. appears. The zone observed after examination of 500 about samples: of overlap, about 4 km long, points to the absence 16 Like in other of there mm). groups amphipods, of severe competition between the two species (see exists a marked sexual So, the clearly dimorphism. also Stock et al., 1966; Pinkster et al., 1970; Gras setation the Al and on peduncle segments of both & Maassen, 1971).

A2 is more strongly than in the male developed Vincent, 1971, in his thesis, comes to the con-

3A and The hand in 1 and (figs. 3B). gnathopods clusion that low temperatures and a low concen-

2 is less and has an almost developed transverse tration of Ca-ions are limiting factors for this without the medial for palm, palmar spine usual species. This view is supported by ecological data, the male (for figures see Pinkster, For the collected in the and 1969). mid-western parts of France

remainder, it can be stated that the characters by many laboratory experiments. In the region

found in the male are less in the pronounced prospected by Vincent, E. berilloni was never

female. the number of setae the So, on metasome, found in waters in which the amount of Ca-ions

the and the basal urosome, especially on segments was lower than 15 mg/1. In his experiments Vin- of P5 P7 is much reduced Pinkster, through (see cent proved that at lower temperatures (4° C),

1969, and E. fig. 3J). berilloni was unable to recompensate the loss

of Ca-ions during the moult, when the outside Immature: In the setation of the is general legs concentration of Ca-ions was below 20—25 mg/1. less in the number of developed juveniles; seg- At higher temperatures (15° C) the minimum out- in the is still ments antennae low and the armature side concentration of Ca-ions, necessary to harden of the dorsal surface of metasome and urosome the carapace, was considerably lower (6 mg/1). consists of small and spines short setae only. This interaction between the hardness of the

During the course of the development, the setae water and its temperature certainly helps to ex- grow out to obtain their characteristic appearance plain the absence of E. berilloni in spring areas of

at the age of sexual maturity figs. 3D the rivers in the (see through western Pyrenees, such as on the The medial of PI be absent 3K). palmar spine can top of Mont Mondarrain. However, it does not in males (fig. 3C). very young explain the distribution pattern of this species in

western France, e.g. its presence in regions such as

— In general the is Variability. variability pattern Brittany (Bretagne) and its absence in the water

similar to that discussed in a previous chapter. system of the rivers Vilaine and Loire. In western

Brittany very dense populations of E. berilloni

Ecology. E. berilloni is a from the have been found in soft species very waters, even with less middle of courses streams and rivers in a large than 6 Pinkster mg Ca/1 (see et al., 1970). On the

area of western When estuarine Europe. species other hand, E. berilloni has not been found (Gras such are absent, as Gammarus chevreuxi, in & in the e.g. Maassen, 1971) drainage systems of the certain places in north-western is Spain, it some- rivers Vilaine and Loire, where high amounts of

times able to into the estuarine of penetrate region Ca-ions are available and the yearly changes in river its distribution systems. Throughout area the temperature are rather moderate. E. berilloni is

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Fig. 3. Echinogammarus berilloni (Catta, 1878), A, B, and E, fifth leg (II); F, sixth leg (II); G, seventh leg (II);

from the C— from the third I, urosome J, urosome J, ￿, neotype locality; K, juvenile H, uropod (III); (II), (II); neotype locality. A, first antenna (scale II); B, second K, epimeral plates (II). antenna (II); C, hand of first leg (III); D, fourth leg (II); Downloaded from Brill.com10/08/2021 03:18:23PM via free access 8 S. PINKSTER - THE ECHINOGAMMARUS BERILLONI-OROUP

also absent from — certain parts of Les Landes (S.W. Material examined.

France). Spain Santander: prov. In the light of the results of Vincent and our — Rio Ebro, W. of Reinosa, 5-VII-1969, rather fast own field observations, we must assume that in running stream, CI 7 mg/1, Ca 36 mg/1, many speci-

as in other E. berilloni, gammarid species (e.g. mens (ZMA).

Gammarus pulex, see Vincent, 1971) physiological — Springs of the Bbro in Fontibre, 5-VII-1969, CI

2 mg/1, Ca 18 mg/1, many of races occur, adapted to waters with high or with specimens, many them in precopulation (ZMA). low amounts of Ca-ions. The same holds true for — About 20 m downstream of the springs of the Ri'o the factor In western pH. exploring Europe, mem- CI Ebro, 5-VII-1969, 2 mg/1, Ca 18 mg/1, many bers of research found E. berilloni in our team specimens (ZMA).

waters varying in pH from 4.5 (in Bretagne) to 8 — Tributary of Rio Besaya, near village Arenas de Iguna (halfway Reinosa and Torrelavega), 16-VII- (localities in the Vendée) (Pinkster & Stock, 1970; 1969, shallow, about 3 m wide, CI 15 mg/1, Ca 66 Gras & Maasen, 1971). mg/1, Ca + Mg 54 mg/1, many specimens (ZMA). The occurrence of E. berilloni in the estuarine — Rio Saja, S. of Renondo (= S. of Cabuérniga), 5-

part of some rivers in north-western Spain and VII-1969, large, fast running river, CI 8 mg/1, Ca 18 Ca Mg 0 (ZMA). underneath pebbles on sandy beaches (near Playa mg/1, + mg/1

— Cascade in tributary of the Rio Saja, on road C de San Antolin, Spain, Prov. Oviedo) are an indi- 625, 14 km S. of Cabuérniga, 5-VII-1969, clean, fast cation that the can stand rather species high CI 7 Ca 8 running, mg/1, mg/1, many specimens,

salinities as well as considerable in of them in changes salinity. many precopulation (ZMA).

This was confirmed by experiments of Vincent, — Upper reaches of Rio Besaya, near village Lantuene

(= N.E. of 16-VII-1969, 40—50 cm wide, 1971, showing that E. berilloni behaved "nor- Reinosa), bottom of lime-concretions, CI 7 mg/1, Ca 82 mg/1, mally" in more concentrated media, up to 80% many specimens (ZMA). sea-water or 15.680 (= approx. 24.8% c S, mg Cl/1). prov. Burgos:

From from all of western samples parts Europe — Rio de Riaza, on road N 122 near Fuentecén

it became clear that this species can withstand a Aranda del Duero, 20-VI-1970, fast running, clear,

30 cm—1 m deep, 10 m wide, 13.4°C, pH substantial amount of organic pollution and high temp. 6.5, Ca 57 mg/1, Cl 11 mg/1, many specimens, many In of it has been temperatures. some parts Spain of them in precopulation (ZMA). found in waters having summer temperatures — Small brook in Quintanaelez (on road Pancorbo- ° around 31 C. CI 9 Ca 50 Ona), 5-VII-1969, mg/1, mg/1, many specimens (ZMA).

Distribution. — E. berilloni is known from the — Valdivielso Tributary of the Ebro near Quintana de north-western of part Spain (see Map III; own (= S.W. of Medina), 5-VII-1969, CI 7 mg/1, Ca

of the 84 exploration); the western part Pyrenees; mg/1, many specimens (ZMA).

— Nameless tributary of Rio Arlanzon, W. of Las those parts of France which drain into the Atlan- Quintanillas, 21 km W. of Burgos, about 3 m wide, tic, except for the higher Plateau Central and the CI 12 Ca 62 of mg/1, mg/1, many specimens, many drainage systems of the Loire and Vilaine (see them in precopulation (ZMA).

II; own the Channel Islands map exploration); — de Rio Pisuerga on road N 120 near village Melgar

(own exploration); Belgium (Stephensen & Hynes, Fernamental, 16-VII-1969, more than 20 m wide, CI 8 Ca 78 (ZMA). 1953); Luxembourg, (Hoffmann, 1963); the south- mg/1, mg/1, many specimens

— Tributary of Rio Arlanzon near village Sarracin eastern provinces of the Netherlands (Holthuis, (7.5 km S. of Burgos), 3 m wide, CI 20 mg/1, Ca the south-western of the German 1956); parts 74 (ZMA). mg/1, 16-VTI-1969, many specimens Federal 1926; Republic (Boecker, Schellenberg, — Rio Hormazuela, near villageVillanueva de Arganó.

In m wide, CI 1942; Besch, 1968). spite of intensive explora- on road N 120, 16-VII-1969, about 3

tion outside this 6 mg/1, Ca 86 mg/1, many specimens (ZMA). area by the present author and

prov. Logrono: other members of the research team of the In-

-— Rio Najerilla, about 5 km S. of Najera (along road stitute of Taxonomie Zoology of the University of C 113), major river, 16-VII-1969, CI 2 mg/1, Ca Amsterdam, the has species never been found 50 mg/1, many specimens (ZMA).

outside this area (see Map II). — Rio Najerilla, S. of village Bobadilla (near road

C 113), 16-VII-1969, large river, CI 6 mg/1, Ca 28 calvus rank. Echinogammarus (Margalef, 1956) new mg/1, 20 specimens (ZMA).

4 A-F — Figs. Rio Tiron, near road N 232 and village Tirgo

Gammarus berilloni subsp. calvus Margalef, 1956 : 33, (= S.W. of Haro), 16-VII-1969, major river, deep,

CI 15 Ca 134 fig- 2. mg/1, mg/1, many specimens (ZMA).

Echinogammarus berilloni subsp. calvus; Margalef, 1970 : — Rio Oja near village Casalaceina on road N 232

170, figs. 2 A, B. (S.W. of Haro), 16-V1I-1969, large river, CI 9 mg/1, Downloaded from Brill.com10/08/2021 03:18:23PM via free access BUDRAGEN TOT DE DIERKUNDE, 43 (1) - 1973 9

Fig. 4 A—F. Echinogammarus calvus (Margalef, 1956), F, pleosome and urosome (II).

￿, from the sources of Río Ebro. A, second antenna G— H. Echinogammarus berilloni (Catta, 1876), ￿, from

mandible hand of first the first second (scale II); B, palp (III); C, leg neotype locality. G, uropod (scale II); H,

(III); D, hand of second leg (III); E, seventh leg (II); uropod (II). Downloaded from Brill.com10/08/2021 03:18:23PM via free access S. PINKSTER - THE ECHINOGAMMARUS BERILLONI-GROUP

10

Ca 34 (ZMA). the basal in mg/1, many specimens are implanted on segment of P7 E.

prov. Vizcaya: berilloni, only a few groups of short setae remain

— Nameless confluent, 0.5 km E. of Bolivar (= 3.5 (fig. 4E). km S.W. of Marquina), 17-VII-1969, 2—3 m wide, The telson third fast CI Ca 58 and both in running, 14 mg/1, mg/1, many speci- uropod resemble,

mens (ZMA). shape and armature, those of E. berilloni.

— the Rio Butrón, near Gâmiz Tributary of village The second and third epimeral plates show the (= N.E. of Bilbao), 17-VII-1969, stagnant, deep, most characteristic features for this species, since CI 22 mg/1, Ca 84 mg/1, 15 specimens (ZMA). have setae the Alava: they very long suspended along prov.

— ventral the surface is - Rio Omecillo, S. of Espejo, 4-VTI-1969, few speci- margin, while lateral com-

mens (ZMA). pletely unarmed. Along the posterior margin only

— Rio Araya, 2 km S. of Araya, about 30 km E. of a few setules occur (fig. 4F). Vitoria, 15-VII-1969, about 3 m wide, slowly run- The urosome segments are armed with small CI 5 Ca 84 ning, mg/1, mg/1, many specimens spines and additional The (ZMA). setae only (fig. 4F).

— of the Rio Bayas, S. of Subljana, 4-VTI-1969, Cl 9 mg/1, armature metasome segments is still more of them in Ca 72 mg/1, specimens, many many poorly developed. This baldness was a reason for precopulation (ZMA). Accompanying species: E. Margalef (1956) to give this form the subspecific longisetosus (see table I). name calvus. Palencia: prov.

— Rio Pisuerga near village Olleros, 16-VII-1969,large

river, about 10 m wide, deep, CI 7 mg/1, Ca 82 Female: The females of this species closely mg/1, specimens (ZMA). many resemble those of E. berilloni. Vitoria: prov.

— Spring area of Rio Deva near Salinas de Léniz (N — This Variability. species, originally described as of Vitoria), 4-VII-1969, moderately fast running, a subspecies of E. berilloni, is a rather stable one. CI 68 mg/1, Ca 120 mg/1, many specimens (ZMA) features prov. Valladolid: Although some are variable to some extent

Tributary of Rio Duero, W. of Fuensaldana, near as discussed in the previous section on variability, Vallodolid,5-IX-1970, slowly running, clear, 30 cm the such the discriminating characters, as armature deep, 50 cm wide, temp. 20.8 °C, pH 6, CI 188 mg/1, of the the absence of setae on Ca 16 of them in epimeral plates, long mg/1, many specimens, many the dorsal of precopulation (ZMA). Accompanying species: Gam- side the body and the setation of

marus gauthieri. the legs, are very stable. Since both E. berilloni

and this form are distributed in the same river

system (Rio Ebro) showing a certain area of

— and since Description. overlap (table I), no isolating mechanisms

Maie: large, largest specimen observed 20 mm can be indicated, we believe that this form, like

long. The shape and size of the eye, the shape of Echinogammarus berilloni, must be considered to

~~ the head and the morphology of the antennae and be a good species: Echinogammarus calvus.

mouth parts are almost identical to those of E.

and — berilloni (for figures see Pinkster, 1969, Distribution. Up to now the species is known

4 A, The first from rather limited present paper figs. B). gnathopod a area in the northwestern part

differs from those in E. berilloni, of (fig. 4C) slightly Spain only (see map III).

in that the setae, implanted on the lateral surface

much The — of the propodus, are longer. same ap- Ecology. Echinogammarus calvus is a species

the from the plies to second gnathopod (fig. 4D). upper and middle reaches of rivers,

The third and fourth legs closely resemble those where it seems to prefer rather fast running waters. of E. berilloni for setation which is except the a The chlorinity and hardness (Ca concentration) of little shorter. the waters inhabited seem to have little influence

seventh fundamen- the of the The fifth through legs are on occurrence species. It was found in tally different from those in E. berilloni in the waters with CI concentrations from 2 to 70 mg/1

and the found the basal number length of setae, on and with Ca concentrations from 8 to 134 mg/1.

the on the Neither the absence of segments: long setae, implanted presence or vegetation, of this posterior margin segment, characteristic for nor the nature of the bottom seems to affect the

E. berilloni, are completely absent in E. calvus. animal since dense populations have been found

Instead of the many groups of long setae, which under extremely varying conditions.

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Fig. 5. Echinogammarus meridionalis n. sp., ￿, from the (II); F, hand of first leg (III); G, second leg (II); H, type-locality. A, head (scale II); B, first antenna (II); C, hand of second leg (III); I, fifth leg (II); J, third uropod second antenna (II); D, mandible palp (III); E, first leg (III); K, pleosome and urosome (II). Downloaded from Brill.com10/08/2021 03:18:23PM via free access 12 S. PINKSTER - THE ECHINOGAMMARUS BERILLONI-GROUP

— — Fig. 6 A D. Echinogammarus aquilifer Pinkster, 1969, E I. Echinogammarus meridionalis n. sp., ￿ from the

￿, from the type-locality. A, second antenna (scale II); type-locality. E, third leg (II); F, fourth leg (II); G. sixth

B, hand of first leg (III); C, hand of second leg (III); D, leg (II); H. seventh leg (II); I, telson (III). telson (III). Downloaded from Brill.com10/08/2021 03:18:23PM via free access BUDRAGEN TOT DE DIERKUNDE, 43 (1) - 1973 13

meridionalis hand of the second 5G, is Echinogammarus n. sp. gnathopod (figs. H)

Figs. 5,6 E-I stronger than that of the first. Its palm is less ob-

lique than in E. berilloni. Like in the first leg, the

Material examined. — medial palmar spine is always absent; a group of Portugal 5 is found near the The setae Leiria: spines palmar angle. prov. short. — on the lateral surface of the are Rio Alcaide, on road N 243, near Alcaria, S. of propodus Batalha, third fourth slender 25-VI-1970, slowly running, slightly pol- The and legs are (figs. 6E, F),

luted stream with gravel bottom, 5—30 cm deep, the setation on the third being longer than that on

10—200 cm wide, temp. 17°C, pH 6.0, CI 15 mg/1, the fourth. Ca 81 of them in mg/1, many specimens, many pre- The basal of 5 has a back- Male and have segment leg (fig. 51) copulation. holotype many paratypes ward lobe. It is armed with been deposited in the Zoölogisch Museum Amster- protruding sparsely dam (ZMA and small Amph. 103.027). some small spinules along the anterior

— Small brooklet near Valado, E. of 25-VI- Nazara, setules the The other near posterior margin. seg- 1970, slowly running, slightly polluted stream with ments bear few groups of spines and setae only, sandy bottom, 20 cm deep, 2—3 m wide, temp. rather short in with other 17.5°C, pH 6.0, CI 52 mg/1, Ca 13 mg/1, 2 speci- being comparison mem-

mens (ZMA). bers of the E. berilloni-group. The sixth and

seventh have legs a slender, elongate basal seg-

— 6G, Description. ment, likewise very poorly armed (figs. H). Male: little smaller A very characteristic species, a The setation of the distal segments is a little longer than E. berilloni. The largest male in the material than in P5. examined measured 16 mm. The uropods closely resemble those of E. beril- with and Head (fig. 5A) truncate lateral lobes, loni (fig. 5J). The telson lobes are only little longer obtuse The are with than Each lobe of 1 angles. eyes relatively large, wide. bears a lateral group a dark colour in live of to 3 deep specimens. spine and some setae, and an apical group 1 The first resembles that of E. antenna (fig. 5B) spines and 2 to 4 setae (fig. 61). berilloni. Its and have 5 accessory main flagellum As in E. calvus, the segments of the metasome

6 and 30 36 to to elements, respectively. and urosome are armed with some small spines The second extremely setose antenna (fig. 5C) and a small number of setules (fig. 5K) only. The

is far the characteristic feature of the by most first urosome segment shows a dorsal concavity animal, and makes the species clearly recognizable ("saddle"). within this It is little shorter than the first group. a The epimeral plates are characterized by the antenna. The flagellum is shorter than the slender absence of setation, and sharp postero-inferior

19 in peduncle, possessing (18 to 20) segments corners (fig. 5K). larger specimens. The distal half of the 4th and

5th the entire peduncle segments, both segments Female: Even more than in other species within almost in are covered with transverse female resembles the male. equal length, this group, the rows of long, slightly curved setae. The flagellum Although the flag-like setation of A2 is less is likewise armed with it remains densely implanted setae, developed than in the male, still very

diminishing in length towards the end. As a whole characteristic.

this setation makes the impression of a flag along

the entire ventral margin of the antenna. Ecology and distribution. — Since the species

The mandible differs from both palp (fig. 5D) only was found in two localities only (map III), the found in berilloni one E. in having 6 long in the province of Leiria, Portugal, we cannot give

terminal the third instead of 4 setae on segment, other information on its ecology than that men-

to 5. tioned undermaterial.

The elongate coxal plates 1 to 4 have rounded

inferior and bear few setules Echinogammarus longisetosus n. corners, very along sp. the lower margin. The hand of the first gnathopod Figs. 7-9

(figs. 5E, F) has an oblique palm. Except for the

normal the consists of Material examined. — groups of setae armature a Spain transverse row of spines near the palmar angle prov. Navarra: and some smaller so-called "Stiftstacheln" only. — Rio Arga, 0.5 km upstream of Urtasun (= N.N.W. The medial is palmar spine always absent. The of Pamplona), 14-VII-1969, moderately fast running

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CI 16 Ca 56 — Rio stream, gravel bottom, mg/1, mg/1, Ursurian, near village Ursurian, 15-VII-1969,

of them in with bottom, about 1.5 m many specimens, many precopulation. polluted stream, stony

and wide, CI 11 Ca 68 8 (The $ holotype, 9 allotype, many paratypes mg/1, mg/1, specimens (ZMA).

Alava: have been deposited in the Zoölogisch Museum prov. Rio Amsterdam, under cat. no. ZMA Amph. 103.025). Bayas, S. of Subljana, 4-VII-1969, CI 9 mg/1,

E. berilloni. Ca 72 of them in Accompanying species: mg/1, many specimens, many

— Rio Irati, near village Arrive, 14-VII-1969, pollut- precopulation (ZMA). Accompanying species: E.

calvus. ed, moderately fast running stream, CI 4 mg/1, Ca

54 mg/1, 4 juveniles (ZMA). — Small brook near Tuesta, 4-VII-1969, about 1 m with — Erreca-Beltz alongside road N 121, about 21 km N. wide, stony bottom, CI 10 mg/1, Ca 74 mg/1,

fast of of Pamplona, 14-VII-1969, clear, moderately many specimens, many them in precopulation

with 3 CI running stream, stony bottom, m wide, (ZMA).

9 mg/1, many specimens (ZMA). prov. Burgos:

— — Small road W.N.W. of Rio Ibur, near village Irurita, 15-VII-1969, clear, brook, on Pancorbo-Ona,

with CI 16 1 CI 8 Ca 84 slowly running stream, gravel bottom, Fuenta, 5-VII-1969, m wide, mg/1,

of them in mg/1, Ca 32 mg/1, 20 specimens (ZMA). mg/1, many specimens, many precopula-

— about tion Rio Leoz, on road N 121, 3 km N. of Be- (ZMA).

— Rio near Terminon, S.W. of Ona, 5-VII-1969, rasain, near Tafalla, 19-VI-1970, clear, fast running Óca,

stream with stony bottom, temp. 16°C, pH 6.5, 20 moderately fast running stream with clay bottom,

5 to 10 CI 93 Ca 154 to m wide, CI 77 mg/1, m wide, mg/1, mg/1, 30 cm deep, 1 20 mg/1, Ca many

of them in of them in many specimens, many precopulation specimens, many precopulation (ZMA).

(ZMA). prov. Zaragoza:

— — of Rio 0.5 km N. of Monasterio Rio Leoz, on road N 121, 1 km N. of Tafalla, Tributary Piédra, Piédra, 15 of de 19-VI-1970, fast running, strongly polluted stream, de km S.E. Alhama Aragon, 13- fast with bottom of clay and stones, temp. 18°C, pH VII-1969, clear, running stream, clay bottom,

about m wide, CI 15 6.5—7, CI 18 mg/1, Ca 127 mg/1, 2 specimens 1 mg/1, Ca 98 mg/1, many of them in (ZMA). specimens, many precopulation (ZMA).

— of Rio Piédra, B. of 13-VII- — Rio Alhama, 1 km N. of Cintruenigo, 19-VI-1970, Tributary Nuevalos,

1969, clear fast stream, with bottom, slightly polluted, moderately fast running stream, running gravel

with CI 300 about 1.5 m wide, pH 6.5, Cl 1 mg/1, Ca 182 mg/1, stony bottom, temp. 20°C, pH 6.6, specimens, of them in precopulation mg/1, Ca 222 mg/1, 20 to 40 cm deep, 4 to 10 m many many species: E. echinosetosus. wide, many specimens (ZMA). (ZMA). Accompanying

— of Rio near — Zuera, 13-VII-1969, Tributary of Rio Araquil, about 4 km N. of Irur- Tributary Gallégo,

30 clear, fast stream, bottom, 5 cm zun, on road N 240, 25-VIII-1968, specimens running gravel

30 cm Cl 1 Ca 128 mg/1, (ZMA). deep, wide, mg/1, many of them in — Confluent of Arroyo Artesiaga, S. of Irurita, 15-VII- specimens, many precopulation (ZMA).

E. berilloni. 1969, clear, fast running stream, with gravel bottom, Accompanying species:

— Tributary of Rio Aragon, on road N 240, near 5 cm deep, 50 cm wide, CI 9 mg/1, Ca 38 mg/1,

of them in Asoberal, 36 km W. of Jaca, 14-VII-1969, clear, many specimens, many precopulation fast with bottom, (ZMA). Accompanying species: E. aquilifer. moderately running stream, gravel CI 90 Ca 102 — Tributary of Rio Arga, about 10 km N.W. of mg/1, mg/1, many specimens (ZMA).

— Ri'o near 84 km E. of Pamplona, onroad N 240, 25-VIII-1968, CI 17 mg/1, Martin, Escatron, Zaragoza,

of them in 14-IV-1967, many specimens (MSNV). many specimens, many precopulation

— Rio (ZMA). Gallégo, near Zuera, 15-V-1967, many speci-

— Rio Arga, on road C 135 near Zubiri, 25-VIII-1968, mens (MSNV).

CI 6 mg/1, 15 specimens (ZMA). — Small brook, about 7 km S.W. of Huesca, on road

stream with — Small brook, about 2 km W. of Aburrea Baja, N 123, 13-VII-1969, clear, fast running

bottom, about 10 cm 80 cm wide, CI 14-VII-1969, 1 to 1.5 m wide, stony bottom, CI gravel deep,

4 Ca 54 of them 23 Ca 92 many of mg/1, mg/1, many specimens, many mg/1, mg/1, many specimens, in precopulation (ZMA). them in precopulation (ZMA).

— Rincón de Ademuz: Small brook between Aburrea Alta and Aburrea prov.

Baja, N.E. of Pamplona, 14-VII-1969, CI 11 mg/1, — Tributary of Rio Turia, near village Las Santos,

Ca near of roads N 420 and N 330, 38 km S. 100 mg/1, many specimens (ZMA). crossing

— of fast Tributary of Rio Salazar, about 2 km W. of Esca- Teruel, ll-VII-1969, clear, moderately run- with bottom, CI 13 Ca 80 roz (about 28 km E. of Burguete), 14-VII-1969, ning stream, stony mg/1, mg/1, about 1 m wide, stony bottom, CI 5 mg/1, Ca 88 many specimens (ZMA). Teruel: mg/1, 15 juveniles (ZMA). prov.

San Sebastian: — Rio about 8 km S. of prov. Guadalope, Alcaniz, near

— of Tributary Rio Deva, upstream of village Men- village Castelseras, 12-VII-1969, slightly polluted,

daro, 4-VII-1969, polluted stream with stony bot- moderately fast running stream, about 10 m wide,

tom, CI 11 Ca 50 CI 18 Ca 52 12 mg/1, mg/1, many specimens mg/1, mg/1, specimens (ZMA).

(ZMA). Accompanying species: E. berilloni. — Ri'o Guadalopillo, about 24 km S.W. of Alcaniz,

7. longisetosus n. from the tenna (II); C, fifth D, third (III); E, Fig. Echinogammarus sp., ￿, leg (II); uropod type-locality. A, first antenna (scale II); B, second an- telson (III).

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fast with about 50 2—3 12-VII-1969, rather running stream, with stony stream gravel bottom, cm deep,

bottom, about 2 m wide, CI 47 mg/1, Ca 208 mg/1, m wide, temp. 13.8°C, pH 5.0, CI 22 mg/1, Ca 102

1 many juveniles (ZMA). mg/1, specimen (ZMA).

— Rio Guadalopillo, 32 km S.W. of Alcaniz, 2 km — Rio Fluvia, under bridge near Esponella, N.E. of

S.W. of village Alcoriza, 12-VII-1969, rather fast Banolas, 18-IV-1971, slightly polluted, fast running

with bottom, to m more running stream, with stony bottom, about 1.5 m stream stony 0.5 2 deep,

CI 23 Ca 110 than 10 CI 100 wide, mg/1, mg/1, many specimens m wide, temp. 16.2°C, pH 6.0, mg/1,

in (ZMA). Ca 138 mg/1, many specimens, many of them

— Tributary of Rio Estercuel, alongside road N 211 precopulation (ZMA).

near village La Zoma, alt. 1100 m, about 14 km E. prov. Lerida:

of Montalban, 12-VII-1969, clear, fast running — Rio Farfana, at Castello de Farfana, W. of Bala-

stream with stony bottom, CI 5 mg/1, Ca 60 mg/1, guer, 19-IV-1971, moderately polluted, moderately

of them in fast stream with 50 deep, many specimens, many precopulation running stony bottom, cm

(ZMA). 1 m wide, temp. 16.2°C, pH 6.5, CI 270 mg/1, Ca

— Rio de Cabra, near village Castel de Cabra, 11 km 198 mg/1, 20 specimens (ZMA).

E. of Montalban, 12-VII-1969, slightly polluted, — Rio Llobregos, at Pons, N.E. of Balaguer, 19-IV-

moderately fast running stream, CI 9 mg/1, Ca 98 1971, clear, moderately fast running stream with

of them in bottom, than 1 m 2 3 m wide, mg/1, many specimens, many precopula- clay more deep, to

tion (ZMA). temp. 14.2°C, pH 6.0, CI 93 mg/1, Ca 148 mg/1,

— of S. of about 25 of them in Tributary Rio Turia, Libros, km many specimens, many precopulation

S. of Teruel, ll-VII-1969, 1 —2 m wide, gravel (ZMA).

CI 34 Ca 84 — Fontana de la well road C bottom, mg/1, mg/1, many specimens Bargasses, alongside 147,

(ZMA). Accompanying species: E. margalefi. 4 km S. of Selles, S. of Tremp, 14-IV-1971, clear,

— Rio Jiloca, near Caminreal, 12-VII-1969, clear, fast moderately fast running stream with gravel bottom,

running stream, about 2.5 m wide, more than 2 m about 5 cm deep, temp. 12.4°C, pH 6.0, CI 6 mg/1,

of them in deep, pH 6.5, CI 34 mg/1, Ca 110 mg/1, many Ca 80.3 mg/1, many specimens, many

of them in (ZMA). specimens, many precopulation (ZMA). precopulation

— Alfarras, of Accompanying species: E. echinosetosus. Ribeira Noguéra Ribagorzana, near N.

— Rio Mijares, near Mora de Rubielos, 7-V-1969, alt. Lerida, 19-IV-1971, slightly polluted, slowly running

to 700 stream with bottom, 50 cm 4 5 m m, many specimens (MSNV). stony deep,

— wide, 15.1°C, 6.0, CI 35 Ca 95.9 Rio Mijares, near Mora de Rubielos, 8-V-1967, temp. pH mg/1,

of them in many specimens (MSNV). mg/1, many specimens, many precopula-

Barcelona: tion prov. (ZMA).

— Ribeira Pontóns, 500 m E. of Pontons, W. of Villa- Description. — franca del Panades, 16-IV-1971, slightly polluted, Male: A medium large species; the maximum moderately fast running stream with stony bottom, length observed after examination of 45 samples 20 to 40 cm deep, 3 to 4 m wide, temp. 15.0°C, pH is 15 mm. In general the body is slender and 5.0, CI 85.7 mg/1, Ca 59.2 mg/1, many specimens,

many of them in precopulation (ZMA). resembles that of E. berilloni, although the ar-

— Small brook in Sierra de Montsény, near Santa Fé rangement of the setae is completely different. de Montsény, alt. 1500 m, 3-V-1967, speci- many The head (fig. 9A) has broadly rounded lateral mens (MSNV). lobes; sinus rather shallow. The are subreni- — eyes Another small brook in Sierra de Montsény, near form in in Santa Fé de alt. 1500 juveniles, elongate and narrow older Montsény, m, 2-V-1967, many specimens (MSNV). animals.

-— Small brook in Sierra de Montsény, alt. 1300 m, The first antenna has no striking differences as

3-V-1967, many specimens (MSNV). compared to that of E. berilloni (fig. 7A). prov. Tarragona: The flagellum of the second antenna (fig. is — Cemented well near Santa Creus, N.E. of Vails, 7B)

to The 15-IV-1971, slightly polluted, moderately fast run- short, compared the peduncle segments.

about 10 cm pH 5.0, CI ning, deep, temp. 14.4°C, second peduncle segment bears the short gland 25 Ca 104 of mg/1, mg/1, many specimens, many Peduncle 4 is little shorter cone. segment a than them in precopulation (ZMA). segment 5. The setae along the ventral margin of — Rio Gaya, 500 m S. of Aguamurcia, N.E. of Vails, 4 and 5 in 15-TV-l971, clear, moderately fast running stream segments are to some extent variable

with stony bottom, 50 cm deep, 2—4 m wide, temp. length.

17.4°C, pH 5.0, CI 6 mg/1, Ca 104 mg/1, many The propodus of the first leg is a little longer specimens, of them in precopulation (ZMA). many than the carpus, elongate, pyriform; armed with a Gerona: prov. medial palmar spine and with some spines near — Tributary of Rio Fluvia, N. of Crespia, S. of the the Figuéras, 18-1V-1971, clear, moderately fast running palmar angle. Moreover, merus, carpus,

Fig. 8. n. sp., from the of second F, urosome G, Echinogammarus longisetosus ￿, leg (III); (II); epimeral plates type-locality. A, mandible palp (scale III); B, first leg (II).

(II); C, hand of first leg (III); D, second leg (II); E, hand

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lateral and propodus of this leg are provided with tufts of setae, both along the margins and on the

very long setae as illustrated in figs. 8B and C. surface (fig. 8G).

The hand of the second gnathopod (figs. 8D, E) The colour of live specimens varies from milky-

is relatively short, always less than twice as long white to greenish with orange dots on the

have as wide. In comparison with E. berilloni, the palm metasome segments. The antennae always an

is less oblique while the medial palmar spine has orange coloration. The proposed specific name,

moved a little towards the articulation with the longisetosus, alludes to the clusters of very long

dactylus. Near the palmar angle two transverse setae on the gnathopods, pereiopods, and uropods.

of rows spines are implanted. The lateral surface

and the posterior margin are covered with many Female: In this species, as in several others, the

groups of very long setae (most of them have been female differs from the male in the gnathopods,

omitted in the figures). which lack a medial palmar spine, and in the

The merus and of P3 and P4 setation, which is better on the anten- carpus (figs. 9B, developed

with dense brushes of but is rather the Nevertheless C) are provided very long nae, poor on legs. females of this make im- setae along the anterior margin. In addition, the species a more "hairy"

merus of P3 bears of setae the than females of other members in the groups long along pression

posterior margin. The coxal plates are relatively berilloni-group.

with short rounded inferior corners bearing short

setules. Variability. — In some populations from the

north-eastern of the the The basal segment of the fifth leg has a back- parts Spain, setae on

ward lobe. of the second antenna protruding The basis, merus, and peduncle segments are

of this with clusters of than in certain from the north- carpus leg are provided longer, populations

western of the Iberian very long setae along the anterior margin (fig. 7C). part peninsula.

The 6th and 7th leg have elongate basal segments.

and distribution. —- This is from Their setation is even more characteristic than that Ecology a species

middle of of P5, since the bundles of long setae do not only the moderately fast running courses

occur along the anterior, but likewise along the freshwater streams. It seems to prefer high con-

of and centrations of Ca-ions. Since it is a species with a posterior margin merus carpus (figs. 9D and

E). Some long setae, increasing in number from P5 rather large range (see map III) in the northern

half of it be found with to P7, are implanted on the lateral surface of the Spain, can sympatrically

various other such basal segments. species, as E. aquilifer in the

1 and 2 identical those in western E. berilloni and E. calvus in the Uropods are to E. Pyrenees,

berilloni. course ofthe Rio Ebro, and E. echinosetosus The elongate outer ramus of uropod 3 upper is armed with and E. margalefi in the south-eastern of its many tufts of very long setae on part Data both the inner and outer margins (fig. 7D). distribution area. on interspecific competition available table Like E. it The telson lobes are about 1.5 times as long as are not (see I). berilloni,

can stand rather of wide, each bearing 1 to 3 apical spines and one a high degree pollution.

lateral spine. The spines are accompanied by a Echinogammarus tarragonensis n. sp. varying number of very long setae. Sometimes Figs. 10A-D some long setae are found on the dorsal surface

Material examined. — of the lobes as well (fig. 7E). Spain The metasome is sparsely armed with very of the Rio km prov. Tarragona: Tributary Ciurana, 4.5 small and spines setules. the urosome Generally S. of Curnudella, W. of Reus, 14-IV-1971, clear, moder- armed with middorsal of segments are a group one ately fast running stream with gravel bottom, temp.

12.4°C, pH 6.0, CI 12 mg/1, Ca 59 mg/1, specimens, spine and some long setae, and some lateral many of them in precopulation. The holotype and There obvious many $ groups. are no elevations or excava- have been in the many paratypes deposited Zoölogisch tions (see fig. 8F). Amsterdam Museum under nr. ZMA Amph. 103.029. The back corners of the second and third

epimeral plates vary from almost rectangular to Description. —

slightly pointed. They are set with rather long This species is very similar to E. longisetosus,

9. from Fig. Echinogammarus longiselosus n. sp., ￿, the fourth leg (II); D, sixth leg (II); E, seventh leg (II). type-locality. A, head (scale II); B, third leg (II); C,

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Downloaded from Brill.com10/08/2021 03:18:23PM via free access Fig. 10 BUDRAGEN TOT DE DIERKUNDE, 43 (1) - 1973 21

Cl Ca 9 described before. It differs from this species in the wide, 11 mg/1, mg/1, many specimens,

of them in 1 many precopulation. Accompanying following characters (based on J ): species: Echinogammarus berilloni (Catta) (ZMA). (1) The setae on the peduncle segments of A2 are — River Laxa, S. of Espelette, 15-VII-1969, clear, fast shorter. running stream, with stony bottom, about 1.5 m (2) The setae implanted on the hands of the first wide, CI 10 mg/1, Ca 46 mg/1, 10 specimens (ZMA)

and second Accompanying species: E. berilloni. gnathopods (fig. 10A) are very

— River Ouhabia, on road from St. Pée to Arbonne, short in E. The against very long longisetosus. 15-VII-1969,slightly polluted, slowly flowing stream, number of spines near the palmar angle of P2 with some pebbles on sandy bottom, about 1.5 m is reduced. CI 21 Ca 40 wide, mg/1, mg/1, many specimens,

(3) The coxal plates of PI and P2 bear long setae many of them in precopulation (ZMA). Accom- species: berilloni. along the ventral margin (see figs. 10B and panying E. dépt. Landes: C). — Small brook, 2 km N.E. of St. Martin de Scignanx The setae the of 3 (4) along margins uropod (fig. of (= N.E. Arbonne), 7-V-1970, temp. 11.8°C, pH

1 in E. short in CI 23 OD), very long longisetosus, are 5, mg/1, Ca 45 mg/1, many specimens, many

this species. of them in precopulation (ZMA). Spain The other such the seta- characters, as very long prov. Navarra: tion of the legs, the structure of the mandible palp, — River Petite Nive, about 5 km S. of Valcarlos,

and the armature of the dorsal surface and telson, 25-VIII-1968, clear, fast running stream, about 50

are identical to those in E. longisetosus. cm wide, stony bottom, CI 27 mg/1. (The $ holo-

been Since the is found in type, V allotype and 27 paratypes have species one locality only deposited in the Zoölogisch Museum Amsterdam (see Map TTI), no other data on its ecology can be cat. under no. ZMA Amph. 102.126, 102.127 and given, than those mentioned under "Material 102.128).

examined". -— Small brook, just S. of village Almandoz (= S.W. of Elizondo), 15-VII-1969, slightly polluted, moder-

Remarks. — This species has characters in many ately fast running stream, with stony bottom, CI

common with E. longisetosus. However, the dif- 10 mg/1, Ca 48 mg/1, many specimens, many of

ferences mentioned them in precopulation (ZMA). in the preceding lines are

— Rio Baztan, N. of Elizondo, 15-VII-1969, clear, fast stable within this sample. They all fall outside the with running stream, stony bottom, 4 m wide, Cl 8 range of variability observed in the 45 samples mg/1, Ca 32 mg/1, 1 specimen (ZMA). Accom- of E. available longisetosus. Since this sample was panying species: E. berilloni.

collected within the of — Confluent of of Irurita, distribution area E. longise- Arroyo Artesiaga, S. 15-VTT-

1969, clear, fast stream, with bottom, tosus we believe that this form must be considered running gravel 5 cm deep, 50 cm wide, CI 9 mg/1, Ca 38 mg/1, valid a species. As the only locality known at of them in many specimens, many precopulation present is situated in the province the Tarragona, (ZMA). Accompanying species: E. longisetosus. specific name tarragonensis is proposed.

Description. —

Echinogammarus aquilifer Pinkster, 1969. The descriptions and illustrations given by

6A-D Pinkster form Figs. (1969) a good basis for the recog- nition of the species. For the sake of complete- E. aquilifer Pinkster, 1969 : 140—142, figs. 1, 3—8; Pink- the most with ster & Stock, 1971: 48. ness, important characters, along illustrations some are repeated.

Material examined. — Male: A relatively small species, maximum France length observed 12 dépt. Basses-Pyrénées: being mm.

— of the The lateral the head have Spring zone river Laxa, on Mont Mondar- lobes of rounded

rain, alt. 625 15-VII-1969, fast m, clear, running corners, the sinus is shallow; the eyes are elongate. stream, about 30 5—10 cm wide, cm deep, CI 11 The third of segment the first antenna is short, mg/1, Ca 8 of them mg/1, many specimens, many less than half as long as the second. in precopulation (ZMA). The — second antenna slender River on (fig. 6A) has Laxa, Mont Mondarrain, alt. 350 m, 15-

VII-1969, clear, fast running stream, about 2 m peduncle segments 4 and 5 armed with 3 rows of

Fig. 10 A—D. E—L. Echinogammarus tarragonensis n. sp., ￿, Echinogammarus margalefi n. sp., ￿, from the from the hand of second type-locality. A, leg (scale III); type-locality. E, first antenna (II); F, second antenna (II); first coxal B, plate (II); C, second coxal plate (II); D, G, detail of second antenna (III); H, head (II); I, hand third uropod (IV). of first leg (III); J, hand of second leg (III); K, urosome

(II); L, epimeral plates (II).

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of the spines and setae. The armature flagellum is Spain, on the foot-hills of the Pyrenees (see Map because of the absolutely distinctive, flag-like III). It is found under stones in narrow, clear, and

brush of setae the inferior which is rather fast along margin, running upper reaches of mountain

reminiscent of the situation found in G. pulex. streams, a biotope somewhat similar to that of

The hand of the first has gnathopod a very Gammarus fossarum in the French Alps. When A medial is the oblique palm (fig. 6B). palmar spine stream gets wider the species is gradually

but unlike in E. berilloniit is outnumbered E. present, not separated by berilloni or Gammarus pulex. from the wide palmar angle spines by a gap.

The hand of the second gnathopod (fig. 6C) is Echinogammarus zebrinus Pinkster & Stock, 1971.

relatively short, 1.5 times as wide as long. A Fig. 11

medial palmar spine is never found. The coxal E. zebrinus Pinkster & Stock, 1971: 43—51, figs. 4-—7. plates of the first two legs are almost rectangular,

while those in legs 3 and 4 are rounded.

Material examined. — see Pinkster & Stock, 1971. The basal segment of legs 5, 6 and 7 is relatively

wide, subrectangular. In leg 5 it °s hardly longer Description. — The than wide. The posterior margin of the basis is recent description of this species may serve

basis finely crenulated and set with short setae. Only as a good for its recognition. Male: It is few groups of setae are implanted on the lateral a large species (males up to 19 mm

surface have and it is the of the basal segments. For the remainder long been observed) only one

these less within this that be in the legs are setose than in E. berilloni and group can distinguished

field with resemblethose of E. calvus. the naked eye, because of its typical

it has The third uropod is more strongly developed colouration. Like a zebra, lighter and darker

than in berilloni. vertical on each E. The length of their setation (almost black) stripes body seg-

is intermediate between E. berilloni and E. lon- ment.

gisetosus. As in E. aquilifer, the flagellum of the second

brush The telson lobes are broad, little longer than antenna bears a flag-like of setae, the indi-

wide (fig. 6D). Apart from the apical and lateral vidual setae are a little shorter, however. On the

which other the fifth groups of elements, are usually found in hand, the setae on peduncle segment

this clusters of are than in the other 11 species group, long setae are im- longer species (figs. A, B).

The hands of both the first and planted on the upper surface as well. second gnatho-

have The metasome and urosome segments have no pods oblique palms, always provided with a

dorsal elevations excavations. medial A of or They are armed pointed palmar spine. group spines

is found the with small tufts of short, slighthly curved setae. always near palmar angle. The coxal

The 1 4 rounded posterior corner of the third epimere is a plates of legs through have corners

little sharper than that of the second. The epimeral and generally long setae along the inferior margin;

armature is rather scanty. legs 5, 6, and 7, and the third uropod are almost

The setation of the second antenna, of P5 identicalto those in E. berilloni.

and of As in E. the through P7, the telson resemble that found aquilifer, telson lobes bear one or

in but dorsal in addition the E. zebrinus, nevertheless the species can two groups of setae, to

be the normal of elements easily distinguished through absence of a groups (fig. 11C). Very characteristic is the medial palmar spine in P2 and the different ar- conspicuous armature of the

mature of the dorsum. dorsal surface of the body (fig. 11D), with its

numerous small curved spines. In addition there

tufts of Female: As in other species, the female is less are some setae, very short on the first

of the characteristic than the male. Nevertheless, females segment metasome, but increasing in length

this towards third from species can be easily identified, because and number the urosome segment. of the brush of flag-like setae on the second an- tenna. Female: Although most of the characteristic

features (such as the "flag" on the second antenna)

Ecology and distribution. — of the male are absent or less developed in the

Intensive sampling has that the still proved species females, this sex can easily be recognized by inhabits a small area only in the south-western very the striped colouration and the armature of the of and in the part France, north-western part of dorsum.

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Fig. 11. Echinogammarus zebrinus Pinkster & Stock, 1971, B, detail of second antenna (V); C, telson (III); D, pleo-

￿, from the type-locality. A, second antenna (scale VI); some and urosome (II). Downloaded from Brill.com10/08/2021 03:18:23PM via free access 24 S. PINKSTER - THE ECHINOGAMMARUS BERILLONI-GROUP

Variability. — Except for the "normal" variability, dépt. Hautes-Pyrénées: River 2 km S.E. of St. in the of l'Arros, Stevens, near Tarbes, this species shows some variability length 1 4-111-1 972, clear, moderately fast running stream, the endopod of uropod 3. It can vary from less with stony bottom, about 2 m wide, 10 to 60 cm than 10% to more than 20% of the length of the deep, pH 7.9, many specimens (ZMA). Accom-

exopod (see Pinkster & Stock, 1971). panying species: Echinogammarus pacaudi, E. beril-

loni.

— River la Bouès, 3 km E. of Autin, near Tarbes, Ecology and distribution. — In spite of intensive 14-111-1972, clear, moderately fast running stream, the found in sampling, species was only a very with stony bottom, about 1 m wide, 10—40 cm limited around Mont-de-Marsan in south- area deep, pH 8.2, many specimens (ZMA). Accom-

western France (see map in Pinkster & Stock, panying species: Echinogammarus pacaudi.

1971). In this region it inhabits a wide range of

in the chemical biotopes not only fluctuating com-

Description. — position of the water (e.g., the amount of Ca-ions

varied from 3.3 mg/1 to 69.8 mg/1), but also in Male: Maximal length observed 16 mm. The head and the of the identical those in the bottom composition (mud, sand, stones) and shape eyes are to E. berilloni. vegetation. Therefore, it is a little astonishing that

The first than a species of such adaptibility could never be traced antenna (fig. 12A) is longer half its restricted the body length. Its third is beyond very area. The species coexists peduncle segment than half the with both E. berilloni and E. pacaudi (see table I). more as long as second. The flagel-

lum and accessory flagellum have 26 to 30 and 5

to 6 The second antenna Remarks. Within the berilloni-group, E. segments, respectively.

meridionalis E. (fig. 13A) has slender peduncle and an aquilifer,and E. zebrinus occupy a segments

since brush 18- to As in the first special place they possess a flag-like 23-segmented flagellum.

the last of setae on the second antenna. They form a series antenna, two peduncle segments are

in which E. zebrinus is still closely related to E. ventrally armed with groups of 1 to 3 rather long

berilloni, while in E. meridianus the relation is setae (longer than the diameter of the segments

on which are Because of these hard to see. In this series there seems to exist a they implanted).

which resemble those of female E. beril- certain regularity, in that along with the presence antennae, loni, the is called of a flag-like brush on the second antenna there species feminatus, latin for

is a tendency towards the loss of the medial effeminate.

E. the palmar spines (and part of the armature of the Within the berilloni- group, feminatus is

with less of dorsum). So, in E. meridianus which has a brush only species a more or irregular row

both the setules the third of the mandible on flagellum and peduncle segments of on segment palp

the setae in the A2, the medial palmar spines are absent in both (fig. 13B), proximal part being

shorter than those in the terminal gnathopods; in E. zebrinus which has a less distinctly part.

Five terminal and of developed "flag", these spines are still present, plumose setae two groups both the inner while E. aquilifer is intermediate. These tendencies setae on and outer surface complete

the armature of this strike me, since both the gnathopods and the segment.

The hand of the first second antenna play such an important role during gnathopod (figs. 12B, 13C)

of has and medial the first stages the precopulation. an oblique palm a palmar spine.

For the remainder PI shows no particularities.

The hand of the second gnathopod (figs. 13D,

Echinogammarus feminatus n. sp. is than twice wide. 12C) more as long as A trans- Figs. 12-13 verse row of spines is implanted near the palmar

angle. In comparison to E. berilloni, the pointed

Material examined. — medial palmar spine has moved towards the France articulation with the In between this dépt. Basses-Pyrénées: dactylus.

— and the River le on road spine palmar angle another Landistou, D35, G.W. of Bruges, group, strong

about 20 km W. of Lourdes, 24-VIII-1968, Cl 7 pointed spine can be found. The coxal plates of

mg/1, 40 specimens, some of them in precopulation. the first four have legs rounded inferior corners, (The $ holotype, 9 allotype, and 38 paratypes and bear only few setae along the margins. have been deposited in the collection of the Zoölo- The setation in P3 is longer than in P4 (figs. gisch Museum Amsterdam, under cat. nr. ZMA These Amph. 103.026). 12D, E). legs, as well as the posterior three

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third fourth 12. Echinogammarus feminatus n. from the C, hand of second (III); D, leg (II); E, Fig. sp., ￿, leg type-locality. A, first antenna (scale II); B, first leg (II); leg (II); F, fifth leg (II); G, seventh leg (II). Downloaded from Brill.com10/08/2021 03:18:23PM via free access 26 S. PINKSTER - THE ECHINOGAMMARUS BERILLONI-GROUP

13. from the third telson and Fig. Echinogammarus feminatus n. sp., ￿, E, uropod (III); F, (III); G, pleosome

type-locality. A, second antenna (scale II); B, mandible urosome (II); H, sixth leg (II). Downloaded from Brill.com10/08/2021 03:18:23PM palp (III); C, hand of first leg (III); D, second leg (II); via free access BUDRAGEN TOT DE DIERKUNDE, 43 (1) - 1973 27

pereiopods, almost resemble those of E. calvus Montalban), 12-VII-1969, slightly polluted, moder- ately fast running stream with gravel bottom, about (see figs. 12F, G, 13H). 1 m wide, 50 cm deep, CI 17 mg/1, Ca 84 mg/1, The third uropod (fig. 13 E) is relatively short. of them in many specimens, many precopulation Its armature is rather for the berilloni- poor group. (ZMA).

for the — Except usual armature, the telson (fig. Rio Pancrudo on road N 234, about 5 km N. of

Cala Mocha, fast 13F) bears a few setae on the dorsal surface of the 12-VII-1969, clear, moderately

running about 2 m wide, gravel bottom, lobes. stream, CI 36 Ca 276 mg/1, mg/1, many specimens, many The metasome and urosome have no segments of them in precopulation (ZMA).

dorsal elevations or excavations and attract atten- — Rio Jiloca near Burbaguena, alongside road N 234, tion because of the setation 12-VII-1969, polluted fast about 3.5 poor only. The third running water,

m wide, CI 32 mg/1, Ca 160 mg/1, specimens epimeral plate is more sharply pointed than the many (ZMA). both bear few of short second; a groups setae on prov. Guadalajara: the lateral surface This "baldness", (fig. 13G). — Rio Jalon, on road Nil, near Medinacelli, 27-VII- in with the setation of combination pattern the 1970, clear, fast running stream with sandy bottom,

20 2 CI Ca antennae, make this species look like females of cm deep, m wide, 17°C, pH 6, 4 mg/1, 90 of them in E. Nevertheless it mg/1, many specimens, many pre- berilloni. can be easily recognized copulation (ZMA). by the structure of the hand of the gnathopods. Soria: prov.

— Rio Jâlon, near Sta. Maria de Huerta, 31-V-1970, Female: It is hard to distinguish females of very of them in many specimens, many precopulation this from thoseof E. berilloni. species (ZMA).

prov. Castellon:

Cemented irrigation ditch, alongside road N 340, Ecology and distribution. — So far this species 600 m N. of Chilches, 35 km N. of Valencia, 19-V- has only been found in three localities in the cen- of them in -1971, many specimens, many precopula- tral part of the Pyrenees. Ecological data, other tion (RMNH). Accompanying species: Echinogam- than those mentioned under "material already marus pacaudi. examined" are not available. Description. —

Male: Maximum length 14 mm; in general smaller. echinosetosus Echinogammarus n. sp. Head with rounded lateral lobes (fig. 14A), Figs. 14-15 eyes reniform, more than twice as long as wide.

Material examined. — The peduncle of Al (fig. 14B) is very short in

Spain comparison to the 33- to 40-segmented flagellum.

prov. Zaragoza: and of Peduncle segments 4 5 A2 (fig. 14C) are — Tributary of Rio Piédra, E. of Nuevalos, 13-VII- long and slender, set with groups of short setae 1969, clear, fast running stream, with gravel bottom, (shorter than the diameter of the segments on about 1.5 m wide, pH 6.5, Cl 1 mg/1, Ca 182 mg/1,

of them in which of the many specimens, many precopulation. they are implanted). The segments (The $ 9 and holotype, allotype, many paratypes 19- to 23-segmented flagellum are relatively short have been in the deposited Zoölogisch Museum and bear many short setae near their distal end. Amsterdam, under cat. nr. ZMA Amph. 103.030). The comb of setae along the ventral margin of Accompanying species: E. longisetosus.

— the third of the mandible differs Rio Piédra, 200 m upstream of the Monasterio de segment palp

Piedra, fast with from members in 13-VII-1969, clear, running stream most other in the group having stony bottom, about 10 m wide, pH 6.5—7, CI 13 a few longer setae intermixed with the elements of Ca 80 of mg/1, mg/1, many specimens, many them the normal regular row (fig. 14D). in precopulation (ZMA). The hand of the first 14E, — gnathopod (figs. 15A) Rio Jiloca, near village Morata de Jiloca, 13 km is and has S.E. of Calatayud, 12-VII-1969, moderately polluted piriform a very oblique palm, so that

stream with clay about 5 m CI 45 bottom, wide, there is no obvious palmar angle. Compared with mg/1, Ca 148 mg/1, specimens, of them many many E. berilloni, the medial palmar spine has moved in precopulation (ZMA). towards the articulation with the dactylus. Like- prov. Teruel:

— wise the but from this Rio Jiloca, near Caminreal, 12-VII-1969, clear, fast on palm, widely separated

about m be running stream, 2.5 wide, more than 2 m spine, a group of strong, graduated spines can

CI 34 Ca deep, pH 6.5, mg/1, 110 mg/1, many found towards the basis of the hand. of specimens, many them in precopulation (ZMA). The basal portion of the enormous propodus of Accompanying species: E. longisetosus. P2 is swollen 14F, 15B). The — reaches (figs. very long Upper of Rio Martin, alongside road N the characteristic of the 211, near Vivel del Rio Martin (= 12 km W. of dactylus, implantation

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29

the and the transverse the bottom, nor the or absence of many spines along palm, presence vege-

seem to have influence on its distribu- row of spines on the wide basal portion make this tation, any

within the berilloni- tion, which is restricted to a rather small area in hand most characteristic group. into the Mediterranean The setae along the palm of both hands often bear Spain, draining (see Map Twice it has been found with E. plumosities. P3 and P4 are not very characteristic; III). together another member of the berilloni- their coxa! plates, like those of the gnathopods, longisetosus, and with have rounded inferior corners, and many short group, once together Echinogammarus

setae along the inferior margin (figs. 14G, 14H). pacaudi (see table I).

The basal segment of P5 is hardly more than 1.5

times has backward as long as wide and a Echinogammarus margalefi n. sp.

protruding lobe (fig. 15C). Tn P6 and P7 (figs. 15D, Figs. ÎOE—L, 16.

15E), where the basal segment is more elongate,

Material examined. — this lobe is less pronounced. The armature of these Spain is characteristic, not account of the legs on posi- Teruel: prov. of the tion of the of but because — groups elements, Rio Guadalajar near Albaracin, 6-V-1967, alt. 1150

short almost in these m, specimens (MSNV). very spine-like setae groups. many

— Frias de Albaracin, Monte Universali, 6-V-1967, alt, The inner ramus of the well-developed third 1600 m, many specimens (MSNV). uropod (fig. 15F) sometimes is a little longer than — Tributary of Rio Turia, S. of Libros, about 25 km in most other species in the group; it can attain S. of Teruel, 11-VII-1969, 1 —2 m wide, gravel bot- about of the The setation CI 34 Ca 84 25% exopodal length. is tom, many specimens, mg/1, mg/1.

rather short. (The $ holotype, 9 allotype, and many paratypes have been deposited in the Zoölogisch has of Museum, The telson (fig. 15G) some groups setae Amsterdam, under no. ZMA Amph. 103.024.) Ac- implanted on the dorsal surface of the lobes. companying species: E. longisetosus.

The first urosome has a dorsal excava- — Moron border of Teruel and segment Rio on prov. prov. tion ("saddle") (fig. 141). The metasome, the Castellon, ll-VII-1969, shallow, stony bottom, CI 14 mg/1, Ca 76 mg/1, 12 specimens (ZMA). urosome, as well as the epimeral plates (fig. 14J) Cuenca: prov. are set with the same spine-like setae that can be

— Rio Riânsares, on road N 400, about 12 km E. of

found on the legs (the proposed specific name Tarancón, 10-VII-1969, about 1,5 m wide, stony echinosetosus alludes to this CI 14 Ca 440 feature). bottom, mg/1, mg/1, many specimens

The short spine-like setation, along with the (ZMA).

— Nameless tributary of Rio Teja, near N 420 at enormous gnathopods, and the characteristic Boniches, 11-VII-1969, limestone, CI 29 mg/1, Ca 82 mandible palp, make this species readily identi- mg/1, many specimens (ZMA). fiable. — Rio Teja, tributary of Rio Gabriel, on road N 420

Female: Can be distinguished from females of near Boniches (87 km S.W. of Teruel), ll-VII-1969,

fast stream with about other species by the structure of the mandible palp clear, running stony bottom, 5 CI 33 Ca 74 m wide, mg/1, mg/1, many specimens only. (ZMA).

— Rio Teja, on road N 420, near Huerguina, ll-VII-

1969, about 2.5 m wide, bottom, CI 24 mg/1, Variability. — In addition to the "normal" stony Ca 84 of them in mg/1, many specimens, many pre- unstable characters, the hand of PI shows varia- copulation (ZMA). bility in the number of participating in the spines — Upper reaches of Rio Gabriel, on road N 420, near row along the palm. The relative length of the Salvacanete, ll-VII-1969, about 3 m wide, shallow,

CI 8 Ca 80 mg/1, endopod, in Ur3, varies from 0.1 to 0.25 times the stony bottom, mg/1, many speci- mens, of them in precopulation (ZMA). length of the exopod. many Rincon de Ademuz: prov.

— Rio Turia near Torre Alta, ll-VII-1969, moder-

ately polluted, fast running stream, bottom, — stony Ecology and distribution. As far as we can 132 about 4 m wide, CI 29 mg/1, Ca mg/1, the the many judge at moment, species seems to prefer specimens (ZMA). rather fast middle courses of streams, with running Ciudad Real: prov. of Ca-ions. Neither the of a high amount structure — Rio Guadiana, about 10 km S.W. of Ciudad Real,

14. echinosetosus from second third fourth Fig. Echinogammarus n. sp., ￿, leg (II); F, leg (II); G, leg (II); H, (II). the type locality. A, head (scale II); B, first antenna (II); leg (II); I, urosome (II); J, epimeral plates

C, second antenna (II); D, mandible palp (III); E, first

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Fig. 15 BUDRAOEN TOT DE DIERKUNDE, 43 (1) - 1973 31

limited number of is found on road N 430, 7-IX-1970, slightly polluted, slowly (fig. 161). A setae on

running bottom, more than 1 m deep, stream, stony the dorsal surface of the rather short telson lobes

more than 10 m wide, temp. 25.5°C, pH 6, CI 86 (fig. 16 J). A saddle is present on the first urosome mg/1, Ca 168 mg/1, many specimens (ZMA). 10 The setation of the segment (fig. K). dorsal sur- — Rio Guadiana, about 10 km S.E. of Malagón, face of the and is little 7-IX-1970, slightly polluted, moderately fast running, urosome epimeral plates a

sandy and stony bottom, 20—60 cm deep, 5 m wide, longer than in E. echinosetosus (figs. 10K, L). temp. 25.2°C, pH 6.0, many specimens, some of Except for the armature of the hands of the them in precopulation (ZMA). this much resembles Toledo: gnathopods, species very prov. E. berilloni or E. echinosetosus. Since — road about Rio Tajo, near Aranguez, on N IV, 40 juvenile only adult used for the km S. of Madrid, 26-VII-1970, moderately fast run- sexually specimens were pres-

ning, slightly polluted, more than 2 m deep, about ent morphological studies, there can be no con- of 20 m wide, pH 6—6.5, many specimens, many fusion. In this species precopulation occurs at a them in precopulation (ZMA). observed in E. body size far smaller than ever prov. Valencia:

— Rio Turia, S. of Liria, on road C 3322, near Liria, berilloni or E. echinosetosus.

9-IX-1970, moderately fast running, clear, stony Female: small, without outstanding characters. and clayish bottom, 1 m deep, 5 m wide, temp. Very

CI 78 Ca 132 23.2°C, pH 6, mg/1, mg/1, many Ecology and distribution. — The species is found specimens (ZMA). in the upper and middle courses of streams, drain-

Description. — ing both into the Mediterranean and into the

All these waters have in Male: The smallest species within the E. berilloni- Atlantic (see map III).

that are mineralized, and group; the maximum length observed was 11 mm. common, they highly

rich in Ca-ions. The bottom The lobes of the head have rounded corners; the particularly always

material. small, than twice consists of or other hard eyes are relatively no more as stones, gravel

was found in a few instances It long as wide. The sinus is rather deep (fig. 10H). Vegetation only.

the has wide The first antenna (fig. 10E) has a short flagel- seems that species a very temperature

since it found in lowland lum, with at most 20 segments. The armature is tolerance, was waters at

but also altitude of 1600 in waters very poor. 25.5°C, at an m,

did exceed Peduncle segments 4 and 5 of A2 (figs. 10F, G) where in summer the temperature not

are armed with 3 rows of short elements. The setae 11°C.

dedicated Dr. R. of of the short flagellum are little longer than the This species is to Margalef,

of his excellent diameter of the segments on which they are im- Barcelona, in recognition ecological

planted. work and in particular for his contribution to the

The mandible palp is in so far particular that it knowledge of Spanish amphipods.

bears some setae on the inner surface of the second

sedis segment (fig. 16A). Species inserta

In this comparison with other members of group, In two localities, both in Spain, somewhat outside the hand of both gnathopods 1 and 2 (figs. 101, J) the main distribution area of the Z>en7/ora-group, are small and have a very oblique palm. In both, animals have been found which certainly are closely the medial palmar spine has moved towards the related to E. berilloni. However, they could not be articulation with the dactylus. Very often another, identified since only females or juveniles were smaller spine (indicated by an arrow in fig. 10 J) available. Intensive search, in two successive years, can be found in P2. The inner and outer surface for more material from these localities or in waters of the hands are set with of setae. many groups belonging to the same drainage system, completely The coxal plates 1—4 have rounded inferior because the Rio failed, of pollution (in Duero) or corners, set with some short setules (figs. 16 B, C, because of man-made water works which had D and E). completely changed or destroyed the old localities. Legs 5, 6, and 7 (figs. 16 F, G, H) are relatively

short and armed with smaller spines and short These localities are: setae. — Prov. Cacares: Abadia, 5-V-1960, 7 specimens The third is short, without uropod particularities (RMNH) (these specimens were erroneously record-

15. echinosetosus from of second fifth sixth Fig. Echinogammarus n. sp., ￿, leg (III); C, leg (II); D, leg (II); E,

the type locality. A, hand of first leg (scale III); B, hand seventh leg (II); F, third uropod (III); G, telson (III).

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Downloaded from Brill.com10/08/2021 03:18:23PM via free access Fig. 16 BUDRAGEN TOT DE DIERKUNDE, 43 (1) - 1973 33

ed E. lusitanus 1968 as uy Stock, : 52). in the entire Euro-Asiatic region during the Terti-

— Prov. Zamora: Auxiliary bed of the Rio Duero, 500 ary period. This idea is supported by Basikalova, m W. of the bridge at Zamora, 10-VII-1969, moder- who assumed that the 1945, genus Eulimnogam- fast about 100 ately running, very polluted stream, marus is one of the five old, genera of m wide, CI 30 mg/1, Ca 70 mg/1, 3 specimens primitive (ZMA). amphipods from which all other gammarids are

descendants. She considers the genus Gammarus (meaning in the Gammarus pulex-group, , in DISCUSSION reality

the context of Bazikalova's work) to be one of the

When reviewing the distribution of freshwater other basic groups from which most of the other freshwater have been This gammarids in the Iberian peninsula (see also gammarids derived.

and idea is not at all astonishing since members of this Pinkster, 1971 in press, and Pinkster & Stock, Gammarus be found in the whole 1970 and 1972) it soon becomes clear that two pulex-group' can

main Euro-Asiatic in Northern Africa and distribution areas can be distinguished: (a) region, even

southern inhabited in North America. The holds true for the a one, by members of the same

and since members of this Echinogammarus simoni-group, (b) a northern genus Eulimnogammarus,

~~ be found in Lake Baikal, one with members of the Echinogammarus beril- genus can not only as

assumed, but also in coastal waters loni-group. Both species-groups show only a nar- was previously in the Western and North row zone of overlap; the only exception being Arctic, Europe, America,

E. lusitanus (a member of the simoni-group) which and moreover in many freshwater localities in the

Iberian and in the Pinkster, is distributed in very soft waters in the extreme peninsula Pyrenees (see in press). north-western corner of the Iberian peninsula,

entirely disjunct from the main range of the simoni- The E. berilloni- and E. simoni-species-groups

and in which group, surrounded by an area mem- must be considered more recent invaders, the latter

bers of the E. berilloni-group are the only fresh- coming from the South, the former coming from

water amphipods. the North. This corresponds well with the fact, that

Apart from these two species-groups, both in- the Echinogammarus simoni-group has its main

habiting a more or less demarcated area, members distribution area in northern Africa, which area

of two other of freshwater have in contact with Eurasia groups amphipods was during many periods been all in less isolated relic-like found, more or of its geological history. The close relationship

localities. These are members of the Gammarus demonstrated between North African and Iberian

pulex-group on the one hand, membersof the genus forms of certain fresh-water gammarids, has been

Eulimnogammarus on the other. The nature of the found in many other groups of organisms as well,

in which of these two biotopes populations species- e.g., in several groups of phanerogams, and verte-

and their of in invertebrates such groups use to live, present degree brates, but also aquatic as

isolation to indicate that the subterranean against geneflow seem amphipod genus Pseudoniphar-

have to do with the relics of we formerly more gus, and the isopod genus Sphaerobathytropa (see

widely distributed forms. The extreme morpholo- Brehm, 1947; de Lattin, 1967).

gical variability found within various populations It is almost sure that in the case of the E. simoni- of these Pinkster, for Gammarus there has been invasion species (see 1971, group, not a single only,

and in for members of gauthieri Pinkster, press, but more likely a series of successive invasions

the Eulimnogammarus), are an indication be faunistical the differ- genus or may exchanges during

that have to dowith isolated we small, populations, ent periods during which connections between the

little numerous that the total is being so gene pool African and European continents did exist. This reduced and that homeostasis (through suppressor also explains the ppecial position of E. lusitanus

genes) is weakened. Pinkster, in press, pointed out in the N.W. part of the Iberian peninsula. This

that these Spanish as was the first member of the Eulimnogammarus-species, species probably group relatives well as their in Lake Baikal, are descend- that invaded the peninsula and lost contact with of the old limnic ants same fauna, that occurred its relatives because of the formation of a sea-arm

16. Fig. Echinogammarus n. from the (II); F, fifth leg (II); G, sixth seventh margalefi sp., ￿, leg (II); H, leg (II); type locality. A, mandible palp (scale III); B, first leg I, third uropod (III); J, telson (III). (II); C, second leg (II); D, third leg (II); E, fourth leg

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between N. Africa and Spain in the valley of the calvus, or E. longisetosus occur in the faster

for instance of these Such present Guadalquivir as happened running, upper courses systems. a

during the Burdigalien (Sollaud, 1939; Mélendez, series of subsequent invasions and successive isola-

1965). Once it was isolated, it could develop into tions thus created favorable conditions for the

that this a highly specialized species, could successfully developing of numerous species within group,

itself invade and maintain in the very poorly all of them with their own, more or less restricted

mineralized and acid waters in the north-western distribution area.

part of the peninsula, a region that did not change In some regions, especially in the western Pyre- distribution since palaeozoic times. Regarding the nees, where many different types of biotopes can

it and pattern of the Echinogammarus berilloni-group, be found, where climatological conditions are

that the western of the and this resulted in distribu- appears parts Pyrenees not too extreme, finally a

the adjacent parts of Spain and France form the tion pattern which is almost similar to that en- within this centre from where most species group countered in the Gammarus pulex-group in other

of originated. parts Europe, including the typical ecological

Since E. live without ill-effects in zonation river berilloni can throughout the system.

80% sea-water (Vincent, 1971) and since it can be In areas with more severe climatological con-

found in estuaries and even on the open beach, on ditions, e.g. the interior of the peninsula with very

condition that absent hot the of the lower the competing species are summers causing desiccation

and with cold winters which (present work, own observation) we must assume stretches, very during

that in this case we have to do with invaders from the most upstream stretches of the river freeze,

the sea. a clearly developed zonation is usually absent and

the distribution Most probably, when regarding the number of species found within one system is

of within reduced pattern the various species this group, usually to one only.

there has not been one single invasion, but a series A problem which is still unsolved is the recent of subsequent invasions. The occurrence of E. discovery of E. pacaudi in a temporary irrigation

in lower stretches of the river berilloni s.str. the ditch, near Chilches in the province of Castellon.

the As this from rather systems are an indication that this species is yet, species was known a

most recent invader. This is supported by the restricted area (from the northern slopes of the

distribution pattern in the western Pyrenees, a Pyrenees to the river Garonne) only. In spite of

several this region where species, belonging to intensive sampling, the species never has been and where E. berilloni inhabits group are found, found in between these two widely separated areas.

always the more downstream parts of the river Maybe, further and more intensive sampling will

while other E. E. the this systems, species as aquilifer, provide answer to new question.

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Table 1

of members of the E. berill Sympatric occurrence oni- group

species found in 1 sample with found in the same stream system with

E. berilloni E. aquilifer E. aquilifer

E. longisetosus E. longisetosus

E. zebrinus E. zebrinus

(E. pacaudi) (E. pacaudi)

E. feminatus E. calvus

E. feminatus

E. calvus E. longisetosus E. longisetosus

E. berilloni

E. margalefi E. longisetosus E. longisetosus

E. longisetosus E. berilloni E. berilloni

E. echinosetosus E. echinosetosus

E. aquilifer E. aquilifer

E. margalefi E. margalefi

E. calvus E. calvus

E. echinosetosus E. longisetosus E. longisetosus

(E. pacaudi) (E. pacaudi)

E. feminatus E. berilloni E. berilloni

(E. pacaudi) (E. pacaudi)

E. zebrinus E. berilloni E. berilloni

(E. pacaudi) (E. pacaudi)

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Amsterdam, 1 (14) : 205—219.

Received: 2 June 1972

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Iberian

the

in

1971

and 1970

1969,

grid Sampling

I. Map peninsula.

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Map II. The distribution of Echinogammarus berilloni (Catta) in Western Europe (hatched area plus the encircled Channel Islands).

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