Bijdragen tot de Dierkunde, 50 (1): 105-144 — 1980

Regression model evolution as exemplified by the genus

Pseudoniphargus (Amphipoda)

by

Jan H. Stock

Institute of Taxonomie Zoology, University of Amsterdam, The Netherlands

Abstract A forma described in 1955 by S. Karaman, was

raised to specific rank by G. Karaman in 1978. The has been considered genus Pseudoniphargus long mono- In the I have revised of the present paper, most specific. Its unique species, Ps. africanus, was supposed to

materials attributed to the as well occur both sides of the the Atlantic existing on Mediterranean, on genus,

Iberian Ma- side of the peninsula, on the Azores, and on as a good number of newly collected samples. This deira, in localities from the shore than ranging sea to more nine named resulted in a classification into species, 1000 m of altitude, and covering almost the entire natural

and a number of unnamed left-overs because of salinity range (0-36‰).

A taxonomie revision revealed that at least nine named insufficient material. The actual names for the

species and several unnamed forms (of which insufficient taxa of Pseudoniphargus, formerly usually recorded material is available) hide under the name Ps. africanus, Ps. shown in table each with and as africanus, are I. a narrow ecological geographical range. The evolutionary scenario of the members of the is Like in the Gammarus identification of genus group, discussed of marine at some length: they are presumably females is much more difficult than that of males. origin, and got adapted to conditions of continental waters both sexes should be available for a during various marine regressions in the Eocene and Oligo- Preferably

cène, but notably in the Miocene. profitable study. Much of the "variability"

recorded in the has been based past on secondary

1. Contents sexual differences within one species.

The evolution of — devel- 2. Introduction 105 Pseudoniphargus the

105 3. Acknowledgements opment of inhabitants of inland waters from 4. model evolution Regression (generalities) . . . 107 marine ancestors — is discussed at some length. 5. Evolutionary scenario and zoogeography of Pseudo- It is that this evolution is of the niphargus 108 likely regression

6. Cladistics 113 model type: stranding of marine populations 7. Place of Pseudoniphargus at level 114 family . . . during sea-level regressions. 8. Taxonomy 116 The has been undertaken to 8.1. Genus Pseudoniphargus Chevreux, 1901. present study com-

Diagnosis 116 the ideas forward pare previously put (Stock, 8.2. Key to the species 116 model ideas 1977b) on regression evolution, 8.3. Description of the species 117 based obtained in the West 9. References 142 mainly on evidence

Indies, with data derived from the Mediterranean

marine area where theories on the stranding of 2. INTRODUCTION populations were first developed.

Since the erection of the genus Pseudoniphargus

Chevreux in almost student of it 3. ACKNOWLEDGEMENTS by 1901, every

has stressed its "morphological variability" and its The author is indebted the and to following persons insti- striking distribution, from the sea shore to caves tutions for providing information or material: at considerable altitudes, from the South European Curators Lennart Cederholm and Hugo Andersson, Zoolo- giska Institutionen, Avdelningen för Systematik, Lund, Swe- and North African continents to oceanic islands den (abbreviated ZIL). like the Azores and Madeira. Dr. E. L. Bousfield, National Museums of Canada, Museum

the remained of Natural Sciences, Ottawa, Canada. Up to 1955, genus monotypic. 106 J. H. STOCK - REGRESSION MODEL EVOLUTION

Table I

Previous records of Pseudoniphargus in chronological order. Unless otherwise stated, all records have been published under

the name of Ps. africanus.

Name in Author, year, page(s) Origin present paper

(t = material re-examined;

* = inferred name)

Chevreux, 1901: 211-216, figs. 1-2 Bône (= 'Annaba) (Algeria) Ps. africanus

Medjez-Sfâ (Algeria) Ps. macrotelsonis

Schellenberg, 1937: 224 CantabrianMts. (Spain) Ps. elongatus

Ps. unisexualis

1939: Bône Ps. Schellenberg, 297-304, fig. 1, 1 map (Algeria) africanus

Medjez-Sfâ (Algeria) Ps. macrotelsonis

Mustapha, Algiers (Algeria) t Ps. sp.

Cantabrian Mts. (Spain) Ps. elongatus

Ps. unisexualis

Split (Yugoslavia) Ps. adriaticus

Funchal Ps. (Madeira) t sp.

Schellenberg, 1943: 1 Cost?, da Arrâbida (Portugal) Ps mateusorum

Schellenberg, 1951: 327-328 Bonifacio (Corsica) Ps. adriaticus

Balazuc & Angelier, 1951: 309-312 Banyuls (France) Ps. adriaticus

S. Karaman, 1953: 139 Dubrovnik (Yugoslavia) Ps. adriaticus

1953: 672 Ps. Ruffe, Philippeville (= Skikda) (Algeria) sp.

Balazuc, 1954: 184-185 Banyuls (France); Bonifacio (Corsica) Ps. adriaticus

44-51 Budva Ps. adriaticus S. Karaman, 1955: 236-241, figs. Dubrovnik, (in text) or Bar (in legends)

(as formaadriatica)

Dahl, 1958: 6-9, fig. 1 Faial, Pico (Azores) Ps. brevipedunculatus

Ruffo, 1960: 172-174 Menorca (Balearic Is.) Ps. adriaticus

Strinati & Coiffait, 1961: 228 Menorca (Balearic Is.) Ps. adriaticus

Sket, 1969: 148 Rupine near Sukosan (Yugoslavia) Ps. adriaticus

Margalef, 1970: 170-171, figs. 1, 2 J-L Basque country (Spain) Ps. unisexualis

I Pi- sp.

Mateus & Afonso, 1974: 10-11 Santa Maria (Azores) Ps. brevipedunculatus

Ginet, 1977: 174-175 Cantabrian Mts. (Spain) Ps. elongatus

Mateus & Mateus, 1978: 16-19 Costa da Arrâbida (Portugal) Ps. mateusorum

Prov. Beira (Portugal) Ps. longispinum

Prov. Estremadura (Portugal) Ps. Sp.

Madeira Ps. sp.

Azores Ps. brevipedunculatus

G. Karaman, 1978: 250-256, figs. VI-IX Bône ( = 'Annaba) (Algeria) Ps. africanus

Oued-ed-Demane (Tunisia) Ps. sp.

Medjez-Sfâ (Algeria) Ps. macrotelsonis

de Cueva San Adrian near Cegame (Spain) Ps. unisexuali s

Do.: 241-250, figs. I-Y (as Ps. adriaticus) Budva (Yugoslavia) Ps. adriaticus

Mola di Bari (Italy) Ps. adriaticus

Isola di Montechristo (Italy) Ps. adriaticus

Funchal (Madeira) Ps. sp.

Menorca (Balearic Is.) Ps. adriaticus

Mustapha, Algiers (Algeria) t Ps. sp.

Prov. Santander (Spain) Ps. elongatus

Split (Yugoslavia) Ps. adriaticus

Dubrovnik (Yugoslavia) Ps. adriaticus

Sukosan Rupinenear (Yugoslavia) Ps. adriaticus

Costa da Arrâbida (Portugal) Ps. mateusorum

Pesce Ps. Bari Ps. et al., 1979: 8 (as sp.) (Italy) sp. BIJDRAGEN TOT DE DIERKUNDE, 50 (l) - 1980 107

the Mme. N. Coineau, Laboratoire Arago, Banyuls-sur-Mer, models, original distribution area of a taxon

France. is divided, the barrier being small at the beginning, Dr. J. Forest, Muséum National d'Histoire Naturelle, Labo- but effective at a later model is ratoire de Zoologie, Arthropodes, Paris, France (MNHN). 100% stage (this

Prof. R. Ginet, Département de Biologie Animale et Zoologie, particularly useful in understanding the develop-

Université Claude-Bernard (Lyon-1), Villeurbanne, France ment of populations isolated under influence of (UL).

Dr. H.-E. Gruner, Zoologisches Museum der Humboldt Uni- plate tectonics). versität, Berlin, D.D.R. (ZMB). A good regression model taxon (genus,

Prof. A. Mateus, Universidade do Porto, Instituto de Zoolo- family...) must satisfy the following five condi- gia "Dr. Augusto Nobre", Porto, Portugal (IZAN). tions: Curator G. E. Maul, Museu Municipal do Funchal, Madeira.

Mr. Jos Notenboom, Biospeleo Werkgroep, Wageningen, The (1) In addition to the species in continental Netherlands. waters, marine relatives must be known. Dr. S. Pinkster, Zoologisch Museum, University of Amster- 2 should occur in areas once covered dam, The Netherlands (ZMA). ( ) They only travel of the University of Amster- above the surface. Furthermore, a grant by sea, but actually dam is (1979) gratefully acknowledged. should outside sub- (3) They not occur formerly remarks made model elements trans- The on regression in merged areas, since then dispersal from continental gression areas are based on unpublished observations made financed ruled during an expedition to the southern Bahamas, by areas to areas emerging cannot be out. the Netherlands Foundation for the Advancement of Tropical (4) They are lacking, or at least exceptional, in Research (WOTRO), The Hague, and the Beijerinck-Pop- areas that show ping Fonds, Amsterdam. transgression (subsidence). If is (5) there more than one inland taxon, these

being developed through independent passive isola- 4. REGRESSION MODEL EVOLUTION tion marine the inland tend from ancestors, taxa (GENERALITIES) to have cladistic relationships indicative for the

model and for the active Several authors (e.g. Bonnet, 1956; Hubault, passive (vicariance) not

model 1938; Fryer, 1965) have emphasized the prob- (dispersal) (see Rosen, 1976, fig. 20). These five be adstructed follows: ability that populations of littoral or shallow-water points may as

in inhabitants of the Ad The marine relatives should animals, more particular 1) by preference

interstitia of stranded to the same the limnic macroporous substrates, be- belong genus as species,

but do have be of the cause of various marine regressions in the Medi- they not per se to ancestors

limnic since the marine forms have terranean region during the Tertiary epoch. These species, may

stranded the further evolution after the isolation of populations got gradually adapted to undergone

conditions of more continental waters and are now the limnic taxa.

Ad 2 Several this distributed in land areas that were previously sub- ) examples prove to a surprising the distributionof the Thermosbaenacea merged. extent, e.g.

Stock (1977b), studying groundwater Amphi- (Fryer, 1965; Stock, 1976), of Typhlocaris (Fryer, of the poda of the hadziid group in the Antilles, accepted 1965), Microparasellidae (Stock, 1977a),

of the hadziid Gammaridae and tectonic uplifts (or sea-level drops) as one of the (Stock, 1977b),

main the evolution several of the causes for of groups genus Pseudonipbargus (present paper). Ad of and thermosbaenacean Crus- 3 on theother that isopod, amphipod ) This, hand, implies regres-

in sion model elements have of tacea stygobiont habitats of the various islands. poor means (active)

For this type of evolution, through stranding of 1 ) Passive model: parts of a population are isolated by marine ancestors, Stock coined the term regression forces; barriers, extrinsic not existing before, come to develop- model evolution. this is a Although essentially ment, dividing or fragmenting the original population. This

model often called passive model of evolution 1 ), the regression is vicariance model nowadays in the New World, but I believe that the vicariance of the biogeographers model differs from the other passive models in of the nineteen-thirties and fourties not only "preoccupies" that the barrier between the ancestral marine and the name, but also had a somewhat different meaning (the derived limnic homonyms are not synonyms). populations developed very slowly Opposed to passive models are active models, according to and and gradually never have been 100% may which of to parts populations actively disperse new, pre- effective flow. In other against gene passive viously uncolonized, areas. 108 J. H. STOCK - REGRESSION MODEL EVOLUTION

For ancestral dispersal. several of them, poor dispersal ments. Apparently marine and polyhaline

means have been made plausible (Remane, 1952; Gammarid species flocks have given rise to more

Delamare than invasion into the continental Deboutteville, I960; Stock, 1979). one epigean

the is low and I the continental Usually number (1 to 4) of eggs waters. Likewise, assume that

pelagic stages are absent (iC-strategists). hypogean waters have been populated along more

is Ad 4) One of the world's largest transgression than one pathway. The word "pathway" used

to be the Bahama here in since under that name (subsidence) areas appears plat- a general sense, may

form (Doran, 1955; Dietz et al., 1970; Uchupi hide various processes: (1) active dispersal,

which is underlain carbonate marine substrates et al., 1971), by a through macroporous (inter-

is than 5 km into the apron that more thick, maybe even stitia) hypogean habitat; or (2) passive

10 km. Extensive search in the Bahamas (own (vicariant) development from epigean populations

observations, unpublished) has shown a striking isolated in groundwater accumulations during

poverty in regression model elements, sharply con- periods of drought or frost when surface waters

trasting with the abundance of such elements in disappeared.

like and various The of member of regression areas Cuba, Hispaniola presence a Pseudoniphargus

Antilles. in Lesser the Azores makes it, as Dahl (1958: 9)

Ad Each isolate in model situation stressed that... 5 ) a regression rightly "highly probable (they) ...

from marine it is reached the Azores of the since the is derived a ancestor; even not by way sea",

has unlikely that the same ancestral marine line "current geological opinion seems to be in favour

than In their oceanic the produced more one stranding. a dispersal of purely origin, probably as

is model situation, a stepstone-like phylogeny more result of submarine volcanic activity in the Mio-

of continental connections likely. cene "proof ap-

in model not to be available". The The most striking point regression evo- pears same arguments

lution is that continuous contact has to the of the seemingly apply presence genus on Madeira,

been possible between the stranded populations and they seem to be supported amply by recent

and the ancestral marine ones. This point will be geological evidence (e.g. Mitchell-Thomé, 1976

in the next section of the and picked up again present 1979).

It that Balazuc & have paper. appears Angelier, 1951,

In model evolution in addition to regression been the first to speculate on the possibility that

groundwater animals, very similar phenomena are members of Pseudoniphargus originate from ma-

known from bird that isolated rine which in the of populations got on ancestors are process adapta-

mountain peaks during orogenesis (see for in- tion towards life in fresh waters. They recorded

Ps. in stance Haffer, 1974: 149). a Pseudoniphargus (as africanus but reality

Ps. from the close " adriaticus) psammon (ll/ 2 5. EVOLUTIONARY SCENARIO AND 6 m) to the sea near Banyuls at salinities from ZOOGEOGRAPHY OF PSEUDONIPHARGUS 24 to 33°/oo- Recent research (present paper),

to the Gam- the Bou-Rouch Pseudoniphargus shows, as compared using phreatic pump technique,

the Ps. in the inter- marus group, a number of apomorphic characters, revealed presence of adriaticus

of both the for instance: disappearance the eyes; reduction stitia of a coarse sand and gravel, between

in size and in number of articles of the tide marks and below low tide accessory just at full marine

of flagellum of the first antenna; abbreviation the salinity (36°/oo)> on the French Provence coast.

Karaman and his second antenna; tendency towards elongation of Later, both S. (1955) son

the third uropod; small body size; suppression of G. Karaman (1978) advocated the view that

in the sexual dimorphism gnathopods; absence of Pseudoniphargus must have a marine origin.

the second I think it that ancestral calceoli on antenna. likely Pseudoniphargus

In lines of such several hypogean Gammaridae (s.l.), lived in macroporous marine substrates, as

these characters have developed independently; gravel or coarse sand, and had a large distribution this can be called convergent or parallel develop- in the early-Tertiary Tethys Sea. Great portions of BIJDRAGEN TOT DE DIERKUNDE, 50 (l) - 1980 109

the northern Mediterranean littoral (France, Cor- These ideas contrast sharply with thoseof Schel-

sica, the Balearic Isles, Yugoslavia) are inhabited lenberg (1939), whoconsidered all populations of

called by morphologically identical populations, Pseudoniphargus as a single species, irrespective

found in localities Ps. adriaticus. This is the only member of the whether they were at the sea-

known far from in altitude of 1000 genus so marine/polyhaline shore, caves at an m, or on

waters. However, morphologically, Ps. adriaticus isolated oceanic islands as Madeira. Schellenberg

does well ancestral considered the be "restricted not classify very as an (or genus to to fresh

plesiomorph) form (see the section on cladistics). waters", "frequently remote from the sea", and

Since Ps. adriaticus has toler- him the a fairly large salinity this brought to often-quoted statement

ance (Balazuc & Angelier, 1951; present paper), that “Pseudoniphargus is not a marine immigrant,

invad- it is more likely one of colonization waves but an old freshwater element that reached its

ing the continental waters, instead of being the present distribution" (northern Mediterranean ancestral form from which all continental and belt, Atlantic side of Spain and Portugal, Balearic

oceanic insular taxa evolved. islands, Algeria, Corsica, Madeira, the Azores),

Fig. 1. Distribution of the genus Niphargus (hatched) and of the genus Pseudoniphargus (dots = taxa from continental

waters; = from Note that there is the of triangles taxa marine/polyhaline waters). hardly any overlap between range

Niphargus in Europe and that of the inland water taxa of Pseudoniphargus. On the other hand, the marine/polyhaline

Pseudoniphargus does occur in Niphargus country.

The of is the of Ruffo Morand-Chevat and Ginet range Niphargus based on publications (1956, 1957), (1972), (1977);

the of is based the data in the unisexualis range Pseudoniphargus on presented present paper. 1, Pseudoniphargus n. sp.; 2,

mateusorum branchiatus Ps. elongatus n. sp.; 3, Ps. longispinum n. sp.; 4, Ps. n. sp.; 5, Ps. n. sp.; 6, Ps. brevipedunculatus

Ps. Ps. macrotelsonis Ps. adriaticus Materials that had n. sp.; 7, africanus Chevreux, 1901; 8, n. sp.; 9, Karaman, 1955.

to remain unnamed are indicated by a crosslet.

Inset bottom right: a male of Pseudoniphargus adriaticus. - 110 ). H. STOCK REGRESSION MODEL EVOLUTION

where still linked with each "when these areas (= Skikda), Algeria and from Mustapha,

other". Schellenberg's statement (1939: 300, in Algeria, Ps. branchiatus, Ps. mateusorum, Ps.

that "the therefore, is not the and Ps. from the translation) sea, longispinum, sp. province bridge between the Mediterranean localities and of Estremadura, Portugal.

Madeira, but the ditch" is not supported by geol- (3) During the Oligocene (fig. 3), most of

and in ogical evidence whatsoever, disproved by these localities were either submerged and

the inter- and discovery of an subtidal species, far off-shore (Ps. branchiatus, Ps. macrotel-

adriaticus. Ps. from land Pseudoniphargus sonis, sp. Mustapha), or on dry

The for div. greatest handicap reaching zoogeo- (Ps. sp. from Portugal).

has The Azores and graphical conclusions been the fact that almost (4) Madeira are oceanic islands

author took it for and every up to now granted that formed by volcanic activity, broke through

Start- surface the in the late Pseudoniphargus was a monospecific genus. the of ocean Oligocene

ing with Chevreux (1901) at the creation of the or early Miocene (Dahl, 1958; Mitchell-

several authors genus, (e.g. Schellenberg, 1939; Thomé, 1976). Subsequently these islands

Ruffo, I960; Dahl, 1958) stressed the mor- underwent a considerable (several hundreds

phological "variability" and ecological plasticity, of metres) uplift (Mitchell-Thomé, 1976).

marine beaches and is ranging from to caves springs It clear that the nonmarinepopulations of

in the mountains. No took the high one step of Pseudoniphargus existing on these islands

trying to split the generotype, Ps. africanus, into could only have evolved after the formation

and of the Miocene geographically, ecologically morphologically terra firma, i.e. in or later.

more coherent units, with the of the 5 The in northern exception ( ) Pseudoniphargus Spain may

Karamans oldest (1955, 1978) who recognized as be the continental populations. Their

distinct the marine/polyhaline form and Mateus localities (two well-distinguished species and who & Mateus (1978) suggested to start a taxo- one population of uncertain taxonomie posi- nomie of certain analysis populations. tion) were submerged under seawater during

It is the Eocene from (unknown) marine, interstitial an- (fig. 2) but emerged during the that the brackish-water cestors, various and inland rest of the Cenozoic. It might not be acciden-

evolved. this tal in this that populations supposingly Apparently, context one of these, Ps. uni- evolution took in different place geological sexualis, is the most plesiomorphic taxon

various discontinuous the periods, by waves. So, actually known, and at the same time recorded animals from the the high mountains of N.W. Spain from one of greatest altitudes known for

are different from those of lower the 1000 morphologically genus (about m above sea level).

altitudes, km In these although living only 30 remote; the general, distribution patterns fit in with

same holds for the mountain and lowland the of true findings Hubault (1938), Bonnet (1956),

in How old various populations Algeria. the popu- Fryer ( 1965) and others, all in the circum- medi-

lations in continental waters are, is a matter o terranean region, and of those of Stock (1976, When check the distribution speculation. we of 1977a, 1977b) in the Antillean region. Adaptation landmasses and sea various to continental during geological waters appears to have taken place in the epochs Tertiary period in correlation with at different periods of marine regression in the

actual localities the for nonmarine Middle the actual localities of Pseudoniphar- Tertiary: many gus (figs. 2-6), we observe some highly suggestive hypogean freshwater crustaceans (the isopods situations: Monolistera and Typhlocirolana, the prawn All localities for 1 ) are on the ( Pseudoniphargus dry Typhlocaris, mysids Stygiomysis and Spelaeo- land during the Pliocene (fig. mysis, of the hadziid 6). , gammarid amphipods group,

(2) During the early Miocene (fig. the loca- or of the 4), genus Pseudoniphargus, microparasellid lities of the following are located on of the Microcharon the order species isopods group, Ther- the of land edge and sea: mosbaenacea... are to Ps. africanus, Ps. ) all close Tertiary Tethys

macrotelsonis, Ps. from shore lines. All found in sp. Philippeville are areas that were sub- BIJDRAGEN TOT DE DIERKUNDE, 50 (l) - 1980 111

2 5

3 6

Figs. 2-6. Approximate distribution of land and sea in four

periods of the Tertiary: the Eocene (fig. 2), the Oligocene

(fig. 3), the early Miocene (fig. 4), the late Miocene

(fig. 5), and the Pliocene (fig. 6). The actual localities for

indicated Pseudoniphargus in continental waters are by dots.

Palaeogeographic data based on Fryer (1965), Termier &

Termier (1960), and Hsü (1972).

4 112 J. H. STOCK - REGRESSION MODEL EVOLUTION

in earlier dian merged beneath the sea Tertiary epochs. islands. Tectonic uplifts played no doubt a

in Since the Pliocene, all these localities are dry more important rôle, but apart from a sharp drop foundoutside the land areas, and none is previously in the sea surface temperature in the Miocene, and

which rules active the submerged areas out dispersal from volcanism in late Tertiary and Recent

elsewhere. Probably the most intensive period of epochs, the area seems to have been much more

"stranding" of these organisms of marine origin stable than the Mediterranean.

has been the late Miocene, when the quasi-totality Another point must be stressed here, first

of the Mediterranean Sea closed off from the S. Karaman It was brought up by (1955: 240-241).

Atlantic, evaporated and transformed into arid seems significant that inland-water populations of

land This in those (Hsü, 1972, and present paper fig. 5). Pseudoniphargus have only been found

level with immense regression (the water dropped areas where "old" hypogean freshwater taxa are

more than 3000 m and the erosion levels of the absent. Karaman mentioned this from his own

such Rhône and lower- in I his larger rivers, as Nile, were experience Yugoslavia, but can confirm

must ed by hundreds if not thousands of metres) observations from two areas, that I intensively

have caused the stranding of great numbers of sampled myself, viz. Mallorca (Balearic Islands)

formerly marine, shallow-water inhabitants. and the French Provence.

It There can be no doubt, and numerous taxo- can hardly be a coincidence that limnic or

of animal inland nomie papers covering a wide range Pseudoniphargus species are known from

that the forms from the Madeira and the groups testify this, con- Algeria, N.W. Spain, Portugal,

least devoid of ancient limnic tinental waters are differentiated at on spe- Azores, members of cific (and often also on generic) level from those groundwater genera. On the other hand, from of marine and near-coast habitats. The question France, Corsica, the Balearic Islands and Yugo-

where the inland well-stocked remains what caused the interruption in gene- slavia, waters are

marine and with flow between the continental popula- ancient forms as Niphargus or crangonyctids,

The coastal tions. above-mentioned late-Miocene Medi- only populations of Pseudoniphargus are terranean crisis might have caused such an inter- known. Fig. 1 shows the actual distribution of ruption. Most of the Mediterranean basin dried Niphargus (hatched) in Europe. The only place

whereas the little that remained was in of with limnic of Pseudo- up, water overlap populations

of nature is narrow zone in the general hypersaline (Hsii, 1972), pre- niphargus a Spanish prov-

inces sumably not suitable to support a varied crusta- Navarra, Guipuzcoa, and Vizcaya. This over-

I that the of the be the result of of cean life. suppose great majority lap may dispersal Niphargus

marine the corridors Tethyan ancestors of the regression ele- along well-known on either end of ments became extinct, whereas the continental the Pyrenees.

in interstitial and populations hiding groundwaters This peculiar, exclusive, distribution pattern can could survive. The extinction idea is corroborated be in two different explained ways: by the absence in the actual Mediterraneanbasin of ( 1 ) The invasionof the limnic domain by Pseudo- marine such hadziid Am- is than the regression elements, as niphargus more recent appearance

in West that phipoda, or Thermosbaenacea, whilst the of Niphargus, implying Pseudoniphargus

Indies, the only other area in the world where could only invade those areas where the

faunas studied regression have been to some ex- competing genera were absent, or tent, marine (not necessarily ancestral) relatives ( 2 ) the older limnic populations of Pseudoniphar- have been found. have been from gus expelled their fresh-

This is in agreement with the fact that the West water habitats by more recent migration

Indian marine fauna has not suffered from a waves of Niphargus.

crisis like the Mediterranean. Certain indications in favour of salinity/evaporation are the first ex-

On the other hand, it is more difficult to envisage planation:

of barrier what kind has caused the evolution of ( 1 ) Several Pseudoniphargus taxa still prefer the elements the various West In- such regression on near-coast localities (e.g. present paper), - 113 BIJDRAGEN TOT DE DIERKUNDE, 50 (1) 1980

of the distal localities for Niphargus are exceptional (Sket, condition, resulting from a reduction

is found 1977). segment, though rare in Echinogammarus,

of relati- for instance in E. sicilianus monomerus (2) The presence Pseudoniphargus on ssp. Stock,

oceanic islands Ma- It is not vely young, (Azores, 1978). sexually dimorphous, except per-

is deira) and the absence of Niphargus on such haps in the setosity/spinosity; the pedunculus

the is not islands, points to a fairly recent development short, exopodite strongly elongated,

of Pseudoniphargus from marine ancestors. straight (not upcurved), and the endopodite is

( 3 ) The morphology of Pseudoniphargus is more short, triangular. This plesiomorphic state is

encountered and in Ps. Gammarus-like, thus more similar to marine in Ps. unisexualis longispi-

taxa, than that of Niphargus. num.

of the male I would like to stress that the Zoogeographie The next step is that the exopodite

distribution of limnic members of the Gammarus third uropod gets elongated and upcurved, the

is that Such female remains in the state. group not unlike of Pseudoniphargus. uropod plesiomorphic

The found in male Gammarusses (5.1.) are likewise known from Ma- exopodal spines, the plesio-

deira madeirensis are setules. (Sarothro gammarus (Dahl, morphic state, partially replaced by This in and Ps. 1958)), from the Azores ( S . guernei (Chevreux, stage is found Ps. mateusorum

from simoni 1889)), Algeria (Echino gammarus brevipedunculatus.

(Chevreux, 1894) and others), from N.W. Spain Then follows a gradual elongation of the pedun- ( Echinogammarus berilloni-complex, vide Pink- culus of the male third uropod (the female third

and lusit still This ster, 1973), Portugal (Ech. anus (Schel- uropod stays unchanged). stage is encoun-

lenberg, 1943) and others). Although members tered in Ps. africanus, Ps. macrotelsonis, and Ps.

of the Gammarus have much distri- adriaticus. group a larger

bution than members of the The most state is reached Ps. genus Pseudoniphar- apomorphic by it be that the and Ps. branc where also the gus, might significant notably eury- elongatus hiatus,

could shows haline gammarids follow, presumably at a female third uropod a distinct elongation.

Other lines more recent geological period, the same invasion evolutionary (development from a

into lines as For the an be Pseudoniphargus. gammarids, a plesiomorphic apomorphic state) may marine has been and I in the Bousfield table origin never doubted, present genus. (1977, II)

neither doubt such an origin for Pseudoniphargus. lists the plesiomorphic and apomorphic condition

of various characters, the telson Note added at the proof-reading: Recently (December 1979), including (deep-

biol. = B. Sket (Bermuda Stat. vol. nr. 2/3) cleft shallow-cleft = Newsletter, 7, plesiomorphic, apomor- recorded an unnamed member of Pseudoniphargus from phic), the shape of the basis of P5 to P7 (with In Walsingham caves, Bermuda. the present paper, Sket's lobe = without could not be taken strong posterodistal plesiomorphic, discovery yet into account. The occurrence, on the other side of the Atlantic, forms quite a lobe = the strong support apomorphic), presence (plesiomor- for idea my that Pseudoniphargus invaded the inland waters phic) or absence (apomorphic) of a basofacial from some marine stock. spine on uropod 1.

6. CLADISTICS Table II summarizes the distribution of plesio-

and apomorphic conditions in the Pseudo- The peculiar elongation of the third uropod, which genus brought Chevreux (1901) to the rather unfor- niphargus.

Note that the marine form Ps. tunate name Pseudoniphargus, is not present in only known, is all within A clear adriaticus, certainly not the most taxa the genus. evolutionary plesiomorphic

one in this line. line is visible from taxa with plesiomorphic to taxa

shows a solution with highly apomorphic third uropods 2 ). The Fig. 7 (left) phylogenetic

the nine taxa of and is of more less involving Pseudoniphargus plesiomorphic appendage a or gener- alized the six characters discussed above table Echinogammarus-type, on the understand- (cf. II).

that has It is clear that character state 4a occurred twice ing the exopodite only one segment (this in the independently cladogram, character state 5 a

2 ) A similar be observed in development can Niphargus. four times, and character state 6a twice. 114 J. H. STOCK - REGRESSION MODEL EVOLUTION

Table II As to the cleavage of the telson (character 5),

I with Bousfield that an Distribution of plesiomorphic (p) and apomorphic (a) agree (1977) entirely conditions in the genus Pseudoniphargus. cleft telson represents a plesiomorphic condition,

but in the of the the case genus Pseudoniphargus, Character * 1 2 3 4 5 6 telson is with less entire a deeper or deep terminal

the notch well notch, and the depth of may be of longispinum a p p p p p value. brevipedunculatus a p p p p p not too great cladistic

mateusorum a p p p a p Based on these considerations, the characters 4 unisexualis p p p a a p to 6 have been omitted in the phenogram presented adriaticus a a p p p a in africanus a a p a p p fig. 7 (right). This phenogram shows a more

elongatus a a a p p p consistent picture. branchiatus a a a p a p

macrotelsonis a a p a a a

7. PLACE OF PSEUDONIPHARGUS AT

* — unmodified and similar Character 1 p: uropod 3 S to FAMILY LEVEL uropod 3 9 .

transformed. a: uropod 3 $

The is one of those Character 2 — jr. only the exopodite of uropod 3 $ modi- genus Pseudoniphargus genera

fied (= elongated). that does not fit into Bousfield's ( 1977) elaborate

a: both exopodite and pedunculus of uro- attempt to subdivide the large of the Gam- modified group pod 3 5 (= elongated).

maridae into a number of Character 3 —■ p: uropod 3 9 unmodified. superfamilies, family

3 9 modified a: uropod (= elongated). groups and subfamily groups, almost as numerous Character 4 —• P7 with posterodistal lobe. p: there as are genera. a: P7 without posterodistal lobe.

He treats in two lines on Character 5 — p: telson deep-cleft. Pseudoniphargus

telson shallow-cleft. his but his is a: p. 304 of account, treatment obscured 6 Character — /;; uropod 1 with basofacial spine. (as more often in the same paper) by several slips 1 basofacial a: uropod without spine. of the the is attributed "Mo- pen. Firstly, genus to

of It is possible to construct a number of alternative nod, 1925" (instead Chevreux, 1901). Sec- possibilities for this nine species/six character ondly, the type-species is called “Gammarus afri-

problem, but always conflicting situations occur. canus Chevreux, 1901", although Chevreux never

This is a strong indication that certain character described such a combination. In the third place

showed "several brackish water" states a parallel development in the genus. subspp., Mediterranean,

I do believe that the of the third attributed the the development were to genus. However, greater

from "normal" into of the uropod, a gammarid type an part the published records are from outside

is line elongated appendage an adaptation from Mediterranean basin (Azores, Madeira, coast of

Cantabrian bottom-dwelling to interstitial habits. The way this Portugal, mountains in N.W. Spain),

from specialized elongation is achieved is unique for some are brackish waters but others are

marine from certain members of the family Gammaridae (s.l.) or fresh waters. No subspecies had and for this I attach value to been described so far other than the reason, greater (nor species the characters 1 to 3, based on uropodal mor- type-species).

than characters 4 In the The allocation of in the phology, to to 6. part, Pseudoniphargus super-

character latter states are based on the loss of family Niphargoidea, as advocated by Bousfield, is

with the a certain morphological structure (loss of baso- conflicting diagnosis of that superfamily facial spine of the first uropod, loss of the postero- (Bousfield, 1977: 303). In contrast with the so- distal lobe on the seventh pereiopod), and it can called Niphargoidea, Pseudoniphargus has strong

that inner the lower dissimilar easily be accepted such losses occurred more lobes on lip, strongly

4 is without than once. Character (lobe on P7) moreover gnathopods sexual dimorphism, narrow sex-linked (only present in males of certain oöstegites armed with long setae, a basofacial spine

and is rather therefore to the first and lacks a 2nd species ), unlikely express on uropod, exopodite seg- cladistic relations. ment on the 3rd uropod. - 115 BIJDRAGEN TOT DE DIERKUNDE, 50 (1) 1980

for the of the The left is of the based Fig. 7. Two possible phenograms 9 species genus Pseudoniphargus. one one possibilities

In similar to that the state of character 5 evolved on a 6-character analysis. this or phenograms, we have assume apomorphic

of characters 4 and 6 each twice. This is indication that the four times independently, the apomorphic state a strong apo-

evolution. based morphic state of characters 4 to 6 must be considered a parallel The right phenogram is on characters

1 offers to 3 only and a more consistent picture.

= = 6 = characters 6 of table = in character Abbreviations: p plesiomorph; a apomorph; 1 to 1 to II; x no change state.

in fundamental between the The absence of sexual dimorphism gnatho- agreement two genera.

pods 1 and 2 and in antenna 2 of Pseudoniphar- Barnard (1976) and Barnard & Karaman (in

well its "normal" 2nd to reduce the number of gus, as as (not elongated) press) propose family

of and wide antennal families and the peduncle segment antenna 1, groups or considerably, place

in Bousfield's of the in the Melitoidea. sinus, are not fitting diagnosis genus Pseudoniphargus Since,

I have shown Bousfield's of Melitoidea. as above, diagnosis

Also with the Gammaroidea (in Bousfield's this superfamily does not fit in completely with

of exist the characters of the sense) a number differences (telson un- Pseudoniphargus (especially

lower with this divided, uropod 3 not natatory, lip gnathopods of genus are more gammaroid

well-developed inner lobes, only 7 teeth on the than melitoid), I feel not able right now to follow

Karaman's outer plate of maxilla 1, etc...). Barnard & allocation. Like me, Barnard

From Bousfield's Crangonyctoidea, Pseudo- & Karaman question the supposed close relation-

in antennal of and niphargus differs having a larger sinus, ship Pseudoniphargus Allocrangonyx.

weak sexual in Like have said before other the showing a very dimorphism having I at occasions,

strong inner lobes on the lower lip and narrow subdivision of the Gammaridae (s.l.) is still in a

in the absence of in the and sound is oöstegites, calceoli, presence flux, as long as no system developed, stick the of a basofacial spine on uropod 1 I prefer to to good old Gammaridae. I consider Pseudoniphargus to be derived from the

Bousfield combines with the Gammarus of Pseudoniphargus group, having obtained a number

American in characters North genus Allocrangonyx "Family apomorphic as a result of adaptations to

2" of the I fail conditions. group Niphargoidea. to see any hypogean 116 J. H. STOCK - REGRESSION MODEL EVOLUTION

8. TAXONOMY Type species. — Pseudoniphargus africanus

Chevreux, 1901.

8.1. Genus Pseudoniphargus Chevreux, 1901

— Other species. Pseudoniphargus macro-

Diagnosis. — Gammaridae (s.l.). Blind, telsonis n. sp., Ps. adriaticus S. Karaman, 1955, depigmented. Body elongate. Antennal sinus shal- Ps. unisexualis Ps. Ps. n. sp., longispinum n. sp., low. Pedunculi of Al and A2 not elongated. Al Ps. mateusorum n. sp., brevipedunculatus n. sp., > A2. Cudgel-shaped aesthetes present on distal Ps. branchiatus and four Ps. elongatus n. sp., n. sp., A2. flagellum segments of Al ( 6, ?), absent on called and linnamed forms Pseudoniphargus sp. 1 Accessory flagellum short, 2-segmented. Ps. 3 2 (near africanus), Pseudoniphargus sp. Upper lip entire; lower lip wing-shaped, inner (near Ps. brevipedunculatus ), and Pseudoniphar- lobes prominent. Mandible palp 3-segmented, well- 4 gus sp. (near Ps. elongatus). developed. First maxilla: outer lobe with 7 spines,

inner lobe with 2 setae, palp 2-segmented with 8.2. KEY TO THE SPECIES 8 distal elements. Second maxilla: outer lobe wide,

with distal of no rows of 2 groups setae; oblique la) Pedunculus of third uropod $ strongly elongated 5

b) Pedunculus of third short 2 uropod $ .... setae; inner lobe with terminal setae only. Maxil- 2a) Telson with a deep, V-shaped distal notch. Posterior with low liped: inner lobe feebly developed, a margin of basis of P7 with numerous (15-20) setules of distal outer number setae lobe and Ps. only; palp brevipedunculatus n. sp. well developed. Gnathopods without sexual di- (Faial, Azores)

b) Telson with shallow distal notch. Setules on posterior P2 > propodus Pi. Palmar morphism. Propodus of basis of margin P7 less numerous (less than 12) 3 armed with bifid or angle (Pi) setule-tipped (P2) 3a) Propodus of gnathopod 2 globular, about as long as wide.

of spines. No mid-palmar spines. Coxal gills with Basis P7 almost rectangular, posterior margin straight. [Male 3rd uropod not differentiated] . well-demarcated basal stalk; on P2-P6. Coxal Ps. unisexualis n. sp. rather 4th without plates small; plate strong poste- (prov. Guipuzcoa, Spain)

of than wide. rior excavation. Oöstegites linear, with 5-12 setae. b) Propodus gnathopod 2 not globular, longer of Basis P7 oval or trapezoidal 4 little P4. P6 P3 and P4 setose, equal. P5 as long as ia) Palm of 2nd gnathopod very oblique, slightly longer than and P7 Posterodistal corner of especially elongate. the posterior propodal margin. Basis of P5 to P7 strongly

Pedunculus and of P5-P7 produced to overhanging, sometimes tapering, trapezoidal. segments 4 5 A2 with Ps. long setae longispinum n. sp. sexually dimorphic. Pleopods biramous, well devel- (prov. Beira, Portugal)

with two retinacula. First oped; pine-tree-shaped b) Palm of gnathopod 2 very little oblique, shorter than the

propodal Basis of P5 to P7 ovoid. uropod well developed, 2nd much smaller; baso- posterior margin. Pedunculus segments 4 and 5 of A2 with short setae Third with facial spine present or absent. uropod Ps. mateusorum n. sp. and long, 1-segmented exopodite parviramus-type (near Setubal, Portugal) 5a) Pedunculus and of third uropod of the adult endopodite; non-natatory. In the most plesio- exopodite 9 5 and Al elongated. Pereiopods to 7 flagellum very state, the pedunculus is short, and the morphic slender 6

is not in exopodite elongated; more apomorphic b) Third uropod $ not elongated. Pereiopods 5 to 7 and

Al "normal" 7 states, the exopodite, and later also the pedun- flagellum times 6a) Basal stalle of coxal gills 2 to 3 as long as wide. culus, get elongated, first in males but in the most Posterodistal corner of P7 not overhanging. Aesthetes of also in apomorphic species females. Al long. Telson cleft wide, U-shaped . . . .

of Ps. branchiatus n. Telson entire, with distal spines and a pair sp. (S. E. Spain) setules on either side, distally notched. sensory b) of about Basal stalk coxal gills as long as wide. Postero-

distal corner of P7 overhanging. Aesthetes of Al short.

Telson cleft Ps. narrow, V-shaped. elongatus n. sp.

Distribution. — Groundwaters of inter- (Cantabrian mountains, N.W. Spain)

7a) Postcrodistal corner of basis of P5 to P7 ) over- stitia, springs, wells, caves. Northern shores of ( $ Posterior of hanging. corner epimeres 2 and 3 rounded Adriatic, N.W. shores of Mediterranean, Algeria, Ps. adriaticus Karaman, 1955 Azores. Marine to Spain, Portugal, Madeira, (Mediterranean, coastal)

Posterodistal of basis of to limnic. b) corner P5 P7 ( $ ) not over- BIJDRAGEN TOT DE DIERKUNDE, 50 (1) - 1980 117

of 2 and 3 maxilla has wide outer hanging. Posterior corner epimeres pointed The second ( fig. 8j ) a

8 armed with 2 of setae. lobe, groups 8a) Distal telson spines about as long as the telson. Uropod The maxilliped (fig 9a) differs from Che- 3 with basofacial spine . Ps. africanus Chevreux, 1901 of (Bône, Algeria) vreux's figure in having a row spines (not Distal telson shorter than the telson. b) spines Uropod 3 the setae) on the medial margin of outer lobe. basofacial without spine Ps. macrotelsonis n. sp. The first differs from (Kef Djemel, Algeria) gnathopod (fig. 9b) in 2 1 and Chevreux's figure having groups (of

8.3. DESCRIPTIONS OF THE SPECIES elements 4 setae, respectively) of on the posterior margin of the propodus. The palmar angle is Pseudoniphargus africanus Chevreux, 1901. marked the anterior by 5 bicuspidate spines; cusp Figs. 8-11. of the is than the spines larger posterior cusp ( fig. Chevreux, 1901: 211-216, figs. 1-2. 9b). G. Karaman, 1978: 250-256, figs. VI-IX

The second gnathopod (fig. 9c) has 3 palmar Material examined. — Many samples from wells absent in Bône (= 'Annaba), Algeria, including Chevreux's type- angle spines; mid-palmar spine (fig. 9d)

MNHN coll. Am. material (Feb. 1899, many specimens, no. The posterior margin of the propodus bears 4-5 Am. 1198; Nov. 1901, 3 5 5,8 9 9, MNHN coll. no. of groups setae. coll. Am. May 1200; 1910, 35 specimens, MNHN no. 1202;

The third has an coll. no. Am. 15 9e) elongate 1924, many specimens, MNHN 1199; Jan. pereiopod (fig.

34 MNHN coll. Am. 1203; without 1943, specimens, no. coxal plate. Its setation is rather limited. date, 100+ specimens, MNHN coll. no. Am. 1201; without The coxal plate of pereiopod 4 (fig. 10b) has date, 1 S, ZMB coll. no. 23950). but a weakly concave posterior margin no strong

Descriptive notes, supplementing Che- incurvation. Otherwise P4 resembles P3-

vreux's description. — Maximum length of male The fifth pereiopod (fig. 10c) is short and

and third 7 of (without antennae uropod) mm, heavy. The basis has a strongly developed poste-

female 5 to 6 mm. The lateral head lobes (fig. rior lobe; the posterior margin of the basis bears

sinus is rather 8a) are rounded, the antennal deep. some 7 distinct notches with a setule. The distal

Preserved show First There is specimens no eye pigment. segments bear few setae. hardly any antenna (fig. 8b) correctly described by Chevreux. sexual dimorphism in the shape of the basis

Accessory flagellum shorter than first flagellum (fig. 11a).

segment (fig. 8c). Aesthetes (fig. 8d) present The sixth pereiopod is shorter than the seventh.

the distal of the female is that on segments (from segment 7 onward); The basis a trifle wider than

the terminal aesthete is about % of the length of of the male (figs, lib, c).

and has distinct the segment; the penultimate antepenultimate The seventh pereiopod (fig. lOd) a aesthetes of the of the the this is are 40-50% length cor- posterior lobe on basis; lobe slightly responding segment (fig. 8e). produced in distal direction ( S ) or not produced

The second antenna (fig. 8f) is not very slen- (5, fig. lid). The distinctly notched posterior

the is setules. der; gland cone low, rounded; the flagellum margin of the basis bears some 10 The ter-

is 6-segmented. minal claw (fig. 10e) is rather slender.

The is of the normal armed with upper lip (fig. 8g) gam- Oöstegites (fig. 10a) linear, some marid The lower has well- P2 type. lip (fig. 8k) 10 long setae. Coxal gills on to P6, oval, with developed inner lobes. short basal stalk (figs. 9c, e).

The mandible palp (fig. 8h) is of the gammarid Epimeres not sexually dimorphic; posteroinfe-

is rior type. Chevreux's figure (2A) erroneous. Seg- corner produced into a small tooth (fig. lOf), ment 3 has 1 A-seta, 1 B-seta, no C-setae, a row posterior margin just above this tooth concave;

and of some 9 D-setae of decreasing length, 3 lower margin with 1 spine; posterior margin with

E-setae. 1 2 each setule. or vague notches, carrying a

The first maxilla (fig. 8i) has 7 strong spines Pleopods with 2 pine-tree-shaped retinacula on the outer lobe, 2 setae on the inner lobe, and (fig. lie).

the with 4 spines + 4 setae on palp. Uropod 1 small basofacial spine (figs. 10g, 118 J. H. STOCK - REGRESSION MODEL EVOLUTION

Fig. 8. Pseudoniphargus africanus Chevreux, 1901 (syntypes from Bône, Algeria): a, head of female, from the left (scale

b, basal of first of first aesthete of first A); portion antenna, ￿ (A); c, accessory flagellum antenna, ￿ (B); d, antenna,

distal of first second ￿ (C); e, flagellum segments antenna, ￿ (B); f, antenna, ￿ (A); g, upper lip, ￿ (B); h, mandible palp, ￿ (B); i, first maxilla, ￿ (C); j, second maxilla, ￿ (C); k, lower lip, ￿ (B). - BIJDRAGEN TOT DE DIERKUNDE, 50 (1) 1980 119

B); distal Fig. 9. Pseudoniphargus africanus Chevreux, 1901 (syntypes from Bône, Algeria): a, maxilliped, ￿ (scale b, second d, margin of second third part of first gnathopod, ￿ (B); c, gnathopod, ￿ (A); palmar gnathopod, ￿ (B); e,

pereiopod, ￿ (A). 120 J. H. STOCK - REGRESSION MODEL EVOLUTION

Fig. 10. Pseudoniphargus africanus Chevreux, 1901 (syntypes from Bône, Algeria): a, oöstegite of second gnathopod, ￿ b, fourth claw of (scale A); pereiopod, ￿ (A); c, fifth pereiopod, ￿ (A); d, seventh pereiopod, ￿ (A); e, seventh f, from the left first h, second pereiopod, ￿ (D); epimeres I-III, ￿, (A); g, uropod, ￿ (A); uropod, ￿ (A). BIJDRAGEN TOT DE DIERKUNDE, 50 (l) - 1980 121

Fig. 11. Pseudoniphargus africanus Chevreux, 1901 (syntypes from Bône, Algeria): a, basis of fifth pereiopod, ￿ (scale

of sixth of sixth basis of seventh A); b, basis pereiopod, ￿ (A); c, basis pereiopod, ￿ (A); d, pereiopod, ￿ (A); e, retinacula of first third third third pleopod, ￿ (E); f, uropod, ￿ (A); g, uropod, ￿ (A); h, uropod, ￿ (A); i, telson, ￿ (B); j, telson, ￿ (B). 122 J. H. STOCK - REGRESSION MODEL EVOLUTION

macrotelsonis 1 if). Contrary to Chevreux's figure, the dorso- Pseudoniphargus n. Sp. Fig. 12.

distal spine on the pedunculus is long (almost

Material examined. — One $ (holotype), thirty- half as long as the ramus). two specimens of both sexes (paratypes). Algeria: subter- and Uropod 2 (fig. 10h) with short pedunculus Kef 4 ranean spring at Djemel near Medjez-Sfâ ) in the Beni

short rami. Salah mountains (alt. about 600 m) (MNHN, cat. no.

Am. 1197). Uropod 3 with strong sexual dimorphism. In

the adult male (fig. Hg) the pedunculus is elon-

— male the is times Description. Largest 5y mm long gated; uniarticulated outer ramus 2 the female the (excluding uropods), largest 5 mm. as long as pedunculus, slightly upcurved (Ni- Closely related to Ps. africanus, but with a num- phargus-like •); the inner ramus is a rounded lobe, ber of apomorphic characters. The features not armed with 1 setule. The length of both pedun-

mentioned resemble to those of Ps. and with in africanus. culus exopodite increase age males; The distalmost aesthete of the flagellum of the so, their ratio stays more or less the same. In the about distal the third elon- first antenna is as long as the segment female, uropod (fig. llh) is never (fig. 12a). gated; it resembles the type found in certain The first uropod (fig. 12d) has a short distal Chaetogammarus species. The armature of the pedunculus spine; the basofacial pedunculus spine pedunculus and the 1-segmented exopodite con- is lacking. sists of spines only.

The 1 to 3 (figs. 12c, d) have a The telson (figs. Hi, j) is as long as wide and epimeres very the tooth has 4 each of strongly produced posterior corner; on a varying number (2 to on side) the the is distal the of these attain posterior margin of plates more strongly spines; longest spines may developed than in Ps. africanus. Between the the same length as the telson itself. The median and the tooth the notch is its reaches pointed corner on posterior mar- V-shaped; deepest point to, or

gin, a rounded sinus is dépassés, the implantation of the spines. There are developed. The telson is than 2 lateral short (fig. 12e) slightly longer sensory setules, implanted at a

wide and bears 2 to 4 distal on either side; distance of the distal spines. spines these shorter in Ps. The spines are than africanus.

mediodistal notch is shallower, more rounded, and

less than in Ps.

Remarks. — V-shaped africanus. The above redescription is based

Chevreux's The is on syntypes. species only known

with certainty from wells in the surroundings of

Remarks. — This sample was already studied Bone (= 'Annaba), Algeria. All other North by Chevreux (1901: 216), who considered it "une African samples I have examined, belong to variété locale". The differences with Ps. morphologically distinct forms. I hesitate, there- africanus in several hundreds are admittedly slight, but of fore, to accept G. Karaman's (1978: 254) iden- examined this 3 specimens of Ps. africanus during tification from Oued-ed-Demane ), Tunisia, alt.

none shows intermediate characters. It 450 m. study, in the depends on material collected future, hope-

fully from localities intermediate between Bône

:i ) It is not sure which is meant this name. I the locality by (on sea coast of Algeria) and Kef Djemel (in been unable have to trace a locality with exactly the same the mountains), whether Ps. macrotelsonis can be spelling, however both Wadi ad Dmaïne (in the El Kef be maintained as an independent species, or must region, 36°08'N 09°07'E) and an unnamed oued (— dry Ed 37°03'N river) near (or Ad) Dmaïne (Tamera region, considered a subspecies.

09°00'E) come near enough in spelling to be considered as possible candidates. Neither of these two localities can be

in but 1 4 This is recorded in the Times but found a regular atlas, they are shown by the : 100,000 ) locality not Atlas, can

181. topographic map of Tunisia. In view of the uncertainty as be found in Andrees Handatlas, ed. 8, sheet It is

to the exact position of the locality, it has not been introduced situated S.E. of Duvivier in the chain of the Monts de

on my maps (figs. 1-6). Medjerda. BIJDRAGEN TOT DE DIERKUNDE, 50 (1) - 1980 123

from Kef distal of first antenna, Fig. 12. Pseudoniphargus macrotehonis n. sp. (paratypes Djemel, Algeria): a, segments

￿ (scale B); b, epimeres I-III, ￿, from the right (F); c, epimeres I-III, ￿, from the left (F); d, first uropod, ￿ (A):

of different e, telson two males (B).

2. Pseudoniphargus sp. 1. Fig. 13. Pseudoniphargus sp.

Material examined. — Two 9 9 (of which one Material examined. — One $. Algeria, from a ovigerous). Algeria: Rhar el Khal, wilaya Tlemcen, daira well in the Mustapha suburb of Alger, leg. H. Gauthier Khemis altitude 1150 Sebdou, (34°34'28"N01°34'17"W); m; coll. (ZMB no. 23951).

14/15 Aug. 1978. Cave, in a small stream connecting the

Galerie de la Rivière with the underground brook; depth Remarks. — This is a mm large (length 5y 2 ), C. 0.2-1.0 m; stony; temp. 15° Biospeleo Werkgroep, sta. no doubt adult, female specimen. It is not identical A 8-VIII (ZMA coll. no. Amph. 107.383). with Ps. because the telson has africanus, a very

-—■ distal notch and the third Remarks. In absence of males, this mate- shallow uropods are females rial is hard to classify. The two specimens differ elongated (in of africanus they are not

The sufficiently from Ps. africanus to prevent inclusion elongated). epimeres on the other hand

in that species. The differences pertain to the resemble those of Ps. africanus. In absence of

Al this has greater length of the aesthetes on (fig. 13a), males, specimen not been named.

the basis in P7 the more elongate shape of P5 to

(figs. 13b-d), the slender claw on the very poste- Pseudoniphargus adriaticus S. Karaman, 1955. rior 13e), the shape of the pereiopods (fig. epi- Figs. 14-15.

meres (posterior margin convex instead of con- Ps. africanus Chevreux forma adriatica S. Karaman, and the of the telson 1955: cave, fig. 13f), armature 236-241, figs. 44-51. reduced to either side a single spine on (fig. 13g). Ps. adriaticus; G. Karaman, 1978: 241-250, figs. I-V.

The locality (cave) where the sample was taken Material examined. — Two $ $, four 9 9 . France, is described in Guldemond et al., 1979: 70. dép. Var: Le Petit Gaou near Le Brüse, steep beach (slope 124 J. H. STOCK - REGRESSION MODEL EVOLUTION

from Rhar el distal of first basis Fig. 13. Pseudoniphargus sp. 1 (￿ Khal, Algeria): a, segments antenna (scale B); b,

of fifth pereiopod (A); c, basis of sixth pereiopod (A); d, basis of seventh pereiopod (A); e, distal segments of seventh

and telson pereiopod (A); f, epimeral plates II III (A); g, (B).

tide 25°) of coarse sand and fine gravel, just below low line Descriptive note. — A small interstitial (behind Studios meublés Le Grand Sarge); 20 Feb. 1979 male female species (adult 4-5 mm, ovigerous Stock. (ZMA coll. no. Amph. 107.336). Leg. J. H. Accom- 3-4 without Number of small panying fauna: Tanaidacea, Idotea, Melita, flatworms, Har- mm, uropods ). eggs pacticoidea, Polychaeta. (up to 4). Eleven specimens. Same locality; 15 Apr. 1979; leg. First antenna (fig. 14a) with long aesthetes J. H. Stock (ZMA coll. no. Amph. 107.337). (Collected with than the distal almost of biophreatic pump type Bou-Rouch.) (longer flagellum segment, Fifteen the and specimens. Same locality; 29 Dec. 1979; leg. as long as penultimate antepenultimate

J. H. Stock (ZMA coll. no. Amph. 107.382). segments). Three specimens. France, dép. Var; Calangue du Port First from that d'Alon, in fine gravel, intertidal; marine; 11 Jan. 1980 gnathopod (fig. 14b) differing

(ZMA coll. no. Amph. 107.381). in Ps. africanus by the shape of the palmar angle Twenty-nine specimens. France, dép. Bouches-du-Rhône: spines (fig. 14c). These are bicuspidate, but the du Cassis, Plage Bestouan; steep (slope 20°), coarse gravel

anterior is shorter than the one. beach, near low tide line; 16 Apr. 1979; leg. J. H. Stock cusp posterior

coll. fauna: (ZMA no. Amph. 107.334). Accompanying The second gnathopod (fig. l4d) bears 2 or Caecum (Gastropoda), Neogammarus, Annelida. 3 of the of the groups setae on posterior margin Thirty specimens. France, dép. Pyrénées-Orientales: Le in Ps. The Troc (Banyuls); 2 May 1950; leg. E. Angelier no. 201 propodus (4-5 groups africanus). pal-

without cat. no.). (ZMB mar margin is slightly sinuous. The shape of the One $. Same locality; 25 May 1950; leg. E. Angelier propodus is slightly more elongate (less globular (MNHN coll. no. Am. 618). than in Ps. Three $ $, three 9 9, one juv. France, Corse: Bonifacio, africanus). Grotte St. Barthélémy; 14 Sept. 1948; leg. P. A. Rémy Pereiopods 5, 6 and 7 of the male (figs 15a, d, (ZMB coll. no. 25469). f have a more distinctly produced posterodistal Three $ $, twenty-five 9 9. Yugoslavia: Split, Kloster- e, )

Pax quelle; 1939; leg. F. (ZMB coll. no. 25046). lobe on the basis and the posterior margin of the BIJDRAGEN TOT DE DIERKUNDE, 50 (1) - 1980 125

Fig. 14. Pseudoniphargus adriaticus Karaman, 1955 (a, c, e, g, h, i, k, m from Le Petit Gaou, France; b, d, f, j from Split,

Le of first distal Yugoslavia; 1 from Troc, Banyuls, France): a, distal segments antenna, ￿ (scale B); b, part of first gnathopod, ￿ (B); c, palmar angle spine of first gnathopod, ￿ (ant. = anterior margin) (E); d, distal part of second

of first gnathopod,￿ (B); e, coxal gills of second gnathopod, ￿ (B); f, oöstegite fourth pereiopod, ￿ (B); g, uropod,

￿ (B); h, third uropod, ￿ (B); i, third uropod, ￿ (A); j, telson, ￿ (B); k, telson, ￿ (C); 1, telson, ￿ (B); m, telson, ￿ (B). 126 j. H. STOCK - REGRESSION MODEL EVOLUTION

Fig. 15. Pseudoniphargus adriaticus Karaman, 1955 (e from Le Troc, Banyuls, France; remaining figures from Le Petit Gaou,

basal of fifth of fifth of sixth France): a, part pereiopod, ￿ (scale D); b, same pereiopod, ￿ (D); c, same pereiopod,

9 (D); d, same of sixth pereiopod, ￿ (D); e, f, same of seventh pereiopod, ￿ (D); g, same of seventh pereiopod, ￿ (D);

of fifth of from h, distal segments pereiopod, ￿ (B); i, same seventh pereiopod, ￿ (B); j, epimeres I-III, the left, ￿ (D). BIJDRAGEN TOT DE DIERKUNDE, 50 (1) - 1980 127

fewer setules than in — Known from basis bears Ps. africanus. Distribution. a large

In the female, the posterodistal lobe of P5 to P7 stretch of coast in the northern Mediterranean,

is not produced (figs. 15b, c, g). from Banyuls (France) toBudva (Yugoslavia) and

The epimeral plates have a rounded posterior from the Balearic Islands and Corsica. Presumably,

corner (fig. 15j). G. Karaman's record (1978: 248) from Funchal The first uropod lacks a basofacial spine (fig. (Madeira) supports on confusion with some other I4g). species.

The third male uropod has a modified (elon-

and In unisexualis gated) pedunculus exopodite (fig. l4i). Pseudoniphargus n. sp. Figs. 16-17. the female, the third uropod is unmodified (fig.

I4h). Material examined. — One 3 (holotype), one 5

(allotype), sixteen paratypes. Spain, Guipuzcoa, field- The telson is slightly wider than long (figs. 14 prov. no. 79-36: Cueva de San Adrian de Cegama (near the south- It is 1 and 1 short j-m). distally armed with long border of ern the province); completely dark; in second

on either side. The setules im- terminal) left of mud, spine sensory are (= basin, the chapel; wood debris,

altitude 1000 1 In stones; (estimated) Aug. 1979. a planted close to the spines. The medial notch is m; straight line, this km from locality is 40 the sea (ZMA coll. shallow: its does reach very deepest point not the no. Amph. 107.341-343). Accompanying fauna: juveniles of line that the connects implantation of the left and Echinogammarus sp., Oligochaeta, Cyclopidae, Harpacticoidea.

Same locality, ("in kleiner Restlache", in the remains right telson spines. Remaining characters similar ganz of small a very pool); 14 Oct. 1935; leg. H. J. Stammer; one to those of Ps. africanus. 2 (ZMB coll. no. 24817).

Same locality ("Grössere Tropfwassertümpel", in larger dripwater pool), 14 Oct. 1935; leg. H. J. Stammer; six Remarks. — There constant dif- are so many specimens (ZMB coll. no. 24816), and two specimens (ZMB ferences between Ps. and the typical africanus coll. no. 24815). form present that I have, in agreement with G.

Karaman doubt it no a distinct tion. — (1978) represents Descrip Length: male 7 mm, female

species. from remote localities, like 6.5 Specimens (ovigerous) mm. Eggs 0.54 x 0.64 mm. Corsica Split, or Banyuls, are morphologically very Secondary sexual differences almost absent. similar. First antenna (fig. 16a) with 16-segmented

After completion of revision, I my present flagellum; accessory flagellum slightly shorter than received G. Karaman's of Ps. first redescription afri- flagellum segment. Aesthetes on all but the

canus and Ps. adriaticus the is called basal three ( 1978; paper segments of the flagellum. Aesthete on for indistinct reasons "Revision", no although penultimate segment longer than the segment it- materials original are re-examined One of the on the at least ). self; other segments % of the length

features discovered by G. Karaman to of segregate the segments. adriaticus from is africanus the of small Second antenna with presence (fig. 16b) 9-segmented cuticular denticulations on the posterior margin of flagellum. the second metasomite in males of the former. Mandible with robust 2nd palp (fig. 16c) seg- Since the samples on which I have based the ment and very robust (almost trapezoidal) 3rd

revision had been returned to their the latter present already segment; armed with 14 D-setae, 2 A-

I have check respective depositaries, been unable to setae, 2 B-setae and 3 E-setae. the validity of this feature for the various taxa First gnathopod (fig. l6d); propodus with 3 recorded here. of groups setae on posterior margin; palmar angle with 6 trifid spines (fig. I6e); spines on palmar

Ecology. — This is a from species marine margin also trifid.

coarse sands and gravels, from just below to Coxal just gills with short basal stalk (fig. I6f). above the tide lines Mediterranean (the tides are Oöstegites linear (fig. l6i).

weak) or from anchihaline localities like the very Second gnathopod: coxal plate longer than wide of sea caves Bonifacio (Corsica) and the "Cloister I6f (fig. ) ; propodus about as long as wide, globu- Spring" near Split, Yugoslavia. lar (fig. 16g); three palmar angle spines, of which 128 J. H. STOCK - REGRESSION MODEL EVOLUTION

16. unisexualis from San Adrian de first Fig. Pseudoniphargus n. sp. (paratypes Cegama, Spain): a, antenna, ￿ (scale F); b, second antenna, ￿ (F); c, mandible palp, ￿ (B); d, distal part of first gnathopod, ￿ (A); e, palmar angle spines of same

coxal and coxal of second distal of second (B); f, plate gill gnathopod, ￿ (G); g, part gnathopod, ￿ (A); h, fourth pereio- pod, ￿ (G); i, oöstegite of fourth pereiopod, ￿ (A). 50 - 1980 129 BIJDRAGEN TOT DE DIERKUNDE, (l)

from San Adrian de fifth Fig. 17. Pseudoniphargus unisexualis n. sp. (paratypes Cegama, Spain): a, pereiopod, ￿ (scale G);

b, distal end of basis of fifth pereiopod, ￿ (A); c, basis of sixth pereiopod, ￿ (F); d, seventh pereiopod, ￿ (G); e, of distal end of basis of seventh pereiopod, ￿ (A); f, same of seventh pereiopod, ￿ (A); g, dactylus seventh pereiopod,

￿ (A); h, first uropod, ￿ (F); i, second uropod, ￿ (F); j, third uropod, ￿ (A); k, third uropod, ￿ (F); 1, epimeres

I-III, from the left, ￿ (F) ; m, telson, ￿ (A); n, telson, ￿ (A). 130 J. H. STOCK - REGRESSION MODEL EVOLUTION

in sexual in one very long; spines on palmar margin trifid; present one having strong dimorphism

between the of and palmar margin slightly S-shaped (concave exopodite uropod 3 (elongated up-

the palmar angle spines and mid-palmar setae, curved in the male). Moreover, Ps. brevipeduncu-

convex between the mid-palmar setae and the im- latus has a more deeply cleft telson than Ps. uni-

plantation of the claw). sexualis, and the propodus of gnathopod 2 is much

Third and fourth pereiopods (fig. l6h) without longer than wide. Ps. mateusorum differs from

particulars. Ps. unisexualis in the overhanging posterior lobe

Fifth pereiopod (fig. 17a): basis (fig. 17b) of the basis of pereiopod 7 ( <3 ), in the more

setules elongate, rectangular; posterior margin with 8 numerous on the posterior margin of the

setules; merus rather slender. same segment, and in the rounded epimeral plates.

Sixth basis to now the is known from the pereiopod (fig. 17c) : very elongate, Up species only

slightly tapering; posterodistal corner only slightly type-locality, the San Adrian cave 5 ), on the produced. southern slopes of the Cantabrian mountain chain.

Seventh pereiopod (fig. 17d) slender; basis The other Pseudoniphargus from the Cantabrian

Ps. in of very elongate, slightly tapering, posterior margin mountains, elongatus, occurs a number

with 7 setules localities the northern only, posterodistal corner only on slopes.

slightly produced (figs. 17e, f); claw slender Margalef (1970) records a Pseudoniphargus

from (% 17g). a cave not far from San Adrian, viz. Aya-

Ataun 6 First uropod (fig. 17h) similar to that of Ps. ). I have been unable to re-examine this

africanus. sample, but Margalef's figures (especially of the

Second armed with seem to indicate that this material uropod (fig. 17i) very P2) belongs

elongate spines. to the present species.

Third uropod (figs. 17j, k): basal segment ( <5, 2) hardly longer than wide; endopodite trian- Pseudoniphargus longispinum 18-19. n. sp. Figs. with gular. Exopodite 3-4 (<3) or 2 (?) lateral

and of 3 <3 or medial groups 2 Material examined. — spines, ( ) (?) One $ (holotype), one $ of Several (paratype). Portugal, Beira Literal: de groups spines. (4-6) distal spines. prov. Ponte Mucela ( D H. of Coimbra), in a well; 14 Telson June 1958 (IZAN). (figs. 17m, n) almost ter- rectangular; Three 2 9 (fragmentary). Portugal, prov. Beira Literal: minal excavation left and very slight; right lobe Anadia (= N.W. of Coimbra); 17 March 1968 (IZAN).

each with 2 or 3 of very long spines; a pair sensory

setae implanted near the middle of the lateral Description. — Body length 4-4.5 mm. First

margin. antenna (fig. 18a) with fairly slender pedunculus;

about Epimeral plates (fig. 171): posterior corners accessory flagellum as long as the first fla-

rectangular, produced into a minute tooth; ventral gellum segment. Flagellum 16-segmented. Distal- aesthete margin unarmed. most slightly longer than distal flagellum

segment, penultimate and more proximal aesthetes about Remarks. — There is sexual 2/3 of the of the no secondary length corresponding seg-

in the and ment (fig. 18b). dimorphism gnathopods pereiopods.

The Second with exopodite of uropod 3 in the female is slightly antenna (fig. 18c) long setae on less slender 4 and and slightly less spinose than in the peduncle segment 5. Mandible palp as in but the Ps. male, pedunculus is not sexually di- brevipedunculatus. First morphous. The absence of noticeable sexual di- gnathopod: shape of palmar angle spines

has the resembling that in Ps. adriaticus. morphism inspired specific name, uni- sexualis, a name reminiscent of the unisex dresses 5 This is shown ) locality not by usual atlas. It be so in of any can vogue a couple years ago. found the scale on Mapa Turistico, 1 : 200,000 (ed. Fire- The two other which have a short stone Hispania), sheet species T-22, on the S. slope of the Monte in the third Aitzgorri. pedunculus male uropod (Ps. mateus- «) Firestone Hispania, sheet situated and Ps. T-22; in the Sierra de orum differ from the brevipedunculatus), S.S.W. of Aralar, Beasain, some 13 km E.N.H. of San Adrian. BIJDRAGEN TOT DE DIERKUNDE, 50 (l) - 1980 131

18. from Ponte de from basal Fig. Pseudoniphargus longispinum n. sp. ( ￿ Mucela, Portugal; ￿ Anadia, Portugal): a, portion of first ￿ distal of of first ￿ second distal antenna, (scale A); b, segments flagellum antenna, (B); c, antenna, ￿ (A); d, part of second gnathopod, ￿ (B); e, second coxal plate, ￿ (ant. = anterior margin) (A); f, coxal gill of fourth pereio-

￿ fifth basis of sixth ￿ of fifth ￿ pod, (A); g, pereiopod, ￿ (F); h, pereiopod, (F); i, dactylus pereiopod, (D); j, seventh pereiopod, ￿ (F); k, distal margin of basis of seventh pereiopod, ￿ (D); 1, epimeral plates I-III, ￿ (D). 132 J. H. STOCK - REGRESSION MODEL EVOLUTION

Coxal plates 1-4 elongate (fig. 18e). Coxal Second uropod likewise armed with long spines gills with short basal stalk (fig. 18f). Oöstegites (fig. 19b).

in linear, non-setose (= non- reproductive stage) Third uropod with slight sexual dimorphism:

specimens examined. exopodite slightly more slender in the male (fig.

Propodus of 2nd gnathopod trapezoidal with 19d) than in the female (fig. 19c). Exopodite

of armed with Pedunculus very oblique palma; length palmar margin spines only. not ($) or longer than that of the posterior propodal margin; hardly ( <3 ) elongate.

with with posterior margin 3 groups of long setae Telson (fig. 19e) broad, not very deep,

(fig. 18d). apical notch. Terminal spines (2 on either side)

Third and fourth pereiopods without particular unusually long. Sensory setules placed at some characters. Basis of 5th to 7th pereiopods (figs. distance of the insertion of the spines.

18g, h, j) tapering, much wider proximally than

armed with 8 10 Remarks. — This form the distally; posterior margin to belongs to group widely spaced setules. Merus short. Claw slender of species in which the pedunculus of the 3rd male

(fig. 18i). Distal propodal setae of normal length. uropod is not strongly elongated. The exopodite

The posterodistal corner of the basis of P5 to of uropod 3 in the male is not strongly elongated

is in the male than which the form P7 more strongly overhanging either, distinguishes present at

in the female (cf. fig. 18k). once from Ps. mateusorum, the only other species

The epimeral plates II and III have rectangular from continental Portugal. The tapering shape of armed with small tooth the basal of the posterior corners a pointed segment pereiopods 5 to 7, very

with in the of the and a setule. Inferior margin 1 or 2 strong oblique palma gnathopod 2, setosity

spines (fig. 181). second antenna, the spinosity of the first uropod,

First with Basofacial form distinct additional uropod very strong spines. etc. differences.

curved. Rami slender The related be Ps. pedunculus spine strong, most closely species may (fig. 19a). unisexualis from northern Spain, which resembles

19. from Ponte de ￿ from first Fig. Pseudoniphargus longispinum n. sp. ( ￿ Mucela, Portugal, Anadia, Portugal) : a, uropod,

￿ second ￿ third ￿ third ￿ (scale D); b, uropod, (D); c, uropod, (D); d, uropod, ￿ (A); e, telson, (D). BIJDRAGEN TOT DE DIERKUNDE, 50 (l) - 1980 133

than the present taxon in the morphology of uropod 3, cuius spine, distal peduncular spine shorter

in other telson half the of the slender rami. but differs many ways (depth length notch, less strongly spinose first uropods, absence Third uropod in the male (fig. 20k) with short

of about the telson + its distal spines on lower margin of epimeral plates, pedunculus ( as long as

of in in shape the basis P5 through P7, more globular spines). Third uropod the female (fig. 201) distal propodus in P2, less elongate coxal plates 1-4 ... ). with still shorter pedunculus; segment

The name, al- proposed specific longispinum , slightly elongated, tapering.

ludes to the armature of the telson, and first and Telson (fig. 20m) wider than long; distally

armed with and shorter second uropods. 1 very long 1 spine on

either setules side; sensory plumose, implanted

the insertion of the distal Medial Pseudoniphargus mateusorum n. sp. Fig. 20. near spines. notch

wide and very shallow.

Material examined. — One $ (holotype), forty-

da seven 9 9 and juveniles (paratypes). Portugal: Costa

— The the Remarks. new species is, by struc- Arrâbida (Bight of Setubal), Lapa dos Morcegos (= bat of the third male and other charac- cave); 26 June 1941; leg. A. Barros Machado (ZMB coll. ture uropod by

no. 25248). ters, closely related to Ps. brevipedunculatus from

One $, same locality; 22 February 1973; leg. A. Barros, the Azores. For the differences between the two, A. Mateus & E. Mateus (IZAN). see the remarks under the latter species.

— Male Description. 4l/2 mm; ovigerous

female female with — This is named in (5 eggs) 41/2 mm; oöstegites Etymology. species

5 mm. honour of Dr. Amilcar Mateus and Emilia de Oli-

First and veira Mateus in of their various antenna: pedunculus accessory flagel- recognition con-

in lum (fig. 20a) as Ps. africanus. Flagellum 15- tributions to the knowledge of the Portuguese

aesthetes distal segmented; on segment 3 to 14; amphipod fauna.

aesthetes (fig. 20b) as long as or slightly longer

than the corresponding segment; proximal Pseudoniphargus brevipedunculatus n. sp. aesthetes half the about as long as segment (fig. Figs. 21-22. 20a).

Second antenna slender, (fig. 20c): pedunculus Material examined. — One $ (holotype), one 9

Azores: little setose. (allotype), fifty-six paratypes. Faial, Horta, coast,

no. 2/3 Apr. Exped. P. Brinck & E. Dahl, it has poço 2; 1957; Mandible palp about as in Ps. adriaticus; loc. 83 (ZIL cat. no. 1). 4 E-setae.

First in adriaticus. — Adult male gnathopod as Ps. Description. 6l/2 -7 mm,

Second of about the female mm gnathopod: propodus same 6l/ 2 -

in Ps. of The first has slender shape as brevipedunculatus; 3 groups antenna pedunculus seg-

setae on its posterior margin. Oöstegites as in ments (fig. 21a); the flagellum is 16-segmented;

Ps. unisexualis. the is 2-segmented accessory flagellum (fig. 21c)

and fourth in Ps. adriaticus. slender and about the first Third pereiopods as as long as flagellum

Fifth The aesthetes (fig. 20d), sixth (fig. 20e) and seventh segment. on the penultimate and

in with than pereiopods (figs. 20f, g) the male a antepenultimate segments are slightly longer

strongly projecting posterodistal lobe. Basis of P6 the corresponding flagellar segment (fig. 21b);

the and P7 of elongate shape; basis of P5 with some 8 the other aesthetes are 3/4 to 4/5 of length of

with of setules on posterior margin, of P6 9, P7 the corresponding segment.

with inner the The second is rather slender 11. No spines on basis. Dactylus antenna (fig. 21 d)

of P7 slender of of and very (fig. 20h). Shape basis setose.

The has P5-P7 in the female as in Ps. adriaticus. mandible palp 7 setae on segment 2

Epimeres (fig. 20i) rounded. and bears 2 A-setae on segment 3; otherwise it is

First uropod (fig. 20j) with basofacial pedun- as in Ps. africanus. 134 J. H. STOCK - REGRESSION MODEL EVOLUTION

and from dos Fig. 20. Pseudoniphargus mateusorum n. sp. ( ￿, holotype ￿, paratype, Lapa Morcegos, Portugal) : a, basal

of first distal of of first second portion antenna, ￿ (scale A); b, segments flagellum antenna, ￿ (B); c, antenna, ￿ (A);

basis of fifth of sixth seventh d, pereiopod, ￿ (F); e, basis pereiopod, ￿ (F); f, pereiopod, ￿ (F); g, posterodistal corner

of basis of seventh seventh II and from the pereiopod, ￿ (D) ; h, dactylus of pereiopod, ￿ (D) ; i, epimeral plates III, right,

first third ￿ ￿ (A); j, uropod, ￿ (A); k, uropod, ￿ (A); 1, third uropod, (A); m, telson, ￿ (B). BIJDRAGEN TOT DE DIERKUNDE, 50 (l) - 1980 135

from basal of first Fig. 21. Pseudoniphargus brevipedunculatus n. sp. (paratypes Horta, Faial, Azores): a, portion antenna,

distal of first second ￿ (scale A); b, segments of flagellum antenna, ￿ (B); c, accessory flagellum, ￿ (B); d, antenna, ￿

of first = anterior distal of second (A); e, palmar angle spine of palma gnathopod, ￿ (H) (ant. margin); f, part gnathopod,

fourth coxal from the first (A); third ￿ (A); g, plate, ￿ (A); h, epimeral plates I-III, left, ￿ (A); i, uropod, ￿ j, uropod, ￿ (A) ; k, third uropod, ￿ (D). 136 J. H. STOCK - REGRESSION MODEL EVOLUTION

The basofacial palmar angle spines of gnathopod 1 (fig. Uropod 1 (fig. 2 li) possesses a

have of the distal is less 2 le) 2 cusps about equal length. peduncle spine; peduncle spine

rami and The second gnathopod has an elongate, trape- than half as long as the rami; pedunculus

zoidal 2If the slender. propodus (fig. ) ; posterior propodal

bears 4 male margin groups of setae; the palmar margin Uropod 3 (fig. 21 j ) : pedunculus short

is regularly curved. (less than twice as long as wide); endopodite

The coxal of 3 and 4 plates pereiopods (fig. as usual; exopodite very elongated, upcurved,

21g) bear a few more setules than in Ps. africanus. armed with long, slender spines; exopodite 6 times

The coxal gills are as in Ps. africanus. as long as the pedunculus.

The of the has short basis 5th to 7th pereiopod a Uropod 3 female (fig. 21k): pedunculus

produced posterodistal corner in the male (figs. (11/2 times as long as wide); exopodite slightly

22a, d). In the female, this corner is only feebly elongate, tapering, 31/2 "3% times as long as the produced (fig. 22g). The posterior margin of the pedunculus, armed with long slender spines.

basis is densely set with setules (up to 20 setules Telson (fig. 22h) wider than long, armed with

and distal have been counted). Moreover, the basis of P7 1 shorter 1 very long spine on either

bears inner subdistal the two spinules. side; sensory setae implanted very closely to

The epimeral plate 1 is rounded; plates 2 and 3 insertion of the spines. Distal notch V-shaped,

have rectangular posterior corners; their lower mar- rather deep; cleft narrower than in Ps. africanus.

gin bears 3 spines (fig. 21h).

22. from Fig. Pseudoniphargus brevipedunculatus n. sp. (paratypes Horta, Faial, Azores): a, fifth pereiopod, ￿ (scale F);

of fifth basis of fifth b, dactylus pereiopod, ￿ (D); c, distal end of pereiopod, ￿ (D); d, seventh pereiopod, ￿ (F); e,

of seventh distal end of seventh of dactylus pereiopod, ￿ (D); f, of basis pereiopod, ￿ (D); g, same ￿ (D); h, telson, ￿ (D). BIJDRAGEN TOT DE DIERKUNDE, 50 (1) - 1980 137

— males to 6 mm Remarks. — The only other species in which Description. Large up

the of in the male is short, adult males from mm in the pedunculus uropod 3 long, 4-41/ 2 present whereas the exopodite is strongly elongated, is samples; ovigerous females from 3 to 5 mm long.

from The of small Ps. mateusorum continental Portugal. Number eggs (2 to 4).

latter has shallow telson First antenna: species a very notch, more pedunculus (fig. 23a) only

rounded slender claw the more slender than in Ps. Acces- epimeres, a very on slightly africanus.

less slender Al the first posterior pereiopods, a pedunculus, sory flagellum as long as or longer than,

of differences and a less setose A2. The number flagellum segment (fig. 23b). Flagellum 19- to

exceed- is rather small, but clear cut. On morphological 20-segmented. Distal flagellum segments

grounds alone, Ps. brevipedunculatus might be ingly slender (fig. 23c); aesthete on penultimate

of the considered a subspecies of Ps. mateusorum. Since I segment about 80% of length distal seg-

less than half do not adhere to the idea of a landbridge con- ment, on the remaining segments as

and the I the necting Portugal Azores, prefer to con- long as corresponding segment (fig. 23c).

sider both populations as separate species. Second antenna (fig. 23d) slightly more slender than in Ps. africanus.

in Ps. Mandible palp (fig. 23e) as africanus.

First in adriaticus. Pseudoniphargus sp. 3. gnathopod as Ps.

Second gnathopod (figs. 23f, g) : propodus of

examined. — One 9. Madeira: Funchal, in Ps. Material very elongate shape. Coxal gills as africanus

cistern of the Seminary; June 1933; leg. J. de Goureia Bar- (fig. 25a). reto (ZMB coll. no. 25082). Pereiopods 3 and 4 without peculiarities; claw

Remarks. — This, unfortunately single speci- of P4 slender (fig. 23h).

the female is of small size Fifth with men of sex (length pereiopod: basis slightly (male, fig.

3 mm). The general morphology resembles that 23i) or not (female, fig. 24a) projecting postero- of the in distal Posterior of basis with Ps. brevipedunculatus from Azores, but corner. margin some absence of males of I setules. Basis P6 or large females, refrain from 8 (P5) to 12 (P7) of male (fig.

identifying the Madeiran material firmly. 23j) and P7 male (fig. 24c) distinctly projecting;

in P6 female and P7 female projecting (fig. 24b).

The posterior pereiopods and their terminal claws,

in in Pseudoniphargus elongatus n. sp. Figs. 23-25. particular P7, very slender.

Epimeral plates rectangular to somewhat

Material examined. —• One $ (holotype), one rounded (figs. 23k, 1); lower margin with 1 spine. 9 Ramales de ovigerous (allotype). Spain, prov. Santander, 1 (fig. 25h) with basofacial la Victoria, Cueva la Cullevera; in shallow pool; 22 July Uropod spine.

J. H., M. & D. Stock (ZMA coll. 1979; leg. no. Amph. Distal peduncle spines short; rami slender. 107.333). Uropod 2 (fig. 25c) with slender rami. leg. Five specimens, same locality; Aug. 1958; Rousset & with De Loriol coll. Uropod 3 (male) strongly elongate pedun- (MNHN no. Am. 617).

De culus Three specimens, same locality; 7 Aug. 1965; leg. and strongly elongate exopodite (fig. 25d). Loriol (UL coll. no. Et. 36). Uropod 3 (ovigerous female) with moderately Four Cueva la Cas- specimens, Spain, prov. Santander, pedunculus and exo- cada; 6 Aug. 1962; leg. De Loriol (UL coll. no. Et. 19). elongate moderately elongate Three Cueva del podite specimens, Spain, prov. Santander, Molino; (fig. 25f). In smaller, non-ovigerous

6 Aug. 1959; leg. De Loriol (UL coll. no. Et. 18). the females, 3rd uropod has a hardly elongated all 9 Twenty-one specimens, 9, Spain, prov. Santander, and a Cueva de la Station de Santa Isabel; 8 Oct. 1936; leg. pedunculus slightly elongated exopodite

H. J. Stammer (ZMB coll. nos. 24812, 24813, 24814). (fig. 25e). de Castillo One $, one 9, Spain, prov. Santander, Cueva Telson (figs. 24d, e) with 2 to 4, not exceed- near Puente Viesgo; 6 Oct. 1935; leg. H. J. Stammer (ZMB setules coll. ingly long, distal spines; sensory placed at no. 24811). Fifty-three this distance from juveniles, probably species, Spain, prov. some the insertion of the spines; Guipuzcoa, Cuevas de Landarbaso, S.E. of San Sebastian, in distal notch V-shaped, medium deep. pool; 18 Sep. 1935; leg. H. J. Stammer (ZMB coll. nos.

24809, 24808, 24810). 138 J. H. STOCK - REGRESSION MODEL EVOLUTION

from Santa from Ramales de la Fig. 23. Pseudoniphargus elongatus n. sp. (b Isabel, Spain, remaining figures Victoria, Spain):

a, basal portion of first antenna, ￿ (scale A); b, accessory flagellum, ￿ (B); c, distal flagellum segments of first antenna,

second mandible distal of second ￿ (B); d, antenna, ￿ (A); e, palp, ￿ (B); f, part gnathopod, ￿ (A); g, palmar margin

of second gnathopod, ￿ (B); h, distal part of fourth pereiopod, ￿ (A); i, distal part of basis of fifth pereiopod, ￿ (A);

j, basis of sixth pereiopod, ￿ (F); k, epimeral plates I-III, from the left, ￿ (A); 1, epimeral plates II and II, from the right, ￿ (A). BIJDRAGEN TOT DE DIERKUNDE, 50 - 1980 (1) 139

24. from Cueva la from Ramales de la Vic- Fig. Pseudoniphargus elongatus n. sp. (f Cascada, Spain, remaining figures fifth ￿ seventh ￿ basis of seventh ￿ d. toria, Spain): a, pereiopod, (scale I); b, pereiopod, (I); c, pereiopod, (I);

telson, ￿ (J); e, telson, ￿ (J); f, anomalous telson, ￿ (J).

from Cueva la from Ramales de la Vic- Fig. 25. Pseudoniphargus elongatus n. sp. (e Cascada, Spain, remaining figures

coxal of second first second toria, Spain): a, gill pereiopod, ￿ (scale A); b, uropod, ￿ (A); c, uropod, ￿ (A); d, third

third small third uropod, ￿ (F); e, uropod, $ (F); f, uropod, large ovigerous ￿ (F). 140 J. H. STOCK - REGRESSION MODEL EVOLUTION

Remarks. — From the most eastern locality flagellum segment (fig. 26b); flagellum 17-seg-

(in the province Guipûzcoa) only juveniles are mented; distal flagellum segments not exceedingly

known. Adults from this distal aesthete than distal province are necessary slender; slightly longer

to ascertain their taxonomie status. flagellum segment; penultimate and antepenul-

A from the Cueva la timate aesthetes than half the male Cascada has an more as long as

anomalous telson (fig. 24f), asymmetrical and corresponding segment (figs. 26d, e).

with instead of with Second a distal projection a distal antenna resembling that of Ps. uni-

notch. sexualis; flagellum rather short, 5- to 7-segmented

This species is characterized by the fact that also (fig. 26f).

the pedunculus of the female third uropod is Mandible palp as in Ps. africanus.

elongated; this elongation is less strong than in the First gnathopod: 6 palmar angle spines; smaller

male, but nevertheless clear in comparison with ones bicuspidate with terminal setule (fig. 26g);

Ps. the is the africanus (in Ps. africanus pedunculus larger ones bifid, posterior cusp longest (fig.

times in Ps. 4 iy 2 as long as wide, elongatus 26g).

times). The proposed specific name alludes to the Second gnathopod: propodus rather elongate;

found in the but also in with elongation 3rd uropod, posterior margin 3 groups of setae (fig. 26i);

the pereiopods 5 to 7, and in the Al flagellum. coxal plate elongate; basal stalk of coxal gill about

times wide 3 as long as (fig. 26h).

armed with 4. Oöstegites linear, 7 setae (fig. 26h). Pseudoniphargus sp. Coxal plates 3 and 4 elongate; stalk of coxal gill

Material examined. — Six juveniles. Spain, prov. elongate (figs. 28a, b). Remaining part of pereio-

Navarra, near French border, Cueva de las ( = W. Brujas 4 in pods 3 and (fig. 26j) as Ps. africanus, but of Zugarramurdi) ; in pool, entirely dark; 19 July 1979; leg. with 5 to 6 the sole. J. H., M. & D. Stock (ZMA coll. spinules on propodal no. Amph. 107.335). Pereiopod 5 (fig. 27a): basis rather elongate

Remarks. —-It is not to these possible identify (fig. 28c), posterior margin armed with 6 setules; juvenile specimens with certainty. They could well propodus with long spines. Stalk of coxal gills be Ps. but being recorded at the far elongatus, elongate. eastern end of the that of species makes me range Pereiopod 6 (figs. 27b, c): basis with 10 setules cautious. on posterior margin. Posterodistal corner of basis The the Las is locality, Brujas shown on caves, of P5 to P7 (6, ?) not overhanging (figs. 27d,

sheet T-23 of the Firestone scale Hispania map, e). The distal spines on the propodus of P5 to P7

1 : 200,000 footnote about (see 5). are as long as the claw in the female holo-

type (fig. 27c), shorter in the paratypes (fig.

of P7 rather slender Pseudoniphargus branchiatus n. sp. Figs. 26-28. 27d). Dactylus (fig. 28d).

Epimeral plates with a small tooth at the poste-

Material examined. — One ovigerous 9 (holo- rior corner. Lower margin of plate II with 2 spines; mouth of river Cullera type). Spain, prov. Valencia, Jugar at of III with 4 S. of 500 from plate spines (fig. 27f). (= Valencia), about m the sea; reeds, stones;

temp. 26.4° C, chlorinity 1400 mg/1; 9 Sep. 1970; leg. Uropod 1 (fig. 28e) rather slender; basofacial J. H. Stock & Pinkster S. (ZMA coll. no. Amph. 106.154). spine present; distal peduncle spine less than half Two S S seven 9 9 which five , (of ovigerous), para- as long as the rami. types. Spain, prov. Malaga, cave system of Hundidero-Gato

altitude of 28f near Benaojan (36°44'N 05°14'W); cave entrance Uropod 2 (fig. ) : pedunculus and rami with

423 In basin m. dripstone ("gours") in the large room; loamy several short marginal spines. gravel; temp. 14.0° C; 31 July 1978; leg. Biospeleo Werk- Uropod 3 of the female (figs. 28h, i): groep, sta. S 4-XI (ZMA coll. no. Amph. 107.384). pedun- twice culus over as long as wide; exopodite

Description. — Ovigerous females: length elongate, about 10 times as long as wide. Uropod

41/2-5V2 mm males: 4 mm. (without uropods); 3 of the male (fig. 28g): both pedunculus and

First antenna with rather slender curved and pedunculus ( fig. exopodite very elongate. 26a, c); flagellum than first accessory longer Telson (figs. 28j-l) with a shallow, rounded - 141 BIJDRAGEN TOT DE DIERKUNDE, 50 (1) 1980

26. branchiatus i from from basal Fig. Pseudoniphargus n. sp. (a, b, d, h, Cullera, Spain; c, e, f, g, j Benaojan, Spain): a,

of basal of first portion first antenna, ￿ (scale A); b, accessory flagellum, ￿ (B); c, portion antenna, ￿ (A); d, distal

flagellum segments of first antenna, ￿ (B); e, same of ￿ (B); f, second antenna, ￿ (A); g, distal part of first gnatho-

of second second pod, ￿ (B); h, coxal plate, oöstegite and coxal gill gnathopod, ￿ (A); i, distal part of gnathopod, ￿

(B); j, distal part of fourth pereiopod, ￿ (A). 142 j. H. STOCK - REGRESSION MODEL EVOLUTION

f from Cullera, b, d from Benaojan, Spain): fifth Fig. 27. Pseudoniphargus branchiatus n. sp. (a, c, e, Spain; a, pereiopod,

sixth ￿ seventh basis of ￿ (scale I); b, basis of sixth pereiopod, ￿ (K); c, pereiopod, (I); d, pereiopod, ￿ (K); e,

seventh pereiopod, ￿ (I); f, epimeral plates I-III, from the left, ￿ (L).

with REFERENCES distal notch; each laterodistal corner 3 to 5 9.

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Received: 5 March 1980