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(Crustacea), with Emphasis on the Iberian Species

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(Crustacea), with Emphasis on the Iberian Species Bijdragen tot de Dierkunde, 58 (2): 159-204 — 1988 Phylogenetic relationships and biogeography of the groundwater-dwellingamphipod genus Pseudoniphargus (Crustacea), with emphasis on the Iberian species by Jos Notenboom Institute of Taxonomic Zoology, University of Amsterdam, P.O. Box 4766, 1009 AT Amsterdam, The Netherlands Abstract El grado de diferenciación morfológica de Pseudoniphar- los gus es bajo y potenciales apomorfismos disponibles son Numerical phylogenetic methods are applied in order to insuficientes resolver las relaciones a ni- para filogenéticas arrive at synapomorphic similarities between species of vel de especie. Ello puede ser atribuible a un modelo de the Character stygobiont amphipod genus Pseudoniphargus. especiación peripátrica subyacente, a problemas de discri- polarity is assessed by comparison with the relevant minación de estados de carácter mutualmente excluyentes Parapseudoniphargus and Allomelita within a outgroups , alta de evolución inde- y homólogos, y a una probabilidad cluster of interrelated of Hadzioidea. presumedly genera de Los la pendiente caracteres. problemas que plantea re- The degree of morphological differentiation of construcción filogenética de niveles taxonómicos bajos son is low and the available Pseudoniphargus potential apomor- universales. En el de el caso un grupo estigobionte, troglo- insufficient resolve phies are to phylogenetic relationships la reducción estructural la morfismo, y morfología sene- to the species level. The inconsistency in the data is high, scente provocan problemas peculiares. which be result of may a an underlying peripatric specia- Los resultados del estudio conducen al reconocimiento tion model, difficulties in discriminating mutual exclusive de monofiléticos bien de los cuatro grupos definidos, tres and homologous character states, and a high probability cuales consisten únicamente en especies hermanas. Los of independent character evolution. Problems of sin taxones restantes quedan agrupar o pertenecen a gru- phylogeny reconstruction at low taxonomic levels are han evidenciar de todas las pos que no se podido a pesar Accidental in the of universal. problems case a stygobiont numéricas. Los monofiléticos de aproximaciones grupos caused structural reduc- group are by troglomorphism, Pseudoniphargus son, en su mayor parte, geográficamente tion, and senescent morphology. circunscriben a los Montes coherentes y se Cantábricos, o The results of the study lead to the recognition of four área Lusitano-Atlántica o Cordillera Bética. Por último, well-corroborated three consist of monophyletic groups; los se discute el posible origen de Pseudoniphargus a partir sister only. taxa are or species Remaining ungrouped de de un ancestro talasoestigobionte originario aguas so- which do after all numerical belong to groups not emerge meras del área oriental atlántica del mar de Tethys. The of approaches. monophyletic groups Pseudoniphargus for the coherent and are greater part geographically con- Cantabrian fined either to the Mountains, or to the Lusitanian-Atlantic region, or to the Betic Cordillera. The Contents from possible origin of Pseudoniphargus a thalassostygobiont 1. Introduction 160 which in the Atlantic of the progenitor arose eastern part 2. Material and methods 161 shallow Tethys sea is discussed. 3. Characteristics of Pseudoniphargus 163 3.1 Comprehensive diagnosis 163 3.2 Distribution and biology 165 4. The phylogenetic position of 167 Pseudoniphargus .... Resumen 4.1 Classificatory problems 167 Se han aplicado métodos filogenéticos numéricos con el fin 4.2 Delimitation of presumedly related taxa 167 de obtener similitudes sinapomórficas entre especies de 4.3 Numerical phylogenetic analysis 169 de La 5. within 174 Pseudoniphargus, género anfípodos estigobiontes. po- Phylogenetic relationships Pseudoniphargus laridad de los ha sido determinada 5.1 Character 174 caracteres por compa- analysis ración Allomelita, 5.2 the characters 177 con Parapseudoniphargus y grupos externos Explanatory notes to especialmente significativos, en el contexto de un cluster 5.3 Numerical phylogenetic analysis 182 de Hadzioidea of of Pseudo- géneros de presumiblemente inter- 5.4 Diagnosis monophyletic groups relacionados. niphargus 187 160 - J. NOTENBOOM PSEUDONIPHARGUS PHYLOGENY AND BIOGEOGRAPHY 6. Discussion of results 188 phylogenetic polychotomous branching were fairly neglected 6.1 Character and unresolved incompatibility in these studies. Furthermore, the groups under relationships 188 the of consideration are rarely defined on basis 6.2 Hypogean features as complicating factors shared derived characters in phylogeny reconstruction 189 (synapomorphies). Modern is based 6.3 Phylogenetic methodology 191 biogeography on phylogenetic 7. of 192 Origin Pseudonipharguss and biogeography hypotheses and inferences of the history of a 199 Acknowledgements from that both group phylogeny (Wiley, 1981), References 200 distributional and paleogeographic data being used. Biogeographic studies of stygobiont amphipods are, however, often based on 1. INTRODUCTION weakly developed phylogenetic hypotheses. The of the is purpose present study to explore have invaded the of the Amphipods very successfully a phylogenetic relationships genus wide of subterranean habitats, with on the Iberian range aquatic Pseudoniphargus , emphasis from marine to continental hypogean waters, species, by using phylogenetic numerical and from of sediments sub- interstitia microporous techniques. Pseudoniphargus is an important in the literature the of to large karstic basins. Those amphipods that ject on biogeography are obligate dwellers of groundwater habitats stygobiont animals. The relationship between constitute (stygobionts) a taxonomically highly hypothesized cladogenetic events within the diverse and the of the distribu- group, with presumed multiple origins genus paleogeography from ancestral tional of the used different stocks. They are areas species was by Stock worldwide in distribution, and known from (1980a) to exemplify the "Regression Model". and This formulated Stock tropical, subtropical, temperate excep- model, as originally by tionally also from cold regions, in general show- (1977), gives explanations for the distribution, of endemism. in and ing a high degree Dispersal brackish fresh continental, but mainly abilities of the stygofauna are limited, owing to insular, groundwaters, of stygobionts from their intrinsic properties and stratigraphie direct marine origin. In the literature devoted barriers. to the biogeography of stygobiont animals, the studies far dis- Many Zoogeographie so dedicated "Regression Model" has frequently been the distribution of cussed Rouch to intriguing patterns (see e.g., Danielopol, 1980; & focus the the few stygobiont amphipods on greater Danielopol, 1987). During past years, Caribbean of have been and peri-Mediterranean regions. many new species Pseudoniphargus from Historical events, such as continental drift, described Atlantic and western peri- from marine regressions, and tectonic uplift are con- Mediterranean regions, especially the sidered have Peninsula to had an important effect on the Iberian (Notenboom, 1986, 1987a, present distribution patterns. Developments in b; Stock et al., 1986; Stock, 1988; Boutin & vicariance biogeography and area cladistics Coineau, 1988). Through these new findings it have had influence became evident of a noticeable on recently that the distribution pattern studies with characters and the the published zoogeographical dealing relationships among taxa of much than groups stygobiont amphipods (Stock, 1980a; are more complex was thought before. For future discussions the Holsinger, 1986; Stock & Rondé-Broekhuizen, on "Regres- 1986; Boutin & Coineau, 1988; Boutin & sion Model" and the evolution of stygobiont the it submit the Messouli, 1988). However, cladograms animals, was deemed desirable to in these be based presented papers seem to on genus to a phylogenetic study. character An obstacle a global concept concerning state important in phylogenetic transformation rather than on a phylogenetic studies of amphipods at lower taxonomic levels analysis. Comparisons with closely related is the highly debated classification into families and and all that of the outgroups, homology of character states, superfamilies, above BIJDRAGEN TOT DE DIERKUNDE, 58 (2) - 1988 161 Gammaridea. suborder Probably none of the The intention of this paper is to find classifications reflects the within actually proposed monophyletic groups Pseudoniphargus. natural relationships and very little is known The methodology used is repeatable and thus about the synapomorphic similarity between the result testable. It willbe shown that the data The external features taxa. morphological cur- are inadequate to provide a single solution. rently used in the systematics of Gammaridea Tree topologies of equally parsimonious solu- to natural tions and conclusions about probably are inadequate provide a are compared classification rather limited of this group. Attempts to explore affinities will be drawn for a character have been made Lincoln number of The method of new sets by taxa only. outgroup & Hurley (1981) and Lincoln (1984): calceoli; comparison is used to assess apomorphous end by Bousfield (1985): reproduction and mating character states, and to that an outgroup and of strategies; by Halcrow & Bousfield (1987): hypothesis Pseudoniphargus was developed. surface microstructures. These developments, Foibles in the data set leading to unresolved in
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