Carles Ribera

A phylogenetic analysis of myosin heavy chain type II sequences corroborates that Acoela and Nemertodermatida are basal bilaterians

Bilateria are currently subdivided into three superclades: Deuterostomia, Ecdysozoa, and Lophotrochozoa. Within this new taxonomic frame, acoelomate Platyhelminthes, for a long time held to be basal bilaterians, are now considered spiralian lophotrochozoans. However, recent 18S rDNA [small subunit (SSU)] analyses have shown Platyhelminthes to be polyphyletic with two of its orders, the Acoela and the Nemertodermatida, as the earliest extant bilaterians. To corroborate such position and avoid the criticisms of saturation and long-branch effects thrown on the SSU molecule, we have searched for independent molecular data bearing good phylogenetic information at deep evolutionary nodes. Here we report a phylogenetic analysis of DNA sequences from the myosin heavy chain type II (myosin II) gene from a large set of metazoans, including acoels and nemertodermatids. Our study demonstrates, both for the myosin II data set alone and for a combined SSU + myosin II data set, that Platyhelminthes are polyphyletic and that acoels and nemertodermatids are the extant earliest bilaterians. Hence, the common bilaterian ancestor was not, as currently held, large and complex but small, simple, and likely with direct development. This scenario has far-reaching implications for understanding the evolution of major body plans and for perceptions of the Cambrian evolutionary explosion.

A Review of Arthropod Phylogeny: New Data Based on Ribosomal DNA Sequences and Direct Character Optimization

Ribosomal gene sequence data are used to explore phylogenetic relationships among higher arthropod groups. Sequences of 139 taxa (23 outgroup and 116 ingroup taxa) representing all extant arthropod “classes” except Remipedia and Cephalocarida are analyzed using direct character optimization exploring six parameter sets. Parameter choice appears to be crucial to phylogenetic inference. The high level of sequence heterogeneity in the 18S rRNA gene (sequence length from 1350 to 2700 bp) makes placement of certain taxa with “unusual” sequences difficult and underscores the necessity of combining ribosomal gene data with other sources of information. Monophyly of Pycnogonida, Chelicerata, Chilopoda, Chilognatha, Malacostraca, Branchiopoda (excluding Daphnia), and Ectognatha are among the higher groups that are supported in most of the analyses. The positions of the Pauropoda, Symphyla, Protura, Collembola, Diplura, Onychophora, Tardigrada, and Daphnia are unstable throughout the parameter space examined.

Radiation of the Spider Genus Dysdera (Araneae, Dysderidae) in the Canary Islands: Cladistic Assessment Based on Multiple Data Sets

The volcanic archipelago of the Canary Islands, 100 km off the northwestern coast of Africa, harbors 43 endemic species of the mostly circum‐Mediterranean spider genus Dysdera (Araneae, Dysderidae). This amounts to approximately one‐fourth of all known Dysdera species in an area that represents 0.1% of the range of the genus. In order to address the origin of this extraordinary number of endemic species, the phylogenetic relationships among all the endemic taxa and a sample of 27 continental species were reconstructed. A simultaneous cladistic analysis was performed on 66 morphological characters, 471 bp of the cytochrome oxidase I and 424 bp of the 16S rRNA mitochondrial genes. The preferred most parsimonious tree supports a single origin for most of the endemic species (84%), although this tree is ambiguous regarding the total number of overseas colonizations (allowing a minimum of two and a maximum of four colonization events). Our data suggest that the Canary Islands have been the source of the colonizers of some of the remaining Macaronesian archipelagoes (certainly for the Selvagem Islands and the Cape Verdes and possibly for Madeira); the Azores have been independently colonized by dysderids from the continent. The present study provides a phylogenetic framework for an exceptional case of insular species radiation, an essential tool for unraveling the factors that have promoted this amazing diversification. Species radiations in oceanic archipelagoes are excellent models for the study of speciation processes.

The dark side of an island radiation: systematics and evolution of troglobitic spiders of the genus Dysdera Latreille (Araneae : Dysderidae) in the Canary Islands

The spider genus Dysdera Latreille is an excellent model for the study of the evolution of cave life: ten species are known to exist exclusively in the subterranean environment of the Canary Islands, where the genus has undergone local diversification. In the present paper, two new troglobitic species (Dysdera madai, sp. nov. and D. sibyllina, sp. nov.) and the previously unknown sex of five additional species are described and illustrated: the males of D. gollumi Ribera & Arnedo, 1994, D. hernandezi Arnedo & Ribera, 1999 and D. labradaensis Wunderlich, 1991; and the females of D. andamanae Arnedo & Ribera, 1997 and D. gibbifera Wunderlich, 1991. The first direct evidence of troglobitic members of Dysdera in micro- and mesocaverns are reported. The evolution of cave life as hypothesised following a combined morphological and molecular phylogeny is investigated. Troglobitic Canarian Dysdera species have colonised the under- ground on eight independent occasions. The Dysderidae groundplan represents a preadaptation to cave life and has facil- itated the colonisation of caves. Canarian members of Dysdera have a predominantly parapatric mode of speciation, although postspeciation changes in distribution may have obscured allopatric processes. Eye regression and, to a lesser extent, larger body size and appendage elongation characterise troglobitic species. The different levels of troglobiomor- phism are interpreted as local adaptations to heterogeneous subterranean conditions. The high levels of sympatry among troglobites are explained by trophic segregation and changes in prey capture strategy were involved in the single identi- fied case of subterranean speciation in the group.

From morphology and karyology to molecules. New methods for taxonomical identification of asexual populations of freshwater planarians. A tribute to Professor Mario Benazzi

Abstract Asexual reproduction by fissiparity is a frequent mode of reproduction in freshwater planarians (Turbellaria, Tricladida, Paludico‐la). Asexual populations pose a main taxonomical problem: to assign them to particular sexual species. We review here two case studies of asexual populations of freshwater triclads belonging to the genera Schmidtea and Dugesia. We show that karyotypes and morphology of the reproductive system of rare ex‐fissiparous specimens led to identifing as Schmidtea mediterranea the few asexual populations of this genus. Instead, morphology and karyology were unable to deal with the frequent asexual populations of Dugesia. We used the internal transcribed spacer region (ITS‐1) of ribosomal DNA to study 20 populations, sexual and asexual, of this genus. The pattern of sequence variation enabled the unambiguous assignment of all asexual populations to particular sexual species. The origin of asexual populations is briefly discussed.

Locomotor Activity Patterns and Feeding Habits in the Prawn Palaemon Serratus (Pennant, 1777) (Decapoda, Palaemonidae) in the Alfacs Bay, Ebro Delta, Spain

Diel activity rhythms of the prawn Palaemon serratus from Alfacs Bay, Ebro Delta, Spain, were studied under laboratory conditions by time-lapse video recording. The activity pattern showed endogenous rhythmicity of a circadian period with maximum activity at night. The feeding habits of P. serratus were studied using the frequency-of-occurrence method and the points method. The food of this species mainly consists of molluscs and crustaceans, and the remains of gastropods, amphipods, isopods, bivalves, mysids, copepods, and decapods were identified. The remaining items consisted of cnidarians, polychaetes, ophiuroids, plant material, sand, and unidentified organic debris. Results indicate that P. serratus is a predator of benthic invertebrates rather than a scavenger or detritus feeder. Diet composition changes with the size of the prawn.

Colonization and diversification of the spider genus Pholcus Walckenaer, 1805 (Araneae, Pholcidae) in the Macaronesian archipelagos: evidence for long-term occupancy yet rapid recent speciation.

Macaronesian archipelagos stand apart from other oceanic islands reputed as laboratories for the study of evolution by their proximity to the mainland, lack of subsidence, and steep ecological gradients. The genus Pholcus Walckenaer, 1805, commonly known as daddy-long-leg spiders, is one of the most speciose arthropod groups in the region, with 25 endemic species. In the present study, we use information from four mitochondrial genes, along with morphological data, to examine the phylogenetic relationships and diversification patterns of the genus in the region. Phylogenetic analyses support monophyly of Macaronesian Pholcus including the Moroccan species Pholcus vachoni and hence a single colonization event in the archipelagos. Madeira colonizers most likely originated from the Canaries, and a back-colonization of the nearby mainland receives further support. Estimated lineage divergence times suggest a long-time presence of Pholcus in the region, but also reveal that most present-day species are the result of recent, and probably rapid, speciation events. Diagnostic characters among Macaronesian Pholcus are confined to structures involved in copulation. Coupled with the extremely high diversification rate, the highest recorded for spiders, these copulatory characters suggest that sexual selection has played a key role in the local diversification of Pholcus in Macaronesia.

Diversity of Loxosceles spiders in Northwestern Africa and molecular support for cryptic species in the Loxosceles rufescens lineage

Until recently, Loxosceles rufescens was the only species known from a geographic range including Northern Africa, Mediterranean Europe and the Middle East. Rich Loxosceles diversity in the New World suggests either that L. rufescens is a young lineage or that its diversity is underappreciated. We use a molecular phylogenetic and morphological approach to examine diversity in L. rufescens and other Loxosceles lineages in Northwestern Africa. Molecular analyses of one nuclear and two mitochondrial genes strongly support a monophyletic clade including L. rufescens, the Northern Brazilian L. amazonica and three other divergent Northwestern African lineages, though relationships among them remain unresolved. A genetically divergent Moroccan individual morphologically consistent with L. rufescens was strongly supported as sister to all other putative L. rufescens, consistent with the presence of at least 2 species in this lineage. COI p-distances and population structuring among remaining putative L. rufescens clades further suggest the absence of gene flow between clades and the possibility that they represent multiple species. Morphological characters of preserved Loxosceles collected in a range of African countries provide additional indication that Loxosceles are more diverse and have a deeper history in Africa than has been previously understood.

SYSTEMATICS OF THE GENUS DYSDERA (ARANEAE, DYSDERIDAE) IN THE EASTERN CANARY ISLANDS

Abstract The circum-Mediterranean spider genus Dysdera has undergone an outsanding species radiation in the volcanic archipelago of the Canary Islands. The present study deals with the endemic species that inhabit the older and ecologically distinct islands of Fuerteventura, Lanzarote and their nearby islets. A new species, Dysdera sanborondon, is described. The male of D. spinidorsum Wunderlich 1991, is described for the first time. Five species are redescribed: D. alegranzaensis Wunderlich 1991; D. lancerotensis Simon 1907; D. longa Wunderlich 1991; D. nesiotes Simon 1907, and D. spinidorsum Wunderlich 1991. The species D. liostethus Simon 1907 is proposed to be a senior synonym of D. clavisetae Wunderlich 1991 and its presence in the eastern islands is considered to be doubtful. A neotype is designated for D. nesiotes. The distribution of D. alegranzaensis is extended to Lanzarote and the other northern islets. Dysdera nesiotes is reported for the first time in the eastern Canaries. Morphological affinities and distribution patterns are discussed. The remarkably lower number of endemic species harbored by the eastern islands, when compared with other Canarian islands similar in size but younger in age, is proposed to be the result of a major extinction event in the eastern Canaries due to climatic change.

Biology of the mysid Mesopodopsis slabberi (van Beneden, 1861)(Crustacea, Mysidacea) in a coastal lagoon of the Ebro delta (NW Mediterranean)

Aspects of the population and reproductive biologyof the mysid Mesopodopsis slabberi were studiedin the western Mediterranean (Encanyissada coastallagoon, Ebre delta, Spain). Misids were sampling atmonthly intervals from August 1993 to June 1995. InDecember 1994 and February 1995 any individuals werefound. The relationship between carapace length (Lc)and total length (Lt) was isometric:logLt = 1.0612logLc + 0.9411 (n = 178,r = 0.9411). Thesize of mature males and females was at a maximum inthe winter, and a minimum in the summer. The longevityof individuals changed throughout the year; those withthe longest life expectancy were members of theoverwintering generation. Brooding females wererecorded throughout the year, except in December 1993and April 1994; they were most numerous in spring andautumn. The number of embryos or larvae (N) carried byfemales was related to the size of the females (range:1–22, mean value: 4.8): logN = 1.8705logLc−0.0985(n = 361, r = 0.4097, P < 0.01). Thesize of the eggs(maximum diameter, Le) was related to the size of thefemales (range: 0.35–0.6 mm in total length):logLe = 0.3404logLc−0.4820 (n = 277, r = 0.5420, P < 0.01).