Received: 17 September 2004 / Accepted: 21 April 2021 / Published online: 17 September 2004
Allopolyploid evolution in Geinae (Colurieae: Rosaceae) – building reticulate species trees from bifurcating gene treesDownload PDFAbstract
A previous phylogenetic study of paralogous nuclear low-copy granule-bound starch synthase (GBSSI) gene sequences from polyploid and diploid species in Geinae indicated that the clade has experienced two major allopolyploid events in its history. These were estimated to have occurred several million years ago. In this extended study we test if the reticulate phylogenetic hypothesis for Geinae can be maintained when additional sequences are added. The results are compatible with the hypothesis and strengthen it in minor aspects. We also attempt to identify extant members of one of the inferred ancestral lineages of the allopolyploids. On the basis of previous molecular phylogenies, one specific group has been proposed to be the descendants of this taxon. However, none of the additional paralogues belong to this ancestral lineage. A general method is proposed for converting a bifurcating gene tree, with multiple paralogous low-copy gene sequences from allopolyploid taxa, into a reticulate species tree.
Cumacea species (Crustacea: Peracarida) from the deep-sea expedition DIVA-1 with RV “Meteor” to the Angola Basin in July 2000. Family LeuconidaeDownload PDFAbstract
Seven new deep-sea species from the Angola Basin are described from the family Leuconidae with the genus Leucon with five species, including the subgenera Crymoleucon Watling, 1991 and Epileucon (Atlantide Rep. 4 (1956) 183) with one species each, and Macrauloleucon Watling, 1991 with three species and the genera Bytholeucon Watling, 1991, and Eudorella Norman, 1867 with one species each. Only one species of the family Leuconidae in the present material—Leucon (Leucon) homorhynchus Bishop, 1981b—was already described.
Received: 16 February 2003 / Accepted: 21 April 2021 / Published online: 16 February 2003
Proper names in twin worlds: Monophyly, paraphyly, and the world around usDownload PDFAbstract
This paper addresses the theoretical relevance of monophyletic, paraphyletic and polyphyletic groups under the paradigm of sophisticated scientific realism. The doctrine of metaphysical realism is introduced using the philosophy of Karl Popper as an example, which is then contrasted with scientific realism. A discussion of the nature of causal relations presents an account of counterfactual conditionals. The current state of art casts the theory of phylogenetic systematics in a stark contrast of classes (universals) and individuals (particulars). In practice, however, individuals piggyback on classes, or sets. Natural kinds are introduced in order to overcome this deep dichotomy. The theoretical relevance of natural kinds lies in their explanatory value, and that may change with changing context. It is for this reason that non-monophyletic groups can have explanatory value (their members can function as tokens of causally relevant kinds) within certain domains of evolutionary biology. Explanatory value is maximized by integration of the genealogical hierarchy of species and monophyletic taxa with other areas of evolutionary biology.
On two rare abyssal Myriotrochidae (Echinodermata: Holothuroidea: Apodida) new to the South Atlantic: Siniotrochus myriodontus Gage and Billett, 1986 and Neolepidotrochus parvidiscus angolensis subsp. nov.Download PDFAbstract
In the course of the DIVA 1 expedition (RV “Meteor”, cruise M48/1) to the Angola Basin (South Atlantic) two myriotrochid holothurians were collected at abyssal depths – Neolepidotrochus parvidiscus angolensis subsp. nov. and Siniotrochus myriodontus Gage and Billett, 1986. The latter, until now, was known only from few specimens from the north-eastern Atlantic Ocean. This new record of S. myriodontus for the South Atlantic Ocean indicates a wide geographical distribution. Both species are described in detail, including parameters for the wheel deposits.
Received: 20 August 2004 / Accepted: 21 April 2021 / Published online: 20 August 2004
The phylogeny of the Hyles euphorbiae complex (Lepidoptera: Sphingidae): Molecular evidence from sequence data and ISSR-PCR fingerprintsDownload PDFAbstract
The evolutionary history of the Hyles euphorbiae complex (HEC) was studied using mitochondrial DNA sequences comprising about 2300bp derived from the genes cytochrome c oxidase subunits I (COX I) and II (COX II) and tRNA-leucine. In addition, we collected genomic fingerprinting data by ISSR-PCR to assess if and how the bi-parentally inherited nucleome may have diverged differently to the maternally inherited mitochondria. The COX sequences revealed a clear geographical pattern of genetic differentiation of the HEC into two main lineages, H. euphorbiae and H. tithymali. Our results provide no evidence that H. dahlii falls within a HEC s. str., although a sister-group relationship cannot be ruled out. The sequence data indicated intraspecific subdivisions and gene flow patterns, and possibly detected both introgression and a major contact zone on Mediterranean islands between these two evolutionary lineages. Hyles tithymali is hypothesized to have been able to retain ancient polymorphisms until the present, whereas H. euphorbiae appears to have (re)colonized its current distribution range after the Ice Ages from a few (or even only one) refugial populations by leptokurtic dispersal, resulting in low diversity. The ISSR-PCR data showed much higher variability among individuals of the HEC than did mtDNA sequence data. They provided insights into the genomic distribution of the simple sequence repeat (GACA)4 and appear to describe a more complex pattern of introgression in the HEC. Our data revealed the HEC as a very young species complex, in which we have detected two distinct mitochondrial lineages, corresponding to H. tithymali (including the deserticola, mauretanica and himyarensis lineages) and H. euphorbiae (including H. robertsi), respectively.
Received: 03 January 2004 / Accepted: 21 April 2021 / Published online: 03 January 2004
Studies on the genus Onisimus Boeck, 1871 (Crustacea, Amphipoda, Lysianassoidea, Uristidae)Part I. The brevicaudatus and sextonae species groupsDownload PDFAbstract
The genus Onisimus Boeck is divided into five species groups to be treated in this and forthcoming publications; a key to these groups includes the genera Menigrates Boeck and Paralibrotus Stephensen. Further keys and redescriptions are presented to all species within two Onisimus species groups. The O. brevicaudatus-group comprises the species O. affinis Hansen, 1886 (=O. dubius Schellenberg, 1935, syn. n.), O. botkini Birula, 1897, O. brevicaudatus Hansen, 1887, O. caricus Hansen, 1887, and O. derjugini Gurjanova, 1929. The O. sextonae-group consists of the species O. abyssi Oldevig, 1959, O. leucopis (G.O. Sars, 1879), and O. sextonae Chevreux, 1926. The full redescriptions of these species are provided in an accompanying Organisms Diversity and Evolution Electronic Supplement.See also Electronic Supplement at: http://www.senckenberg.de/odes/05-07.htm
Received: 10 May 2003 / Accepted: 21 April 2021 / Published online: 10 May 2003
Reinterpretation of an alleged marine hexapod stem-group representativeDownload PDFAbstract
The appendages of the recently described lower Devonian arthropod Devonohexapodus, originally assigned to the stem group of Hexapoda, probably have been misinterpreted. There is no convincing evidence for the evolutionary loss of the first post-antennal pair of appendages and the post-mortem loss of another two pairs of anterior appendages. Three pairs of particularly long appendages in the anterior body portion possibly are not homologous to the thoracic legs of insects. The fossil does not belong in the stemgroup of Hexapoda.
A parthenogenetic, simplified adult in the life cycle of Pliciloricus pedicularis sp. n. (Loricifera) from the deep sea of the Angola Basin (Atlantic)Download PDFAbstract
A new species, Pliciloricus pedicularis (Pliciloricidae, Loricifera), is described inhabiting fine-grained clayish sediments in the deep sea of the Angola Basin. This is the first report of a Pliciloricus-species with a simplified parthenogenetic adult in its life cycle. The simplified adult is a non-free-living stage differing morphologically considerably from the free-living bisexual adults. It has a sack-like body without an introvert but with a persisting neck region covered with hooks or spiny pads. The sack-like body contains mainly the mature ovary. Large eggs are released into a shelter formed by the exuvium of the last or seventh instar Higgins-larva. Both types of adults, the parthenogenetic as well as the bisexual ones, are surrounded during metamorphosis by two exuviae: a simple inner one as rest of the postlarval stage and an outer one belonging to the seventh instar Higgins-larva. The bisexual adult of the new species is characterized by type B spinoscalids in the fourth row basally equipped with a ventral row of minute denticles; long rigid trichoscalids basally with numerous strong cross walls; small cuticular bars directly above the well-defined edge of the lorica, and a lorica consisting of 44 primary plicae. Distinguishing features of the Higgins-larva of the new species are: very long toes, and clavoscalids with a dorsal row of spinules and an additional fourth segment. The most conspicuous feature of the Higgins-larva is the stem-like basal part of the anterior setae called pedicels. Especially the enlarged pedicels of the posterolateral setae enable the larva to be determined even when the introvert is retracted. The study revealed new information about embryology and development, which have never been observed in any Loricifera-species so far. The great abundance of the new species in samples from the Angola Basin allows the identification of most life history stages and developmental instars.
Received: 16 March 2004 / Accepted: 21 April 2021 / Published online: 16 March 2004
Species diversity within the Morchella esculenta group (Ascomycota: Morchellaceae) in Germany and FranceDownload PDFAbstract
The distribution of the Morchella esculenta group in Germany and France is examined based on 22 samples, a sample from Montenegro is studied as well. In the recent literature the group was often treated as a single species, M. esculenta sensu lato. Our study, based on the polymorphism of the internal transcribed spacer (ITS) region within the nuclear ribosomal DNA (nrDNA), indicates the presence of three distinct species: M. esculenta (L.) Pers., M. crassipes (Vent.) Pers. : Fr., and M. spongiola Boud. They can be identified easily by restriction fragment length polymorphisms (RFLP) of the ITS region.
Notes on the Tanaidacea (Crustacea: Peracarida) of the Angola BasinDownload PDFAbstract
In July 2000 the DIVA-1 Expedition collected deep-water benthos (5125–5449m) in the Angola Basin, South Atlantic Ocean. The tanaidacean material is composed of 434 individuals with a body length between 1.2 and 12mm. Sixty-eight species, belonging to 10 families were found. Tanaidomorpha with 63 species represent 96% of the material, Neotanaidomorpha with one species 3% and Apseudomorpha with 4 species 1%. The most diverse genus is Leptognathia (22 species). The material is dominated by Paranarthrura angolensis with 58 individuals. The Tanaidacea of the Angola Basin share only one species (Neotanais americanus) with other deep-sea areas of the Atlantic Ocean, 97% of the material is composed of new species. The different sampling gear results are discussed.
Received: 27 August 2004 / Accepted: 21 April 2021 / Published online: 27 August 2004
Evolutionary relationships in the Asteraceae tribe Inuleae (incl. Plucheeae) evidenced by DNA sequences of ndhF; with notes on the systematic positions of some aberrant generaDownload PDFAbstract
The phylogenetic relationships between the tribes Inuleae sensu stricto and Plucheeae are investigated by analysis of sequence data from the cpDNA gene ndhF. The delimitation between the two tribes is elucidated, and the systematic positions of a number of genera associated with these groups, i.e. genera with either aberrant morphological characters or a debated systematic position, are clarified. Together, the Inuleae and Plucheeae form a monophyletic group in which the majority of genera of Inuleae s.str. form one clade, and all the taxa from the Plucheeae together with the genera Antiphiona, Calostephane, Geigeria, Ondetia, Pechuel-loeschea, Pegolettia, and Iphionopsis from Inuleae s.str. form another. Members of the Plucheeae are nested with genera of the Inuleae s.str., and support for the Plucheeae clade is weak. Consequently, the latter cannot be maintained and the two groups are treated as one tribe, Inuleae, with the two subtribes Inulinae and Plucheinae. The genera Asteriscus, Chrysophthalmum, Inula, Laggera, Pentanema, Pluchea, and Pulicaria are demonstrated to be non-monophyletic. Cratystylis and Iphionopsis are found to belong to the same clade as the taxa of the former Plucheeae. Caesulia is shown to be a close relative of Duhaldea and Blumea of the Inuleae-Inulinae. The genera Callilepis and Zoutpansbergia belong to the major clade of the family that includes the tribes Heliantheae sensu lato and Inuleae (incl. Plucheeae), but their exact position remains unresolved. The genus Gymnarrhena is not part of the Inuleae, but is either part of the unresolved basal complex of the paraphyletic Cichorioideae, or sister to the entire Asteroideae.
Received: 30 April 2004 / Accepted: 21 April 2021 / Published online: 30 April 2004
Caprellids (Crustacea: Amphipoda: Caprellidae) from shallow waters of the Caribbean coast of VenezuelaDownload PDFAbstract
The caprellidean fauna of Venezuela is investigated. Twenty-seven stations in shallow waters of the states Falcón, Carabobo, Aragua, Anzoátegui, Sucre and Nueva Esparta were sampled, and the caprellids were sorted and identified. A total of 3984 specimens were identified; 12 species in eight genera are reported. Two species, Caprella penantis Leach, 1814 and Paracaprella digitimanus Quitete, 1971, are new records for Venezuelan waters. The electronic supplement to this paper includes illustrations and a key to all caprellid species known from Venezuela, as well as comments on their distribution and ecology.
Received: 07 April 2004 / Accepted: 21 April 2021 / Published online: 07 April 2004
Redescription of Ischyromene lacazei Racovitza, 1908 (Isopoda: Sphaeromatidae) from the Mediterranean Coast of southern FranceDownload PDFAbstract
The sphaeromatid species Ischyromene lacazei Racoviza, 1908 is rediagnosed, and new data on its morphology and distribution are given, based on new material from the type locality, the intertidal zones at Banyuls-sur-mer, southern France. Groups of specimens were found in empty shells of Balanus perforatus and Chthamalus stellatus (Crustacea: Balanomorpha); the formation of harems was not observed. The full redescription, further data and a discussion are provided in an accompanying Organisms Diversity and Evolution Electronic Supplement.See also Electronic Supplement at: http://www.senckenberg.de/odes/05-08.htm
Diversity of peracarid crustaceans (Malacostraca) from the abyssal plain of the Angola BasinDownload PDFAbstract
During the expedition DIVersity of the abyssal Atlantic benthos (DIVA-1) with RV “Meteor” in July 2000, samples were taken at seven stations by means of an epibenthic sledge north of the Walvis Ridge in the Angola Basin off Namibia in 5125–5415m depth. Two hundred and forty one species of Peracarida are identified from the material so far. Dominant elements of the peracarid fauna were Isopoda, which were most abundant and diverse, 100 species were identified from 1326 individuals, followed by Tanaidacea with 50 species and 194 individuals, and Cumacea with 45 species and 479 individuals. Amphipoda were less frequent with 39 species and 150 individuals, Mysidacea were rarest yielding only 7 species and 34 individuals. The fauna is characterized by 118 rare species, most of them occurring only with single specimens at one station. Only 123 species occur at more than one station and only two species of the Eurycopinae (Isopoda) at all stations. The few species which are already known are either cosmopolitan or typical for the Atlantic Ocean, while elements known from the Southern Ocean are rare indicating that the Walvis Ridge is an effective distribution barrier for deep-sea organisms.
Received: 18 April 2004 / Accepted: 21 April 2021 / Published online: 18 April 2004
Phylogenetic analysis of the Malvadendrina clade (Malvaceae s.l.) based on plastid DNA sequencesDownload PDFAbstract
Phylogenetic relationships within Malvaceae s.l., a clade that includes the traditional families Bombacaceae, Malvaceae s.str., Sterculiaceae, and Tiliaceae, have become greatly clarified thanks to recent molecular systematic research. In this paper, we use DNA sequences of four plastid regions (atpB, matK, ndhF, and rbcL) to study relationships within Malvadendrina, one of the two major clades of Malvaceae s.l. The four data sets were generally in agreement, but five terminal taxa manifested highly unexpected affinities in the rbcL partition, and the non-coding sequences of the trnK intron were found to provide limited phylogenetic information for resolving relationships at the base of Malvadendrina. The remaining data strongly support the existence of six major clades within Malvadendrina: Brownlowioideae, Dombeyoideae, Helicteroideae, Malvatheca (comprising Bombacoideae and Malvoideae), Sterculioideae, and Tilioideae. These data also resolve the placement of two problematic taxa: Nesogordonia (in Dombeyoideae) and Mortoniodendron (in Tilioideae). The relationships among the six clades are not definitively resolved, but the best-supported topology has Dombeyoideae as sister to the remainder of Malvadendrina (posterior probability PP=80%) and Sterculioideae as sister to Malvatheca (PP=86%). This early branching position of Dombeyoideae is supported by similarities in floral characters between members of that clade and outgroup taxa in Byttnerioideae. Similarly, the sister-group relationship of Sterculioideae and Malvatheca receives support from androecial characteristics, like subsessile or sessile anthers and an absence of staminodes, shared by these two clades.
Received: 27 July 2004 / Accepted: 21 April 2021 / Published online: 27 July 2004
A new eocene Chascacocolius-like mousebird (Aves: Coliiformes) with a remarkable gaping adaptationDownload PDFAbstract
A skull of a new species of mousebird (Aves: Coliiformes) is described from the Middle Eocene of Messel in Germany. Chascacocolius cacicirostris n. sp. is the fifth coliiform species described from the Messel deposits, and a further example of the remarkable similarity between the early Eocene avifaunas of North America and Europe. As for its much smaller North American counterpart, C. oscitans Houde & Olson, 1992, the new species has greatly elongated retroarticular processes on the mandible, which are an adaptation to gaping, i.e. opening of the bill in a substrate. The cranium and upper beak of Chascacocolius are seen for the first time in the new specimen; the latter shows a striking resemblance to the upper beak of some modern New World Blackbirds (Passeriformes: Icteridae). C. cacicirostris n. sp. thus provides another example of the diversification of early Tertiary Coliiformes, some taxa of which apparently occupied feeding niches that today are filled by songbirds (Passeriformes).
Received: 16 December 2002 / Accepted: 21 April 2021 / Published online: 16 December 2002
Squamate phylogeny, taxon sampling, and data congruenceDownload PDFAbstract
To investigate the affinities of snakes, amphisbaenians and dibamids, the phylogenetic relationships among the major lineages (families) of extinct and extant squamates are assessed through a combined analysis of 248 osteological, 133 soft anatomical, and 18 ecological traits. The osteological data set represents a revision of previous data, taking into account recent criticism; the ecological data set is new. In addition, potentially critical fossil taxa (polyglyphanodontids and macrocephalosaurs) are included for the first time. The osteological and soft anatomical data sets each place snakes within anguimorphs, with dibamids and amphisbaenians near gekkotans. The putative primitive fossil amphisbaenian Sineoamphisbaena groups with macrocephalosaurs and polyglyphanodontids, together the sister group to scleroglossans. All three data sets are congruent, and these results are reinforced by combined analyses. In these, as in the osteological analyses, snakes are nested within marine lizards. However, exclusion of fossil taxa from the osteological data set results in a ‘limbless clade’ consisting of snakes, amphisbaenians and dibamids, and introduces significant conflict between osteology and soft anatomy. Also, deletion tests and character weighting reveal that the signal in the reduced osteological data set is internally contradictory. These results increase confidence in the arrangement supported by the all-taxon osteological, the soft anatomical, and the combined data, and suggest that exclusion of fossils confounds the signal in the osteological data set. Finally, the morphological data support the nesting of snakes within marine lizards, and thus a marine origin of snakes. This result still holds when relationships between living forms are constrained to the topology suggested by molecular sequences: if marine lizards are allowed to ‘float’ within this molecular framework, they form the stem group to snakes, and do not group with varanids as previously suggested.See also Electronic Supplement at: http://www.senckenberg.de/odes/05-04.htm.
Chauliodoniscus coronatus sp. nov., a new deep-sea species from the Angola Basin (Crustacea, Isopoda, Asellota, Janiroidea, Haploniscidae)Download PDFAbstract
A new isopod species of the family Haploniscidae is described. The new species was discovered in samples collected from the Angola abyssal plain (south-east Atlantic). Differences to related species of the genus are discussed. A row of several small tooth-like processes at the anterior margin of the cephalon is a conspicuous character of Chauliodoniscus coronatus sp. nov., which distinguishes it from all other members of the genus Chauliodoniscus.
Received: 17 June 2004 / Accepted: 21 April 2021 / Published online: 17 June 2004
Proventricular structure in solitary bees (Hymenoptera: Apoidea)Download PDFAbstract
Proventricular structure, analyzed by scanning electronic microscopy, is compared among 28 species of solitary bees representing four families. Observations on the shapes of proventricular folds and on hair-like cuticular projections are presented, discussed, and suggested as useful to future studies of bee systematics.
Received: 20 May 2004 / Accepted: 21 April 2021 / Published online: 20 May 2004
Origin and age of Australian ChenopodiaceaeDownload PDFAbstract
We studied the age, origins, and possible routes of colonization of the Australian Chenopodiaceae. Using a previously published rbcL phylogeny of the Amaranthaceae–Chenopodiaceae alliance (Kadereit et al. 2003) and new ITS phylogenies of the Camphorosmeae and Salicornieae, we conclude that Australia has been reached in at least nine independent colonization events: four in the Chenopodioideae, two in the Salicornieae, and one each in the Camphorosmeae, Suaedeae, and Salsoleae. Where feasible, we used molecular clock estimates to date the ages of the respective lineages. The two oldest lineages both belong to the Chenopodioideae (Scleroblitum and Chenopodium sect. Orthosporum/Dysphania) and date to 42.2–26.0 and 16.1–9.9Mya, respectively. Most lineages (Australian Camphorosmeae, the Halosarcia lineage in the Salicornieae, Sarcocornia, Chenopodium subg. Chenopodium/Rhagodia, and Atriplex) arrived in Australia during the late Miocene to Pliocene when aridification and increasing salinity changed the landscape of many parts of the continent. The Australian Camphorosmeae and Salicornieae diversified rapidly after their arrival. The molecular-clock results clearly reject the hypothesis of an autochthonous stock of Chenopodiaceae dating back to Gondwanan times. Instead, they indicate that most lineages arrived in Australia via long-distance dispersal. Some lineages (e.g. the Halosarcia lineage) may have used the Indonesian archipelagos as stepping stones. The authors are aware that estimates of diversification times using a molecular clock can be subject to considerable levels of error. Our estimates of the age of Australian chenopod lineages based on three alternative fossils were made independently from any knowledge about shifts in climatic and geographical conditions in Australia during the times of arrival. In most cases, however, the paleoclimatic scenario indicates habitat shifts suitable for the respective chenopod colonizer, which corroborates our findings and provides a plausible scenario.
A new species of Paranannopus Lang, 1936 (Copepoda, Harpacticoida, Pseudotachidiidae) with atrophic mouthparts from the abyssal of the Angola BasinDownload PDFAbstract
Within the multicorer deep-sea samples of the DIVA 1 expedition (METEOR cruise M48/1) some male harpacticoid specimens with strongly reduced mouthparts have been collected, each of them representing a hitherto unknown species. Despite their atrophic mouthparts, they can be identified easily as members of the Paranannopinae Por, 1986 because of showing the characteristic swimming-leg sexual dimorphisms of the latter.In the present paper, a species of Paranannopinae with reduced mouthparts is described for the first time. At the present state of systematic discussion the new species has to be placed within Paranannopus Lang, 1936.Males with atrophic mouthparts have been sporadically described also from other harpacticoid families (such as e.g. Argestidae Por, 1986 and Aegisthidae Giesbrecht, 1892) and the phenomenon is likely to be sexually dimorphic. This kind of sexual dimorphism within harpacticoid copepods combined with enlarged antennular aesthetacs seems to be predominantly a deep-sea phenomenon.
A short note on the cephalopods sampled in the Angola Basin during the DIVA-1 expeditionDownload PDFAbstract
Five cephalopods, all belonging to different species, were identified from deep-sea trawl samples conducted during the DIVA 1-expedition of RV “Meteor” in the Angola Basin in July 2000. These were the teuthoid squids Bathyteuthis abyssicola, Brachioteuthis riisei, Mastigoteuthis atlantica, Galiteuthis armata, and the finned deep-sea octopus Grimpoteuthis wuelkeri. The present study contributes information on size, morphometry, biology and distribution of the species form this unique cephalopod collection.
Abyssal benthic Syllidae (Annelida: Polychaeta) from the Angola BasinDownload PDFAbstract
Three species of Syllidae were obtained from the deep sea expedition DIVA-1 to the south-eastern sector of the Atlantic Ocean. Anguillosyllis capensis Day, 1963 has previously been recorded only from the Agulhas Bank, Parexogone wolfi San Martín, 1991 is also known from the northern Atlantic Ocean and the Gulf of México, and Typosyllis sp. might be a new species.
Three new species of carnivorous deep-sea sponges from the DIVA-1 expedition in the Angola Basin (South Atlantic)Download PDFAbstract
DIVA 1 is the first in a series of expeditions dedicated to the study of benthic diversity in the deep sea of the Atlantic Ocean. On RV “Meteor” we collected samples along a transect of about 700km in the Angola Basin. The collection of Porifera from this expedition is composed of 50 specimens or in some cases pieces of specimens collected at depths between 5162 and 5460m. This work concerns only carnivorous sponges of which we collected 16 specimens belonging to four species. Here we describe three new species: Chondrocladia levii sp. nov., C. vaceleti sp. nov., and C. nicolae sp. nov.; we also found another species, C. cf. guiteli Topsent, 1904 recorded for the first time from a very far abyssal basin (NW of the Galician coast). The main distinctive features of the three new species are the anchorate isochelae categories, the number of teeth on each, and the presence/absence of sigmas; C. levii has three unguiferous isochelae classes with 5, 5, and 9–10 teeth, respectively; C. vaceleti has two unguiferous isochelae classes with three teeth each; C. nicolae has only one unguiferous isochelae category with three teeth. The habitus of C. nicolae (orange in colour) is also very different from that of most of the known Chondrocladia species.
A new taxon of Idyanthidae (Copepoda, Harpacticoida) from the deep sea of the Angola BasinDownload PDFAbstract
Multicorer samples from the deep sea of the Angola Basin yielded 19 species of Idyanthidae Lang, 1994 belonging to five genera. The taxon membership of the species was determined following (Phylogeny of Harpacticoida (Copepoda): Revision of “Maxilliped-asphalea” and Exanechentera. Cuvillier Verlag, Göttingen, 259pp.). Except for one species, Styracothorax gladiator Huys, 1993, all species are new to science. One of them is described here on the basis of females and males. It represents a new genus, Nematovorax gen. nov. One female of the new species N. gebkelinae, was found to hold a nematode between its mouthparts.
Received: 03 March 2004 / Accepted: 21 April 2021 / Published online: 03 March 2004
Comparison of the serotonergic nervous system among Tunicata: implications for its evolution within ChordataDownload PDFAbstract
Using immunohistochemistry in combination with confocal laser scanning microscopy, the serotonergic nervous systems of major tunicate taxa were studied in three-dimensional detail. Organisms analyzed included aplousobranchiate, phlebobranchiate, and stolidobranchiate ascidian larvae, appendicularian juveniles and adults, and doliolid oozooids. Outgroup comparisons to notochordates showed that the serotonergic nervous system of the last common ancestor of Chordata consisted of two elements: (i) an anterior concentration of serotonergic cell bodies, and (ii) a fiber network that followed posteriorly and gave rise to fiber tracts that descended towards the effective somatic lateral musculature. Within Tunicata, the nervous systems of Appendicularia and Aplousobranchiata appear serotonin-reduced or negative. This could be interpreted as a heterochronic reduction and a synapomorphy between Appendicularia and Aplousobranchiata. In this hypothesis, free-living Appendicularia are derived within Tunicata, and a biphasic life cycle with a free-swimming larva and a sessile, ascidian-like adult is most parsimoniously reconstructed for the last common ancestor of Tunicata. The close spatial relation of the serotonergic cell cluster with the statocyte complex suggests a function as an integrative control center for the coordination of locomotion. A similar anterior concentration of serotonergic nerve cells is known from tornaria larvae.See also Electronic Supplement at: http://www.senckenberg.de/odes/05-02.htm
A method for comparing within-core alpha diversity values from repeated multicorer samplings, shown for abyssal Harpacticoida (Crustacea: Copepoda) from the Angola BasinDownload PDFAbstract
A methodology for comparing repeatedly sampled multicorer stations as to significant differences in alpha diversity of selected cores is presented. This is demonstrated for Harpacticoida of the Angola Basin which were sampled during the DIVA-1 campaign of RV “Meteor” in the year 2000 (M48/1).Two replicatedly sampled multicorer stations were compared as to their species-level alpha diversity values of all adult Harpacticoida in single cores. This was done by a newly developed procedure: based on a rank-ordered alpha diversity matrix, using each a species richness, evenness, and dominance diversity index, a minimum spanning tree test (MST-test) was performed to test for significant diversity differences between the replicates of stations 325 (depth: 5448m) and 346 (depth: 5389m). The Canberra Metric was used as a measure of dissimilarity between multicorer deployments. With this procedure, any choice of combination of diversity indices can be made, according to the desired emphasis on certain aspects of diversity. This freedom of choice, together with the possibilities to test for significant diversity differences and to visualize this test, are desirable features of the presented procedure for diversity comparisons. Testing for diversity differences may be useful in the context of conservational purposes when politicians need clear statements from scientists.Due to sufficient replicates, for the first time a significant diversity difference between two abyssal (>2000m depth) multicorer stations was detectable. Station 346 (eight replicates) was significantly more diverse in harpacticoid species than station 325 (seven replicates). Regional-scale differences in food availability are assumed to be of importance for the different patterns of diversity at stations 325 and 346.The slope of the line of regression in a species/individuals plot on single-core level was not far from 1 (R2=0.990;y=0.877x), indicating that most species were represented by singletons and the rest only by very few specimens.The data supported scale-dependent differences of harpacticoid diversity in the Angola Basin. Local-scale (between replicates) differences in harpacticoid within-core species diversity were lower than regional-scale (between stations) differences.At least 134 species of Harpacticoida were found at the two stations, of which the subgroups of Pseudotachidiidae, Argestidae, Ameiridae, and Ectinosomatidae turned out to be richest in species and individuals.
Received: 12 February 2004 / Accepted: 21 April 2021 / Published online: 12 February 2004
On the taxonomic status of the Antarctic amphipod crustacean genera Eclysis (Astyridae) and Bathypanoploea (Stilipedidae), with partial redescription of their type species and description of Bathypanoploea polarsterni n. sp.Download PDFAbstract
The monotypic genus Eclysis K.H. Barnard, 1932 and its type species, E. similis Barnard, are redescribed based upon a newly discovered second specimen. The genus Bathypanoploea Schellenberg, 1939 is reviewed; B. schellenbergi Holman and Watling, 1983 is fixed as the type species. B. polarsterni n. sp. is described; Alexandrella pulchra Ren in Ren and Huang, 1991 is a new junior synonym of B. schellenbergi. The morphology of Eclysis and Bathypanoploea is examined, as well as their relationships to the Astyridae and Stilipedidae.See also Electronic Supplement at: http://www.senckenberg.de/odes/05-03.htm
Received: 25 April 2004 / Accepted: 21 April 2021 / Published online: 25 April 2004
Lungs and mesopneumonia of scincomorph lizards (Reptilia: Squamata)Download PDFAbstract
Mesopneumonia of 28 and lungs of 44 species of scincomorph lizards are described, representing mesopneumonia in six of the seven scincomorph families and lungs of all seven families of this taxon. Except for gymnophthalmids and scincids, a family typical organization of mesopneumonia occurs. In cordylids, gerrhosaurids, xantusiids and lacertids the complete right ventral mesopneumonium (VMp) inserts cranially on the posterior vena cava and caudally on the dorsal surface of the right liver lobe. The left VMp is attached to the ventrolateral body wall in cordylids and lacertids; in gerrhosaurids the left VMp is short and inserts on the pericardium and the posterior vena cava; in xantusiids the left VMp is reduced and the lungs are fused cranially with the pericardium. In scincids the VMp of both sides vary in length and insert on the ventral mesentery, or may be lacking completely. The visceral topology of the gymnophthalmids Calyptommatus and Notobachia differs from the general scincomorph pattern, with the liver and stomach elongated and situated on the right and left side of the body cavity, respectively. The left and the right VMp extend over the entire length of the lungs and insert on the ventral mesentery. All lungs examined in the study are single-chambered and show no major structural variability of the inner surfaces, except in lacertids, some gerrhosaurids, Cordylus, and two genera of gymnophthalmids (Echinosaura and Neusticurus). In these groups, rows of dorsomedial niches are present.See also Electronic Supplement at: http://www.senckenberg.de/odes/05-06.htm
Giant Higgins-larvae with paedogenetic reproduction from the deep sea of the Angola Basin – evidence for a new life cycle and for abyssal gigantism in Loricifera?Download PDFAbstract
A new genus and species of Loricifera, Titaniloricus inexpectatovus (Pliciloricidae) represented by a new type of Higgins-larva is described from the deep sea of the Angola Basin. The new larva is characterized by its gigantic size which is unusual for larval Loricifera, by six rows of scalids on the unit of introvert and neck, by an additional transversal row of scales marking the anterior rim of the collar, by a high number of thoracic plates, by more than 160 longitudinal ridges forming plicae on the loricate abdomen, by tubular toes with a broad basal part and undulated cuticle, and by two pairs of long anterior setae. The exuvium of the presumable sixth instar Higgins-larva of T. inexpectatovus gen. et sp. n. functions as a shelter for several instars: a large simplified seventh instar larva which is paedogenetic and contains some rests of smaller Higgins-larvae, and rests of adult specimens. The paedogenetic or seventh instar larva is morphologically not identical with the sixth instar Higgins-larva from which it moults because of the following transformations: all body regions form a sack-like trunk on which only the scalids of introvert and neck persist as small protoscalids.One of the smaller Higgins-larvae inside contains an adult ready to leave it. The large sixth instar Higgins-larva and the simplified seventh instar paedogenetic larva it contains seem to function together as a “mother larva” for a new larval generation, which has the ability to moult directly into a postlarva and then into an adult. These adults have a size not different from that of other adults of Pliciloricidae. It remains unclear whether these instars represent a new life cycle or just an additional phase in a more complex life cycle such as is often found in other taxa of Pliciloricidae especially from the deep sea. An adult specimen from the same locality as the giant larva is briefly described and assigned with caution to the same species T. inexpectatovus sp. n. The larval gigantism of a deep-sea loriciferan probably reflects adaptations to the deep-sea habitats they inhabit.
A new deep-sea isopod species from the Angola Basin: Prochelator angolensis sp. nov. (Asellota: Desmosomatidae)Download PDFAbstract
A new isopod of the genus Prochelator (Desmosomatidae) is described from the abyssal plain of the South East Atlantic Ocean. This is the first record of Prochelator in the Angola Basin. The new species differs from all other species particularly in antennular setation, the shape of pereonite four and operculum, and the presence of very small posterolateral spines. Furthermore, the species Desmosoma serratum is transferred to the genus Prochelator.
Received: 27 April 2004 / Accepted: 21 April 2021 / Published online: 27 April 2004
Fridericia larix sp. nov. (Enchytraeidae, Oligochaeta) from Irish soilsDownload PDFAbstract
A new species of Fridericia (Enchytraeidae, Oligochaeta) is described from soils in Ireland. It was found during sampling campaigns in the framework of a comprehensive taxonomic revision of the genus (Schmelz, 2003. Taxonomy of Fridericia (Oligochaeta, Enchytraeidae). Revision of species with morphological and biochemical methods. Abh. Naturw. Ver. Hamburg, N.F. 38, 1–415, figs. 1–73), but it was not included in that study. Fridericia larix sp. nov., named in reference to the type locality, belongs to the large and taxonomically difficult group of species with two diverticula per spermatheca. It is distinguished from all known congeners by the following combination of characters: (1) a maximum of four chaetae in ventral preclitellar bundles; (2) oesophageal appendages poorly branched; (3) no pharyngeal glands in segment VII; (4) coelomocytes without refractile vesicles; (5) clitellum girdle-shaped, cell distribution alike on all sides; (6) bursal slit of male copulatory organ mainly transverse; (7) no subneural glands; (8) spermathecal diverticula not stalked. Further distinguishing characters are: (9) an asymmetrical arrangement of chaetae in the first lateral postclitellar bundles, with one large chaeta and one small chaeta per bundle (Fig. 1b, “lc XVI”); (10) the length ratio of spermatozoa to spermatozoal nuclei (6:1–7:1); and (11) a wavy inner surface in parts of the epithelium of the spermathecal ampulla (Fig. 1i, arrow).See also Electronic Supplement at: http://www.senckenberg.de/odes/05-05.htm
Cumacea species (Crustacea: Peracarida) from the deep-sea expedition DIVA-1 with RV “Meteor” to the Angola Basin in July 2000. Family NannastacidaeDownload PDFAbstract
Seven new deep-sea species of the family Nannastacidae from the Angola Basin were observed. In total three species, one in open nomenclature, of the genus Campylaspis Sars, 1865, and one Cumella Sars, 1865, two Styloptocuma Bacescu and Muradian, 1974, and one Platycuma Calman, 1905, are described.
Melinnopsis angolensis (Annelida: Polychaeta: Ampharetidae), a new species from the Angola BasinDownload PDFAbstract
A new species of ampharetid polychaetes, Melinnopsis angolensis sp. nov. is described. It was collected with an Agassiz trawl at seven stations in the Angola Basin in depths between 5385 and 5448m. The new species differs from the few other species described in this genus by the number of thoracic setigers, the number of enlarged tentacles, presence of reduced notosetae in the anterior thorax, and morphological details of the gills and postbranchial membrane. A brief overview of the other species of the genus is given, and the synonymization of Amelinna and Melinnopsides with Melinnopsis is discussed.
Received: 26 January 2004 / Accepted: 21 April 2021 / Published online: 26 January 2004
Evolution of Suessenguthia (Acanthaceae) inferred from morphology, AFLP data, and ITS rDNA sequencesDownload PDFAbstract
The phylogeny and evolution of Suessenguthia (Acanthaceae), a genus of six species from the Andean foothills and adjacent Amazonia in Bolivia, Peru and western Brazil, are discussed based on morphological and molecular (amplified fragment length polymorphism, ITS rDNA) data. Suessenguthia forms a paraphyletic group at the base of the larger genus Sanchezia. The non-overlapping geographical distribution of closely related species suggests that parapatric or allopatric speciation is the major mode in the genus. A major evolutionary tendency promoting diversification of the group presumably was a change from bee- to hummingbird pollination, resulting in a successive adaptation of flower morphology and inflorescence structure.
Received: 13 May 2004 / Accepted: 21 April 2021 / Published online: 13 May 2004
Are Polyommatus hispana and Polyommatus slovacus bivoltine Polyommatus coridon (Lepidoptera: Lycaenidae)? The discriminatory value of genetics in taxonomyDownload PDFAbstract
Taxonomic confusion exists in several sibling species groups. The Polyommatus coridon species complex (Chalk Hill Blues) serves as a model group of sibling species in which genetic analyses provide suitable means for taxonomic clarification. We studied the allozyme patterns of the two described bivoltine species of this complex, Polyommatus hispana and Polyommatus slovacus, and compared them to the two genetic lineages of the univoltine P. coridon. P. hispana is well distinguished from P. coridon (genetic distance: 0.081), and most probably is a sibling species that has evolved during glacial isolation on the Iberian Peninsula. P. slovacus is genetically indistinguishable from the eastern, Pontic–Mediterranean lineage of P. coridon; therefore we suggest that it represents a local bivoltine population only. Since the spring generation of P. slovacus was much less common than the summer generation and showed less genetic diversity, it is probable that uni-/bivoltinism is a dimorphism affecting only part of the whole population. We suggest that the higher genetic diversity of the second generation may be a consequence of gene flow from adjacent single-brooded populations.
First deep-sea record of the genus Kliopsyllus Kunz, 1962 (Copepoda: Harpacticoida) with the description of Kliopsyllus diva sp. n. – the most abundant member of Paramesochridae at two different sites of the Angola BasinDownload PDFAbstract
Two of the sites sampled during the DIVA-1 cruise (Meteor 48/1) to the Angola Basin in 2000 have been studied intensively in order to highlight the diversity of meiofauna in this deep-sea region. This work contributes to the final aim to describe the species of the community of harpacticoid copepods in the Angola Basin and adds a new member to the few hitherto known deep-sea Paramesochridae. The new species belongs to the genus Kliopsyllus, until now only reported from beaches, the intertidal and shallow waters and thus thought to be representing a mainly interstitial group. Furthermore, Kliopsyllus diva sp. n. is the dominating paramesochrid copepod at the two studied sites with depths of more than 5400m. Among other features, K. diva sp. n. is unique in the armature of its fourth leg, the presence of hedgehog-like arranged spinules on the exopod of the fifth leg and the relation of length and width of its furcal rami.
Received: 12 May 2004 / Accepted: 21 April 2021 / Published online: 12 May 2004
Systematics of the Damon variegatus group of African whip spiders (Chelicerata: Amblypygi): Evidence from behaviour, morphology and DNADownload PDFAbstract
The African whip spider, Damon variegatus, exhibits a broad, discontinuous distribution from the Congo, through western Tanzania and Zimbabwe, to South Africa and Namibia. Variation in size, number of antenniform leg segments, and colouration, taken together with a discontinuous distribution, suggest that allopatric populations of D. variegatus may be reproductively isolated, and more than one species may be involved. Furthermore, many morphological characters of D. variegatus appear to be plesiomorphic if compared to closely related species, suggesting that D. variegatus might be paraphyletic, regardless of whether it is a single panmictic species or a group of partly or entirely reproductively isolated populations. This contribution attempts to determine whether D. variegatus is monophyletic and comprises more than one species, by investigating three sources of evidence: behaviour, morphology and DNA. Mating behaviour is observed and mate-recognition trials conducted between males and females from several populations of D. variegatus and related species of Damon. The morphology of spermatophores obtained during these matings is studied and a matrix of somatic and genitalic characters produced. These morphological data are analysed separately and in combination with DNA sequences from loci of three genes in the nuclear genome (18S rDNA, 28S rDNA and Histone H3) and three genes in the mitochondrial genome (12S rDNA, 16S rDNA and Cytochrome Oxidase I). Neither the comparative behavioural evidence gathered nor the spermatophore morphology conclusively suggest that D. variegatus comprises more than one species. However, the molecular data, analysed separately and in combination with the morphological data, reveal that D. variegatus is monophyletic and that the population of D. variegatus to the west of the Kalahari sand system (Namibia and southern Angola) is specifically distinct from those to the east. This new species is described as Damon sylviae, the diagnosis of D. variegatus s. str. is revised, and a key to the species of the D. variegatus group is provided.
Received: 22 April 2004 / Accepted: 21 April 2021 / Published online: 22 April 2004
Phylogeny and biogeography of Valerianaceae (Dipsacales) with special reference to the South American valeriansDownload PDFAbstract
Species of Valerianaceae are a common component of the alpine flora throughout the Northern Hemisphere as well as the Andes of South America. Sequence data from three chloroplast markers (psbA-trnH intron, trnK-matK intron, and the trnL-F region) along with the internal transcribed spacer region (ITS) of nuclear ribosomal DNA were used to infer relationships within Valerianaceae. Both genomes, as well as a combined data set, provide support for the major clades within the group and do not support a monophyletic Valeriana. In addition, these data indicate that Plectritis is nested within South American Valeriana, as opposed to being sister to Centhranthus as previously hypothesized. Valerianaceae appear to have originated in Asia, probably in the Himalayas, and subsequently to have dispersed several times to Europe and to the New World. Our results imply that Valerianaceae colonized South America on multiple occasions from the north. In one of these cases there appears to have been a substantial and rapid radiation, primarily in the high elevation paramo habitat. A variety of methods were used to estimate divergence times to determine when Valerianaceae might have colonized South America. Regardless of the method and fossil constraints applied, our estimates suggest that Valerianaceae colonized South America prior to the formation of the Isthmus of Panama.
Received: 17 June 2005 / Accepted: 21 April 2021 / Published online: 17 June 2005
Bracca olafhenkeli sp. nov., a new species of the Ennominae (Lepidoptera: Geometridae) from Sulawesi (Indonesia)Download PDFAbstract
Bracca olafhenkeli sp. nov. is described from the island of Sulawesi, Indonesia, to which it is considered endemic. Morphological characters such as the basic pattern and coloration as well as the genitalia clearly indicate that the species is a typical member of the genus Bracca Hübner, but large, white apical spots on the forewings and extremely broad, white marginal bands on the hindwings render it unmistakable among its congeners. The new species is abundant in montane areas of North, Central and South Sulawesi and seems to be restricted to primary forests. The biology is unknown.
Received: 11 May 2004 / Accepted: 21 April 2021 / Published online: 11 May 2004
Evolution of the New Zealand mountain flora: Origins, diversification and dispersalDownload PDFAbstract
The New Zealand mountains provide a unique system in which to study the evolution of alpine plants. The relationship between the recent uplift of mountain habitats (5–2 million year ago (mya)) and floristic diversity has polarized hypotheses on the evolution of the alpine flora; suggestions have ranged from an ancient history in New Zealand to recent arrival by long distance dispersal from the Northern Hemisphere. Molecular phylogenies are now available for numerous New Zealand alpine plant groups and these provide insights into the evolution of this unique flora. Taken together with the fossil record, these studies suggest that many alpine lineages first arrived in New Zealand during the late Tertiary and subsequent radiations accompanied environmental upheaval in the Pliocene and Pleistocene. Ongoing studies are investigating the processes that contribute to morphological and ecological diversity in the New Zealand alpine flora.
Received: 04 August 2005 / Accepted: 21 April 2021 / Published online: 04 August 2005
Ehrendorferiana, a new genus of Neotropical jumping plant lice (Insecta: Hemiptera: Psylloidea) associated with conifers (Cupressaceae)Download PDFAbstract
Ehrendorferiana and its two included new species, E. austrocedri (the type species) and E. fitzroyae, are diagnosed. The new genus is a member of the predominantly temperate Neotropical subfamily Aphalaroidinae (Psyllidae). Based on the absence of metabasitarsal spurs it forms a likely monophyletic group with Baccharopelma, Panisopelma and Russelliana. The absence or paucity of long dorsal or marginal setae on the wing pads and caudal plate suggests a probable sister-taxon relationship with Russelliana. Apart from two species of Triozidae from New Zealand on Podocarpaceae, Ehrendorferiana is unique within Psylloidea in its host association with conifers, viz. Austrocedrus and Fitzroya (both Cupressaceae).See also Electronic Supplement at: http://www.senckenberg.de/odes/05-12.htm
New Cumacea species (Crustacea: Peracarida) from the deep-sea expedition DIVA-1 with RV “Meteor” to the Angola Basin in July 2000. Families Lampropidae, BodotriidaeDownload PDFAbstract
Six new deep-sea species from the Angola Basin are described: the family Lampropidae with two species of the genus Hemilamprops Sars, 1883, and one Paralamprops Sars, 1887; the family Bodotriidae with one new species each of the genera Atlantocuma Bacescu and Muradian, 1974, Bathycuma Hansen, 1895, and Cyclaspoides Bonnier, 1896.
Received: 29 July 2004 / Accepted: 21 April 2021 / Published online: 29 July 2004
Phylogenetic analysis of genera of ‘Akanthophoreinae’ (Crustacea: Tanaidacea)Download PDFAbstract
Phylogenetic relations among tanaidacean genera within ‘Akanthophoreinae’ are addressed using computer-assisted parsimony methods. The morphology-based analysis includes 10 well-defined and described genera: Araphura, Chauliopleona, Collettea, Paragathotanais, Metagathotanais, Paraleptognathia, Paranarthrura, Portaratrum, and Tanaella in Tanaidomorpha, and Glabroapseudes in Apseudomorpha as the outgroup. Chauliopleona and Paraleptognathia form a monophylum; Portaratrum cannot be placed in any known family. These three genera are considered as incertae sedis. The analysis does not support the monophyly of ‘Akanthophoreinae’, and further questions the monophyly of Tanaellidae.
Received: 04 October 2004 / Accepted: 21 April 2021 / Published online: 04 October 2004
Molecular evidence that phoronids are a subtaxon of brachiopods (Brachiopoda: Phoronata) and that genetic divergence of metazoan phyla began long before the early CambrianDownload PDFAbstract
Concatenated SSU (18S) and partial LSU (28S) sequences (∼2kb) from 12 ingroup taxa, comprising 2 phoronids, 2 members of each of the craniid, discinid, and lingulid inarticulate brachiopod lineages, and 4 rhynchonellate, articulate brachiopods (2 rhynchonellides, 1 terebratulide and 1 terebratellide) were aligned with homologous sequences from 6 protostome, deuterostome and sponge outgroups (3964 sites). Regions of potentially ambiguous alignment were removed, and the resulting data (3275 sites, of which 377 were parsimony-informative and 635 variable) were analysed by parsimony, and by maximum and Bayesian likelihood using objectively selected models. There was no base composition heterogeneity. Relative rate tests led to the exclusion (from most analyses) of the more distant outgroups, with retention of the closer pectinid and polyplacophoran (chiton). Parsimony and likelihood bootstrap and Bayesian clade support values were generally high, but only likelihood analyses recovered all brachiopod indicator clades designated a priori. All analyses confirmed the monophyly of (brachiopods+phoronids) and identified phoronids as the sister-group of the three inarticulate brachiopod lineages. Consequently, a revised Linnean classification is proposed in which the subphylum Linguliformea comprises three classes: Lingulata, ‘Phoronata’ (the phoronids), and ‘Craniata’ (the current subphylum Craniiformea). Divergence times of all nodes were estimated by regression from node depths in non-parametrically rate-smoothed and other chronograms, calibrated against palaeontological data, with probable errors not less than 50My. Only three predicted brachiopod divergence times disagree with palaeontological ages by more than the probable error, and a reasonable explanation exists for at least two. Pruning long-branched ingroups made scant difference to predicted divergence time estimates. The palaeontological age calibration and the existence of Lower Cambrian fossils of both main brachiopod clades together indicate that initial genetic divergence between brachiopod and molluscan (chiton) lineages occurred well before the Lower Cambrian, suggesting that much divergence between metazoan phyla took place in the Proterozoic.See also Electronic Supplement at: http://www.senckenberg.de/odes/05-11.htm