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Sibling Echinoderm Taxa on Isolated Ordovician Continents: Problem of Center of Origin

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Sibling Echinoderm Taxa on Isolated Ordovician Continents: Problem of Center of Origin Sibling echinoderm taxa on isolated Ordovician continents: Problem of center of origin SERGEY V. ROZHNOV Morphologically similar echinoderm genera are mostly known from the Ordovician of two isolated continents, Baltica and Laurentia, although these landmasses were strictly isolated biogeographically by the Iapetus Ocean and had differ- ent climates during the Early and Middle Ordovician. The morphological characters of most of these echinoderm genera exclude the possibility that Baltic genera evolved from Laurentian forms or vice versa and suggest that their common an- cestral group was from a different region. I proposed to name such genera ‘sibling genera’, or twin genera by analogy with sibling species. Eocrinoid sibling genera are representatives of the cryptocrinitid-rhipidocystid clade. Baltic Paracryptocrinites and Cryptocrinites are sibling genera of Laurentian Columbocystis, Springerocystis and Foerstecystis, Baltic Rhipidocystis is a sibling genus of Laurentian Mandalacystis and Baltic Neorhipidocystis is sibling genus of Laurentian Batherocystis. It is assumed that North American platicystid paracrinoids, cryptocrinid and rhipidocystid eocrinoids evolved from a common ancestral eocrinoid, similar to Paracryptocrinites. Sibling genera are also known among other Ordovician echinoderms, for example, crinoids (hybocrinids), edrioasteroids (edrioblastoids) and rhombiferans. Some genera migrated from Baltica to Laurentia and vice versa. The temperate warm water seas of eastern Gondwana in the northern China-Australian region were a possible biogeographical center of origin, diversifica- tion or distribution of Laurentian and Baltic sibling genera. Biogeographical analysis of Baltic echinoderms shows that the Baltic Region could be viewed as a ‘museum’ or ‘storehouse’ for many of them, they immigrated and survived there for some considerable time, rather then a ‘cradle’, where they arose and from where they migrated to other continents. • Key words: Ordovician, echinoderms, sibling taxa, biogeography, Baltica, Laurentia, Gondwana. ROZHNOV, S.V. 2010. Sibling echinoderm taxa on isolated Ordovician continents: Problem of center of origin. Bulletin of Geosciences 85(4), 671–678 (4 figures). Czech Geological Survey, Prague. ISSN 1214-1119. Manuscript received February 1, 2010; accepted in revised form September 6, 2010; published online November 30, 2010; issued December 20, 2010. Sergey V. Rozhnov, Borissiak Paleontological Institute RAS, Profsoyuznaya str. 123, Moscow, 117997, Russia; [email protected] Biogeographical analysis of the distribution of taxa is parti- rally valid patterns of macroevolution. The present paper cularly important at the point of significant evolutionary develops an approach which takes into account probable radiation as it provides a more reliable foundation for one biogeographical links between the epeiric seas of separated or another phylogenetic reconstruction based on morpho- continents. logical data. The Ordovician evolutionary radiation during which many metazoan classes, including Recent taxa, ap- peared, is of particular interest in this respect (review in Geographical, climatic Webby 2004). Echinoderms compose one of the most im- and sedimentological features portant benthic groups of Ordovician invertebrates (Rozh- of Ordovician continents nov 2002, 2007a; Sprinkle & Guensburg 2004). In the Or- dovician, ten echinoderm classes originated, including all In the Ordovician, there were four large isolated continents the present day retained forms, i.e., crinoids, sea urchins, (Fig. 1) positioned relatively far from each other: Baltica, starfishes, ophiuroids and, probably, holothurians (Rozh- Laurentia, Siberia and Gondwana (Cocks & Torsvik 2002). nov 2002). The relationships between these classes and Siberia was close to the equator in the tropical climatic the higher taxa containing them, centers on their origin, fea- zone (Cocks & Torsvik 2007). The echinoderm fauna of tures of geographical distribution and development in the the Ordovician seas of this continent has not been studied epeiric seas of separated continents. They are important in detail; however, it was probably relatively poor. Other- problems in the study of echinoderms which display gene- wise, they would be well represented in collections of DOI 10.3140/bull.geosci.1181 671 Bulletin of Geosciences Vol. 85, 4, 2010 Figure 1. Early Ordovician geography (after Cocks & Torswik 2002). Large ar- rows show possible migration routes of sibling echinoderm genera from East Gondwana to West Gondwana and Laurentia. Small arrows show possible oceanic currents based on the data from Christiansen & Stouge (1999) with addi- tions. marine invertebrates from hard-to-reach areas of Ordovi- simultaneously with the beginning of carbonate sedimenta- cian deposits of this continent, collected by the many stra- tion and rapidly became a dominant group in many benthic tigraphers and paleontologists by the middle of the 20th communities (Rozhnov 2007a). Echinoderm faunas of century. Gondwana was the largest continent, extending these continents differ considerably in composition and in- from the South Pole to tropical latitudes. Cold climatic clude many endemic genera. Nevertheless, they show cer- conditions were characteristic for most of its area in the tain similar features, in particular, morphologically similar Ordovician, although eastern Gondwana extended to the taxa, which are undoubtedly closely related. The center of equator, with a much warmer climate (Cocks & Torsvik origin and distribution of these taxa and their migration 2002). The most thoroughly investigated echinoderm routes are an interesting biogeographical problem, and are fauna comes from the cold marginal areas of West Gond- important for the resolution of phylogenetic problems of wana (Gondwanan Africa and peri-Gondwanan Europe), various taxa, that were prominently manifest during the with mostly terrigenous sedimentation (Lefebvre & Fatka Ordovician evolutionary radiation. 2003). Laurentia was near the equator during the entire Or- dovician. A tropical climate prevailed, with mostly carbo- nate type sedimentation. This continent has yielded the Opportunity of dispersal richest echinoderm fauna (Sprinkle & Guensburg 2004). of Ordovician echinoderms During the Ordovician, Baltica moved from high latitudes towards the equator (Cocks & Torsvik 2005). Consequent- The majority of Ordovician echinoderms were sessile ani- ly, at the beginning of the Middle Ordovician, cold epeiric mals, and migrated only during the larval stage. Free-living seas were replaced by temperate seas; they became warm Ordovician echinoderms migrated mostly by means of from the end of the Middle Ordovician and tropical by the planktonic larvae since the adults were slow moving. Ex- middle of the Upper Ordovician (Dronov & Rozhnov 2007). pansion usually occurred along continental coasts due to Carbonate sedimentation was established and prevailed transportation of larvae by coastal currents, and was rather from the very end of the Early Ordovician, even under rapid, covering all shelves. In some cases, migrations may cold-water conditions. Echinoderms appeared in Baltica have occurred in directions opposite to the prevailing 672 Sergey V. Rozhnov Sibling echinoderm taxa on isolated Ordovician continents currents due to the displacement of larvae during storms having a common ancestor but isolated reproductively, and unusual changes in the prevailing winds. Only consi- Mayr (1942) introduced the term twin (or sibling) species. derable changes in water temperature and salinity preven- These similar species often inhabit different isolated re- ted expansion along a coast. Larvae could be transported gions or biotopes. In paleontology the majority of benthic from one continent to another only by oceanic currents, as species with wide geographical and stratigraphical ranges all Ordovician echinoderms were shallow-water animals may in fact be mixed sibling species in the biological sense, and it was impossible for them to cross oceans in the man- since they were established based on a limited number of ner of a step-by-step migration. Migration of echinoderms characters, as compared with living taxa, and they could from one continent to another required either the opportu- have been isolated reproductively both geographically and nity for larvae to remain in currents for some considerable chronologically. By analogy with sibling species, morpho- time or successive migration via a series of islands and logically similar genera and higher rank taxa, for example, microcontinents. All modern crinoid larvae are lecithotrop- families having a common ancestor at the same taxonomic hic, that is unable to eat independently (Holland 1991, level can be termed sibling taxa. In other words, morpholo- Smith 1997). Therefore, they are unable to endure gically similar genera, with a common ancestral genus, are long-term travel in oceanic currents. Paleozoic crinoids, designated sibling genera. The families having a common like other pelmatozoan echinoderms, were probably consi- ancestral family are sibling families. The presence of sib- derably restricted with reference to migration using ocea- ling taxa on separated continents suggests that they migra- nic currents. Lecithotrophic larvae live only a number of ted from a specific third region where their common ances- days whereas planktotrophic larvae usually live for several tor dwelt (Rozhnov 2007b). At the same time, the ancestral weeks before setting (Ivanova-Kasas
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