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Metrarabdotos and Related Genera (Bryozoa: Cheilostomata) in the Late Paleogene and Neogene of Tropical America
- Alan H. Cheetham, Joann Sanner, Jeremy B. C. Jackson
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- Journal:
- Journal of Paleontology / Volume 81 / Issue S67 / January 2007
- Published online by Cambridge University Press:
- 11 August 2017, pp. 1-91
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Metrarabdotos Canu, 1914 and the related genera Escharoides Milne Edwards, 1836b, Adeonellopsis MacGillivray, 1886, and Reptadeonella Busk, 1884 were key taxa in the decline of Bryozoa with erect, arborescent colonies and concomitant increase in numbers of species with encrusting colonies in the late Paleogene and Neogene of tropical America. In particular, the abundance, continuity of occurrence, and diversity of Metrarabdotos before its decline have made this genus, over the past 20 years, a model taxon for detailed morphometric studies of fine-scale evolutionary tempo and mode. During the same period, significant new occurrences of both Metrarabdotos and its near relatives have been documented with detailed collections from tropical American areas not represented in the original studies.
In this paper we present a complete reanalysis of the original morphometric data (Cheetham, 1986a), incorporating the new tropical American material and comparative material of selected Metrarabdotos species from Europe and Africa in order to formalize the taxonomy of the genus, many species of which have been in open nomenclature since 1986, and to explore their possible phylogenetic relationships to each other and to Escharoides, Adeonellopsis, and Reptadeonella. The new analysis, with procedures slightly modified from those used in the original analysis, is based on more than twice the number of specimens and 20% more morphological characters. Although the results include changes in species assignments for 13% of the specimens in the original analysis, the pattern of intraspecific morphological stasis previously identified in the tropical American Neogene species, and thus the concomitant interpretation of evolutionary tempo and mode, are unaltered.
Cladistic analysis resulted in a single most parsimonious tree for the 22 tropical American Metrarabdotos species, arranged in two monophyletic crown groups and a paraphyletic stem group. The stem group, ranging from latest Eocene to late Early or early Middle Miocene in age, includes four species, two of which are new: M. aguilerai from Venezuela and M. hispaniolae from the Dominican Republic and Haiti. Crown group A comprises seven species occurring in deposits of Late Miocene to Recent age, four of which are new: M. arawakorum from Venezuela; M. boldi and M. saundersi from the Dominican Republic; and M. coatesi, which occurs in the Dominican Republic, Costa Rica, and Panama. Crown group B, comprising 11 species, ranges from Early Miocene to Recent and includes five new species: M. cubaguaense from Venezuela; M. vokesorum from the Dominican Republic, Costa Rica, and Panama; and M. tainorum, M. jungi, and M. lopezense from the Dominican Republic. Incorporation of six European and African (eastern Atlantic) species, ranging in age from Early Miocene to Recent, into cladograms with the tropical American species produced more variable results in terms of both numbers of alternative trees and the positions of the tropical American species in them. One of these species, M. thomseni from the Pleistocene of Greece, and possibly a second, M. cf. M. maleckii Cheetham, 1968 from the Miocene of the Czech Republic, are new. With one exception, the eastern Atlantic species were placed in more or less basal positions in one or the other of the crown groups, suggesting that, as hypothesized by Cheetham (1968), the “Old World” and “New World” species of Metrarabdotos represent largely independent radiations. The exception is the living species M. cookae Cheetham, 1968, from West Africa and the Cape Verde Islands, which is placed unequivocally in a terminal position in crown group B. Although the three groups of tropical American species correspond in part to subgenera of Metrarabdotos erected by Cheetham (1968), the paraphyly of the stem group and the unresolved identity of the type species of the subgenus apparently corresponding to crown group A suggest that formalizing the taxonomy of these groups is unwarranted at present.
A further cladistic analysis, incorporating the three groups of Metrarabdotos species and Escharoides, Adeonellopsis, and Reptadeonella, yielded two trees, both of which support Cheetham's (1968) hypothesis that Metrarabdotos is more closely related to Escharoides than to Adeonellopsis and Reptadeonella. Escharoides is represented in tropical American Neogene deposits by two species, one of which, E. guraboensis from the Dominican Republic, is new. Of the six tropical American Neogene species of Adeonellopsis, four are new: A. cribrospiramen and A. guraboensis from the Dominican Republic, A. limonensis from Costa Rica, and A. antilleana from Cuba and Jamaica. Three of the six tropical American Neogene species of Reptadeonella, R. buddae from the Dominican Republic, R. collinsae from the Dominican Republic, Panama, Costa Rica, and the southeastern United States, and R. buricaensis from Panama, are also new.
Evolutionary significance of sexual and asexual modes of propagation in Neogene species of the bryozoan Metrarabdotos in tropical America
- Alan H. Cheetham, Jeremy B. C. Jackson, Joann Sanner
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- Journal:
- Journal of Paleontology / Volume 75 / Issue 3 / May 2001
- Published online by Cambridge University Press:
- 20 May 2016, pp. 564-577
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Three new Miocene-Pliocene species of the cheilostome bryozoan Metrarabdotos from Venezuela are atypical in showing significant evidence that as many as half the colonies originated asexually (clonally) by “regeneration” from previously existing colonies, rather than almost exclusively from ancestrular zooids (products of metamorphosis of sexually produced larvae), as is characteristic of the genus. The extremely low proportion of zooids (less than two percent) recognizably committed to producing larvae (ovicelled) in these Venezuelan species agrees with that reported in a variety of Danian (Paleocene) genera in which clonal propagation has been reported to predominate. However, all but two of 17 other living and fossil species of Metrarabdotos also have fewer than two percent of their zooids ovicelled, even though all but one of more than 250 colony bases examined originated from ancestrulae. The lack of significant correlation in Metrarabdotos between frequencies of ovicelled zooids and of ancestrular colonies suggests that clonal propagation may not have diverted resources from sexual reproduction. This inference is supported by the retention in these species of a level of heritable morphologic variation (estimated by partitioning among-colonies and within-colonies variance in zooid characters) that is commensurate with that estimated for species of Metrarabdotos in which propagation was apparently entirely by sexual means. Thus, sexual reproduction throughout the genus was apparently sufficient to maintain the genetic diversity from which speciation could proceed at normal rates. As estimated by both cladistic and nearest-neighbor morphologic-stratigraphic methods, the three Venezuelan species occupy quite different positions in the inferred phylogeny of Metrarabdotos. Thus, the elevated level of clonal propagation in these species appears to be a response to local conditions, most probably high productivity associated with upwelling, that promoted more rapid vegetative growth while leaving the level of sexual reproduction unchanged.
Neogene Cupuladriidae of tropical America. II: taxonomy of recent Discoporella from opposite sides of the Isthmus of Panama
- Amalia Herrera-Cubilla, Matthew H. Dick, Joann Sanner, Jeremy B. C. Jackson
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- Journal:
- Journal of Paleontology / Volume 82 / Issue 2 / March 2008
- Published online by Cambridge University Press:
- 20 May 2016, pp. 279-298
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We used up to 30 morphological characters to discriminate and describe species of the genus Discoporella based on complete colony specimens collected from both coasts of the Isthmus of Panama. The characters included zooidal characters and colony-level characters such as colony size and basal granule density. Species were classified by a series of multivariate cluster and linear discriminant analyses until the majority of specimens were assigned to their putative species with high confidence. In the first phase of the analyses, the colonies were grouped by ocean (Caribbean versus eastern Pacific), discriminated predominantly by colony size and basal granule density, characters that might reflect ecophenotypic responses to different conditions in primary productivity and predation between the two oceans. Further analyses of these two groups separately resulted in the discrimination of seven species. Five new species from the Caribbean (D. scutella, D. peltifera, D. bocasdeltoroensis, D. terminata and D. triangula), and two from the eastern Pacific (D. marcusorum and D. cookae). Of these, D. cookae had been identified previously as D. umbellata, a species once considered cosmopolitan, with a range spanning the Caribbean and eastern Pacific coasts of America. With the exception of one genetically defined clade represented by only two specimens, the correspondence of classification between groups discriminated morphometrically by separate step-wise multivariate analyses and those detected by a previous genetic analysis, ranged from 91% to 100%. In analyses of all specimens combined or separated by ocean, but using the total number of characters, 20% to 30% of the specimens could not be distinguished morphometrically from extremely similar sympatric species or cognate (“geminate”) species from the opposite ocean. Diversity was higher in the Caribbean compared to the eastern Pacific, which reflects a similar pattern recently described for the genus Cupuladria from the same region.
Morphological differentiation of Avicularia and the proliferation of species in mid-Cretaceous Wilbertopora Cheetham, 1954 (Bryozoa: Cheilostomata)
- Alan H. Cheetham, Joann Sanner, Paul D. Taylor, Andrew N. Ostrovsky
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- Journal of Paleontology / Volume 80 / Issue 1 / January 2006
- Published online by Cambridge University Press:
- 20 May 2016, pp. 49-71
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Discovery of avicularium-like polymorphs in Wilbertopora mutabilis Cheetham, 1954 has provided not only a new opportunity for revising the genus Wilbertopora Cheetham, 1954, but also a more detailed basis for documenting the series of morphological changes by which avicularia differentiated from ordinary feeding zooids in what appears to be the first occurrence of these characteristic cheilostome bryozoan structures in the fossil record.
Eighteen of a total 60 quantitative characters measured on avicularia and ordinary and ovicell-bearing autozooids were sufficient to distinguish eight species of Wilbertopora by discriminant function analysis of zooid data from 93 colonies from the mid-Cretaceous (Albian-Cenomanian) Washita Group in northeastern Texas and southeastern Oklahoma. Eighteen of a total of 20 of the quantitative characters that could be statistically coded for cladistic analysis proved to be informative with respect to parsimony, providing two maximally parsimonious trees for the eight species. Two-thirds of the diagnostic characters involve avicularia. An additional 55 colonies too poorly preserved for morphometric analysis could then be assigned to species qualitatively, with 170 more colonies lacking species-diagnostic characters.
The cladistic trees strongly suggest that most or all of the species diverged before the end of the Albian, but stratigraphic resolution is insufficient to test this hypothesis. Nevertheless, the series of morphological changes differentiating avicularia from ordinary autozooids in these species, based on the cladistic relationships, is highly significant statistically, and may be a pattern later repeated in other cheilostomes.
Wilbertopora and W. mutabilis are emended, and seven new species are described: W. listokinae, W. tappanae, W. spatulifera, W. attenuata, W. improcera, W. acuminata, and W. hoadleyae.
Neogene Cupuladriidae of tropical America. I: Taxonomy of Recent Cupuladria from opposite sides of the Isthmus of Panama
- Amalia Herrera-Cubilla, Matthew H. Dick, Joann Sanner, Jeremy B. C. Jackson
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- Journal:
- Journal of Paleontology / Volume 80 / Issue 2 / March 2006
- Published online by Cambridge University Press:
- 14 July 2015, pp. 245-263
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We used up to 28 morphological characters to discriminate and describe species of the genus Cupuladria based on entire colony specimens collected from both coasts of the Isthmus of Panama. The characters included a combination of zooidal features traditionally used in cheilostome taxonomy and nontraditional characters such as colony size, shape, and an index of calcification of the colony, as well as the size of the basal sectors and their number of pores. Species were discriminated by a series of repeated multivariate cluster and discriminant analyses until the majority of specimens were assigned to their putative species with high statistical confidence. Nontraditional characters contributed significantly to the power of the analyses. Colonies fell into two highly distinct groups most clearly recognized by the presence or absence of vicarious avicularia, which agrees well with previous molecular genetic analyses. Further analyses of each of these two groups considered separately resulted in the discrimination of eight species. These include two previously described Caribbean species, C. biporosa Canu and Bassler, 1919 and C. surinamensis Cadée, 1975, and six new species: C. multesima, C. incognita, C. cheethami, and C. panamensis from the Caribbean, and C. pacificiensis and C. exfragminis from the eastern Pacific. There was also good correspondence between major clades within these morphologically defined groups and the previous molecular analysis, although 20% of the specimens could not be distinguished from their cognate (“geminate”) species from the opposite ocean. The high ratio of undescribed to described species and higher diversity in the Caribbean than eastern Pacific agree well with newly described patterns from other cheilostome genera based on similar analyses. Quantitative morphometric studies are essential to study biologically meaningful patterns of cheilostome speciation and macroevolution in the fossil record.