Mid-Cretaceous angiosperm radiation and an asterid origin of bilaterality: diverse and extinct "Ericales" from New Jersey.

PREMISE OF THE STUDY Numerous fossils from the Upper Cretaceous have been confidently placed within modern crown groups. Many 95-75 Myr-old taxa, however, including the taxon described herein, do not fit well with known extant crown or stem groups. Understanding such fossils and their possible affinities would certainly enhance our understanding of the circumstances involved in a major eudicot radiation. METHODS Bulk samples from the Old Crossman Clay Pit were prepared using standard methodology, which includes several washing and sieving steps, and a treatment with hydrofluoric acid. The fossil taxon was coded into a matrix built from the combination of two previously published morphological matrices and was analyzed using the parsimony criterion with the computer program TNT. KEY RESULTS The fossils have a unique combination of characters relative to living and fossil Ericales taxa, and therefore, a new genus, Teuschestanthes, is erected. Mosaic evolution and rapid parallel changes in such groups blur taxonomic distinctions, and these issues are exacerbated by limited numbers of characters available in fossils. Teuschestanthes flowers are slightly bilaterally symmetrical and somewhat variable with regard to petal disposition, suggesting an early stage in transition to bilaterality from radial symmetry early in eudicot history under pollinator selective pressure. CONCLUSIONS While Teuschestanthes shares characters with modern Ericales and Sapindales, there are significant non-overlapping differences between Teuschestanthes and modern Sapindales (notably, among others, ovule number). Based on available evidence, however, the position of Teuschestanthes is likely as an early offshoot of the stem clade of core Ericales (Ericales sensu stricto). Its relatively unstable floral plan may presage subsequent bilaterality associated with growing selective pressure by advanced pollinators.

[1]  M. Gandolfo,et al.  A late Cretaceous fagalean inflorescence preserved in amber from New Jersey. , 2018, American journal of botany.

[2]  P. Stevens,et al.  Evolution of the process underlying floral zygomorphy development in pentapetalous angiosperms. , 2017, American journal of botany.

[3]  M. Gandolfo,et al.  A new species of Athrotaxites (Athrotaxoideae, cupressaceae) from the upper cretaceous raritan formation, New Jersey, USA , 2016 .

[4]  K. Nixon,et al.  Rariglanda jerseyensis, a new ericalean fossil flower from the Late Cretaceous of New Jersey1 , 2016 .

[5]  J. Lundberg,et al.  An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants : APG II THE ANGIOSPERM PHYLOGENY GROUP * , 2003 .

[6]  K. Nixon,et al.  Fossil Ericales from the Upper Cretaceous of New Jersey , 2013, International Journal of Plant Sciences.

[7]  Xianghui Xiao,et al.  Glandulocalyx upatoiensis, a fossil flower of Ericales (Actinidiaceae/Clethraceae) from the Late Cretaceous (Santonian) of Georgia, USA. , 2012, Annals of botany.

[8]  Peter R. Crane,et al.  Early Flowers and Angiosperm Evolution , 2011 .

[9]  K. Nixon,et al.  Pentapetalum trifasciculandricus gen. et sp. nov., a thealean fossil flower from the Raritan Formation, New Jersey, USA (Turonian, Late Cretaceous). , 2009, American journal of botany.

[10]  Pablo A. Goloboff,et al.  TNT, a free program for phylogenetic analysis , 2008 .

[11]  B. Danforth,et al.  A Fossil Bee from Early Cretaceous Burmese Amber , 2006, Science.

[12]  M. Gandolfo,et al.  An extinct calycanthoid taxon, Jerseyanthus calycanthoides , from the Late Cretaceous of New Jersey. , 2005, American journal of botany.

[13]  J. Schönenberger,et al.  Molecular Phylogenetics and Patterns of Floral Evolution in the Ericales , 2005, International Journal of Plant Sciences.

[14]  M. Gandolfo,et al.  Cretaceous flowers of Nymphaeaceae and implications for complex insect entrapment pollination mechanisms in early angiosperms. , 2004, Proceedings of the National Academy of Sciences of the United States of America.

[15]  M. Gandolfo,et al.  Divisestylus gen. nov. (aff. Iteaceae), a fossil saxifrage from the Late Cretaceous of New Jersey, USA. , 2003, American journal of botany.

[16]  M. Gandolfo,et al.  Triuridaceae fossil flowers from the Upper Cretaceous of New Jersey. , 2002, American journal of botany.

[17]  M. Källersjö,et al.  Phylogenetic relationships in the order Ericales s.l.: analyses of molecular data from five genes from the plastid and mitochondrial genomes. , 2002, American journal of botany.

[18]  K. Nixon,et al.  The earliest fossil evidence of the Hamamelidaceae: Late Cretaceous (Turonian) inflorescences and fruits of Altingioideae. , 2001, American journal of botany.

[19]  E. M. Friis,et al.  Fossil flowers of ericalean affinity from the Late Cretaceous of Southern Sweden. , 2001, American journal of botany.

[20]  M. Gandolfo,et al.  Turonian Pinaceae of the Raritan Formation, New Jersey , 2001, Plant Systematics and Evolution.

[21]  M. Gandolfo,et al.  Sorophores ofLygodium Sw. (Schizaeaceae) from the Late Cretaceous of New Jersey , 2000, Plant Systematics and Evolution.

[22]  K. Nixon The Parsimony Ratchet, a New Method for Rapid Parsimony Analysis , 1999 .

[23]  K. Nixon,et al.  Two new fossil flowers of magnoliid affinity from the Late Cretaceous of New Jersey. , 1998, American journal of botany.

[24]  K. Nixon,et al.  Fossil Clusiaceae from the late Cretaceous (Turonian) of New Jersey and implications regarding the history of bee pollination. , 1998, American journal of botany.

[25]  M. Gandolfo,et al.  A new fossil flower from the Turonian of New Jersey: Dressiantha bicarpellata gen. et sp. nov. (Capparales). , 1998, American journal of botany.

[26]  M. Gandolfo,et al.  Tylerianthus crossmanensis gen. et sp. nov. (aff. Hydrangeaceae) from the Upper Cretaceous of New Jersey. , 1998, American journal of botany.

[27]  M. Gandolfo,et al.  A new fossil fern assignable to Gleicheniaceae from Late Cretaceous sediments of New Jersey. , 1997, American journal of botany.

[28]  P. Herendeen,et al.  Fossil flowers and fruits of the Actinidiaceae from the Campanian (Late Cretaceous) of Georgia , 1996 .

[29]  W. Crepet Timing in the evolution of derived floral characters: upper cretaceous (turonian) taxa with tricolpate and tricolpate-derived pollen , 1996 .

[30]  K. Nixon,et al.  Fossil flowers and pollen ofLauraceae from the Upper Cretaceous of New Jersey , 1994, Plant Systematics and Evolution.

[31]  A. Anderberg Cladistic interrelationships and major clades of theEricales , 1993, Plant Systematics and Evolution.

[32]  K. Nixon,et al.  Chloranthus-like stamens from the upper Cretaceous of New Jersey , 1993 .

[33]  K. Nixon,et al.  LATE CRETACEOUS FOSSIL FLOWERS OF ERICALEAN AFFINITY , 1993 .

[34]  W. Judd,et al.  Circumscription of Ericaceae (Ericales) as determined by preliminary cladistic analyses based on morphological, anatomical, and embryological features , 1993, Brittonia.

[35]  K. Nixon,et al.  Oldest fossil flowers of hamamelidaceous affinity, from the Late Cretaceous of New Jersey. , 1992, Proceedings of the National Academy of Sciences of the United States of America.

[36]  C. Michener,et al.  The oldest fossil bee: Apoid history, evolutionary stasis, and antiquity of social behavior. , 1988, Proceedings of the National Academy of Sciences of the United States of America.

[37]  E. M. Friis Actinocalyx gen. nov., sympetalous angiosperm flowers from the upper cretaceous of southern Sweden , 1985 .

[38]  E. M. Friis Upper Cretaceous (Senonian) floral structures of juglandalean affinity containing normapolles pollen , 1983 .

[39]  R. Christopher Normapolles and triporate pollen assemblages from the Raritan and Magothy Formations (Upper Cretaceous) of New Jersey , 1979 .

[40]  J. Doyle,et al.  Angiosperm pollen zonation of the continental Cretaceous of the Atlantic coastal plain and its application to deep wells in the Salisbury embayment , 1977 .

[41]  K. Nixon,et al.  Flowers of Turonian Magnoliidae and their implications , 1994 .

[42]  Larry Hufford,et al.  Rosidae and their relationships to other nonmagnoliid dicotyledons : a phylogenetic analysis using morphological and chemical data , 1992 .

[43]  D. Grimaldi,et al.  Occurrence, chemical characteristics, and paleontology of the fossil resins from New Jersey. American Museum novitates ; ; no. 2948. , 1989 .

[44]  R. Christopher The occurrence of the Complexiopollis-Atlantopollis zone (Palynomorphs) in the Eagle Ford Group (Upper Cretaceous) of Texas , 1982 .

[45]  P. Callahan The evolution of insects , 1972 .

[46]  G. J. Brenner The spores and pollen of the Potomac group of Maryland , 1963 .