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Fossil Record and Age of the Asteridae

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Abstract

The Asteridae is a group of some 80,000 species of flowering plants characterized by their fused corollas and iridoid compounds. Recent phylogenetic analyses have helped delimit the group and have identified four main clades within it; Cornales, Ericales, Lamiids and Campanulids, with the last two collectively known as the Euasteridae. A search for the oldest fossils representing asterids yielded a total of 261 records. Each of these fossils was evaluated as to the reliability of its identification. The oldest accepted fossils for each clade were used to estimate minimum ages for the whole of the Asteridae. The results suggest that the Asteridae dates back to at least the Turonian, Late Cretaceous (89.3 mya) and that by the Late Santonian-Early Campanian (83.5 mya) its four main clades were already represented in the fossil record.

Resumen

Las Asteridas son un grupo de unas 80,000 especies de plantas con flor caracterizadas por sus corolas fusionadas y compuestos iridoides. Análisis filogenéticos recientes han ayudado a delimitar al grupo y han identificado cuatro clados principales en él; Cornales, Ericales, Lamiidas y Campanulidas, con las últimas dos conocidas colectivamente como Euasteridas. Una búsqueda por los fósiles más antiguos que representan asteridas produjo un total de 261 registros. Cada uno de estos fósiles fue evaluado en cuanto a la confiabilidad de su identificación. Los fósiles aceptados más antiguos de cada clado se usaron para estimar edades mínimas para las Asteridas. Los resultados sugieren que las Asteridas datan al menos del Turoniano, Cretácico Tardío (89.3 ma) y que para el Santoniano Tardío-Campaniano Temprano (83.5 ma) sus cuatro clados principales ya estaban representados en el registro fósil.

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Acknowledgments

The author would like to thank the following colleagues for helping locate type specimens and for providing access to the collections in their institutions: Magali Volpes (AIX, Aix-en-Provence), Dr. Dario De Franceschi, Dr. Jean Dejax and Mme. Christiane Gallet (MNHN, Paris), Dr. Paul Kenrick, Dr. Peta Hayes and Mr. Cedric Shutte (NHM, London), and Dr. Dena Smith and Dr. Jonathan Marcot (CU Museum, Boulder CO). Special thanks to the librarians at the Hortorium, Mann, Kroch and Engineering Libraries, the Library Annex and Interlibrary Loan for providing prompt access to obscure materials that would have been otherwise impossible to obtain. The author would also like to thank Dr. William Crepet for comments and suggestions that greatly improved this paper, and Dr. Steven Manchester for this thorough review of the manuscript. This research was supported by grants from the following sources: Sigma Xi Cornell Chapter, the Mario Einaudi Center for International Studies, Grants-in-Aid of Research (Sigma Xi), The Paleontological Society, the Mid-America Paleontology Society (MAPS), the American Society of Plant Taxonomists (ASPT), the Botanical Society of America (BSA), the Harold E. Moore, Jr Memorial and Endowment Funds, the Department of Plant Biology, Cornell University and NSF grant DEB 01-08369.

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  1. L. H. Bailey Hortorium, Department of Plant Biology, Cornell University, 412 Mann Library Building, Ithaca, NY, 14853-4303, USA

    Marcela Martínez-Millán

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Appendices

Appendix A

Circumscription of Asterids

The concept and circumscription of asterids has changed dramatically from the days of classification systems based on morphology only. Table 16 shows three widely known systems and their concept of asterids and other groups that are now considered asterids.

Table 16 Circumscription of Asterids Under Three Classification Systems. Placement by Cronquist and Takhtajan of Families that Today are Considered To Be Asterids are Also Given
Full size table

Appendix B

Phylogenetic Analysis of Silvianthemum Friis, 1990

Although the fossil taxon Silvianthemum suecicum Friis, 1990 was included in a phylogenetic analysis (Backlund, 1996 using the Backlund & Donoghue, 1996 matrix of morphological characters), it is appropriate to revisit this taxon as the aforementioned analysis is not compatible with more recent and robust hypotheses of phylogenetic relationships. For this new analysis, characters 32–60—representing floral morphology, androecium, gynoecium and fruit characters—of the Backlund and Donoghue (1996) matrix of morphological characters were used. Silvianthemum was coded based on its original description (Friis, 1990). The resulting matrix has 59 taxa, including the fossil taxon and 29 characters (Table 17):

Table 17 Characters 32 to 60 of the Backlund and Donoghue (1996) Matrix of Morphological Characters with Silvianthemum suecicum Friis, 1990
Full size table
  1. 32.

    Sexual distribution: bisexual = 0; dioecious = 1; gynodioecious = 2; trioecious = 3.

  2. 33.

    Perianth position: hypogynous = 0; semi-epigynous = 1; epigynous = 2.

  3. 34.

    Flower/corolla orientation: one petal adaxial = 0; one petal abaxial = 1.

  4. 35.

    Sepal size: absent or not visible = 0; very reduced, inrolled plumes or minute teeth = 1; well developed prominent = 2.

  5. 36.

    Sepals, number of: 2 = 0; 3 = 1; 4 = 2; 5 = 3; 6 or more, indefinite = 4.

  6. 37.

    Sepal vascularization: 1 trace = 0; 3 traces = 1; 4 traces = 2; 5 traces = 3.

  7. 38.

    Sepal modification for fruit dispersal: none = 0; developing into a plumose seed/ fruit = 1; developing to seeds/fruits with awns/bristles = 2; enlarged and leaflike aiming for wind dispersal = 3.

  8. 39.

    Petal and sepal folding-pattern in buds: valvate = 0; imbricate = 1.

  9. 40.

    Petal fusion: fused = 0; free = 1.

  10. 41.

    Petals, number of: three petals/lobes = 0; four petals/lobes = 1; five petals/lobes = 2; six or more petals/lobes = 3; unnamed state = 4 [sic].

  11. 42.

    Corolla tube: petals weakly connate or no tube = 0; tube rotate/small but distinct = 1; tube well developed/long = 2.

  12. 43.

    Corolla symmetry: actinomorphic = 0; weakly zygomorphic = 1; strongly zygomorphic/bilabiate = 2.

  13. 44.

    Corolla nectary type: absent = 0; nectar disk = 1; multicellular hairs = 2; unicellular hairs = 3.

  14. 45.

    Corolla nectary number: 1, or fewer than number of lobes = 0; 1–5, or equal to number of lobes = 1.

  15. 46.

    Corolla vascularization: lacking lateral connections = 0; with lateral connections = 1.

  16. 47.

    Stamen number: 1 = 1; 2 = 2; 3 = 3; 4 = 4; 5 = 5; 6-more, indefinite = 6.

  17. 48.

    Stamen relative length: equal in length = 0; prominently unequal in length = 1; didynamous = 2.

  18. 49.

    Staminal filament indumentum: glabrous = 0; hairy = 1; unnamed state = 2.

  19. 50.

    Filament attachment: free from corolla = 0; weakly fused to corolla = 1; prominently fused to corolla = 2.

  20. 51.

    Staminal modifications: all stamens fertile = 0; sterile staminodia present = 1.

  21. 52.

    Anther attachment: dorsifixed = 0; basifixed = 1; sagittate = 2.

  22. 53.

    Anther orientation at dehiscence: extrorse = 0; introrse = 1.

  23. 54.

    Sporangium number in thecae: 1 = 0; 2 = 1.

  24. 55.

    Carpels, number: 1 = 1; 2 = 2; 3 = 3; 4 = 4; 5 = 5.

  25. 56.

    Carpel abortion: all fertile = 0; one aborted = 1; two adjacent aborted = 2; two adjacent aborted and ovule displaced = 3; two opposite aborted = 4.

  26. 57.

    Sterile loci: none = 0; present but much reduced and visible only as minor openings = 1; normal/prominent in cross sectioning of ovary = 2.

  27. 58.

    Carpel vascularization: free adaxial and abaxial = 0; adaxial bundles only = 1; only free abaxial, adaxial not recessed = 2.

  28. 59.

    Stigma shape: entire and slender = 0; capitate = 1; bilobate = 2; trilobate = 3; pentalobate = 4.

  29. 60.

    Fruit type: capsule loculicidal = 0; capsule septicidal = 1; berry = 2; drupe = 3; cypsela, with persistent remains of calyx = 4; cypsela, lacking remains of calyx = 5; schizocarp = 6.

The analysis of Bremer et al. (2002) showed that the Escalloniaceae is a polyphyletic group and that part of it, namely Quintinia is more closely related to the Dipsacales than to the rest of the family. It also showed that the Columelliaceae and the Bruniaceae do not belong with the Dipsacales and that Polyosma and Tribeles do not belong with the Apiales. These results differ enough from Backlund and Donoghue (1996) that a reevaluation of the position of Silvianthemum is justified.

In this work, the 59 extant taxa used by Backlund and Donoghue (1996) were rearranged in clades following the results of Bremer et al. (2002), Donoghue et al. (2003) and Zhang et al. (2003). The backbone tree and its group membership matrix were constructed in Winclada version 1.00.08 (Nixon, 2002). In order to allow Silvianthemum to “float” free among all possible positions in the cladograms, all its cells were changed to “?”. Similarly, those taxa not included in the Bremer et al. (2002), Donoghue et al. (2003) or Zhang et al. (2003) works—Zabelia, Knautia, Succisa, Belonanthus, Phyllactis and Stangea—were allowed to “float” free inside the Dipsacales clade by changing their scores to “?” in all characters that defined relationships within that clade. All characters were assigned a weight of 50.

The group membership matrix and the matrix of morphological characters matrix were combined in Winclada. A total of ten heuristic searches were run in NONA version 2.0 (Goloboff, 1999) where each run consisted on 1,000 replications of SPR searches on randomly generated initial wagner trees, holding up to ten trees per replication with an additional TBR on the resulting trees (>h10001; rs0; h/10; mult*1000; max*). The resulting trees were combined in Winclada where suboptimal and duplicate trees were eliminated and a strict consensus was calculated.

The analysis yielded 312 trees whose strict consensus was rerooted to match the basal polytomy of the Campanuliid clade showed in Bremer et al. (2002). In this strict consensus (Fig. 16), Silvianthemum and Quintinia are sister groups with dorsifixed anther attachment as synapomorphy (ch 52). The Dipsacales is the sister group of this clade.

Fig. 16
figure 16

Strict consensus of 312 trees (L = 204, CI = 32, RI = 68 each) rerooted to match the basal polytomy of the Campanuliid clade found by Bremer et al. (2002). The consensus shows the position of the fossil Silvianthemum as sister to Quintinia with dorsifixed anther attachment as synapomorphy (ch 52) and the Dipsacales (black circle) as their sister group

Full size image

Appendix C

Phylogenetic Analysis of Scandianthus Friis and Skarby, 1982

In order to test the assignment of the fossil genus Scandianthus to the Vahliaceae in a phylogenetic context, the characters used by the authors in their original description of the fossil taxon were used to create a data matrix. In Friis and Skarby’s (1982) original table, Scandianthus was compared to 28 families then assigned to the Saxifragales. In the table most columns represent character states (absent/present scoring) and not independent characters, here, this situation was addressed by creating multistate unordered characters. The resulting data matrix has 12 characters (Table 18):

Table 18 Data Matrix of Morphological Characters Derived From the Table Presented By Friis and Skarby (1982)
Full size table
  1. 1.

    Flower sex: bisexual = 0; unisexual = 1.

  2. 2.

    Flower part position: epigynous = 0; perigynous = 1; hypogynous = 2.

  3. 3.

    Perianth part connation: floral parts free = 0; floral parts fused = 1.

  4. 4.

    Androecium number: diplostemonous = 0; obdiplostemonous = 1; haplostemonous = 2; numerous stamens = 3.

  5. 5.

    Carpel number: 2 = 0; 3–5(−15) = 1.

  6. 6.

    Apo/Syncarpic gynoecium: apocarpous = 0; syncarpous = 1.

  7. 7.

    Locule number: 1 = 0; 2–5(−15) = 1.

  8. 8.

    Style number in syncarpous gynoecium: 1 = 0; 2 = 1.

  9. 9.

    Capsule as fruit: capsule = 0; other than capsule (berry, drupe or nut) = 1.

  10. 10.

    Placentae pendant: absent = 0; present = 1.

  11. 11.

    Ovule relative number: few = 0; many = 1.

  12. 12.

    Nectary disc: absent = 0; present = 1.

The families used by Friis and Skarby (1982) were then thought to form the saxifragalean complex. The phylogenetic analyses by Soltis et al. (2000) dismembered this “saxifragalean complex” by showing that these families are distantly related. The works of Soltis et al. (2000), Bremer et al. (2002) and APG (2003) were used here to create a fixed backbone compatible with these newer hypotheses of relationships. This backbone tree was constructed in Winclada version 1.00.08 (Nixon, 2002) and a group membership matrix was derived from it. In this matrix (29 taxa, 20 characters) all character states for Scandianthus were changed to “?” and all characters were assigned a weight of 20. The group membership matrix and the morphological characters matrix were combined in Winclada and ten heuristic searches were run in NONA version 2.0 (Goloboff, 1999). Each run consisted on 1,000 replications of SPR searches on randomly generated initial wagner trees, holding up to ten trees per replication with an additional TBR on the resulting trees (>h10001; rs0; h/10; mult*1000; max*). The resulting trees were combined in Winclada where suboptimal and duplicate trees were eliminated and a strict consensus was calculated.

The searches resulted in 32 trees whose strict consensus was rerooted in the node between the Saxifragales and the Rosid-Asterid clade (Fig. 17). The strict consensus shows Scandianthus as sister to Vahliaceae based on both having one locule (ch 7) and pendant placentae (ch 10).

Fig. 17
figure 17

Strict consensus of 32 trees (L = 62, CI = 22, RI = 28 each) rerooted in the node between the Saxifragales and the Rosid-Asterid clade. Scandianthus was resolved as sister to Vahliaceae based the synapomorphies one locule (ch 7) and pendant placentae (ch 10)

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Martínez-Millán, M. Fossil Record and Age of the Asteridae. Bot. Rev. 76, 83–135 (2010). https://doi.org/10.1007/s12229-010-9040-1

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