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A Review of the Fossil Record of the Genus Itea (Iteaceae, Saxifragales) with Comments on its Historical Biogeography

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Abstract

Itea is a genus of about 20 species of trees and shrubs that are today native to southeastern North America, eastern Asia, and eastern Africa. In this paper, I review the fossil record of Itea, which is based on four types of fossils: diporate, psilate pollen attributed to Itea or the dispersed pollen genus Iteapollis; carpofossils representing fruits and seeds attributed to Itea europaea; flowers preserved in amber and assigned to Adenanthemum iteoides; and leaf impressions attributed to Itea. The distributions of these fossils indicate that Itea was present in western North America from the early Eocene to Miocene, in eastern North America beginning no later than the early Miocene, and in western Eurasia from the late Eocene to Pliocene. Only one datapoint is known from eastern Asia; it is early Miocene in age. Based on the fossil record, it can be inferred that Itea crossed between continents over both the Bering Land Bridge and North American Land Bridge, and that it reached Africa from Europe via Anatolia. Thus, it is predicted that the sole extant North American species, I. virginica, may be most closely related to the sole extant African species, I. rhamnoides. The potential application of Itea fossils to calibrating phylogenetic trees generated from molecular sequence data is also discussed.

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Acknowledgments

This project includes information from the second chapter of my dissertation (Hermsen, 2005) as well as much new research done since 2005. I would especially like to recognize W.C. Wehr, now deceased, for contributing to my understanding of the putative Itea leaves from the Republic flora. Thank you to the staff members of all facilities which I visited to examine collections or from which I borrowed material; I would especially like to acknowledge L. Barksdale, C. Brown, B.A.R. Mohr, R. Russell, R.A. Stockey, W.C. Wehr, and S. Wing for their assistance in accessing collections and/or providing loans of material. I would like to thank the following libraries and their staffs for assistance with gathering literature for this review: the Cornell University Libraries (Ithaca, NY), the San Jose Public Library (San Jose, CA), the United States Geological Survey Library (Menlo Park, CA), and the Branner Earth Sciences Library at Stanford University (Palo Alto, CA). I would also like to extend my gratitude to those who have provided meaningful discussion pertaining to this research through the years. Special thanks to I. Petrescu for clarifying the Romanian fossil record of Itea, S.R. Manchester for comments on the identity of the leaves from Republic, W.K. Buechler for discussion of leaf clearing techniques, E. Martinetto for providing literature, the NESCent working group "Synthesizing and Databasing Fossil Calibrations" for relevant discussions, and J.R. Hendricks for reading drafts of the manuscript and providing assistance with GIS. Thank you also to those who collected material from the Stonerose Fossil Site in Republic, Washington, particularly L. Barksdale and E. Lester, whose discoveries are figured in this paper. Funds from NSF grant DEB 0206185 and graduate research grants from the Paleontological Society and the American Society of Plant Taxonomists covered travel and other expenses incurred in examining specimens of fossil and extant Itea. Tropicos® was used as a source for extant taxon authorities.

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Authors and Affiliations

  1. Department of Environmental and Plant Biology, Ohio University, Porter Hall 401, Athens, OH, 45701-2979, USA

    Elizabeth J. Hermsen

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  1. Elizabeth J. Hermsen
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Correspondence to Elizabeth J. Hermsen.

Appendices

Appendix 1

List of reported occurrences of fossils assigned to or morphologically consistent with Itea. Fossils are listed by relative age of first occurrence, then alphabetically by locality/flora. Information is given in the following format: Taxon name, age or age range, locality/flora name (primary geographic subdivision—e.g., state/province/region/county, country). Citations (including explanations for any name changes). Additional Occurrence References (“Add. Occ. Refs.,” additional references pertaining to the age, stratigraphy, or geographic occurrence of a given locality/flora that do not mention the presence of Itea fossils are given after the abbreviation). Occurrences mapped in this paper may include one to several separate samples or closely spaced localities.

Flowers

  1. 1.

    Adenanthemum iteoides, late Eocene or older, Baltic Amber (thought to have originated in Fennoscandia). Conwentz (1886). Add. Occ. Refs.: Schlüter (1990), Ritzkowski (1997, 2008), Perkovsky et al. (2007), Kasiński and Kramarska (2008). Neotype of Adenanthemum iteoides here designated: The illustrations of one specimen of a floral inclusion in a piece of amber provided by Conwentz (1886), pl. 9, figs. 16, which is an illustration of the entire pedicellate flower. Additional figures of the same specimen include Conwentz (1886), pl. 9, figs. 15, 17, & 18–24. Conwentz (1886) pl. 9, fig. 25, is a floral diagram for the taxon.

Fruits & Seeds

  1. 1.

    Itea europaea, Miocene, locality unknown (Belarus). Mai (1985), Dorofeev (1988), Pingen (1996). Not mapped.

  2. 2.

    Itea europaea, early Miocene, Berzdorf (Saxony, Germany). Mai (1985, 2000a, b), Czaja (2003). Add. Occ. Ref.: Mai (1999). Holotype: Mai (1985), pl. 6, fig. 1.

  3. 3.

    Itea europaea, early Miocene, Hartau (Saxony, Germany). Mai (1985, 2000a, b). Add. Occ. Ref.: Mai (1999).

  4. 4.

    Itea europaea, early Miocene, Reichwalde (Saxony, Germany). Mai (1985, 2000a, b). Add. Occ. Ref.: Mai (1999).

  5. 5.

    Itea europaea, middle Miocene, Chelnovskih (Tambov, Russia). Dorofeev (1988) as Carpolithus sp. 1, transferred to Itea europaea by Pingen (1996).

  6. 6.

    Itea europaea, middle Miocene, Kleinleipisch (Brandenburg, Germany). Mai (1985, 2001a, b). Add. Occ. Ref.: Mosbrugger et al. (2005a).

  7. 7.

    Itea europaea, middle Miocene, Klettwitz (Brandenburg, Germany). Mai (1985, 2001a, b). Add. Occ. Ref.: Mosbrugger et al. (2005b).

  8. 8.

    Itea europaea, middle Miocene, Kreuzau (North Rhine-Westphalia, Germany). Pingen (1987) as Ericaceae div. sp., transferred to Itea europaea by Pingen (1996). See also Ferguson et al. (1998).

  9. 9.

    Itea europaea, middle Miocene, Lavrovo (Tambov, Russia). Dorofeev (1988) as Carpolithus sp. 1, transferred to Itea europaea by Pingen (1996).

  10. 10.

    Itea europaea, middle Miocene, Plessa (Brandenburg, Germany). Mai (1985, 2000a, b, 2001a, b). Add. Occ. Ref.: Mai (1999, 2000c).

  11. 11.

    Itea europaea, late Miocene, Baldevo (Blagoevgrad, Bulgaria). Mai & Palamarev (1997), Palamarev et al. (2005).

  12. 12.

    Itea europaea, late Miocene, Gozdnica (Lubusz, Poland). Mai (2001a). Add. Occ. Refs.: Dyjor (1992), Mai (2001b).

  13. 13.

    Itea europaea, late Miocene, Hambach (North Rhine-Westphalia, Germany). Pingen (1996). Add. Occ. Refs.: Gee (2005).

  14. 14.

    Itea europaea, late Miocene, Kausche-Klara II (Brandenburg, Germany). Mai (1985, 2001a, b).

  15. 15.

    Itea europaea, late Miocene, Melnik (Blagoevgrad, Bulgaria). Mai & Palamarev (1997), Palamarev et al. (2005).

  16. 16.

    Itea europaea, late Miocene, Welzow (Brandenburg, Germany). Mai (1985, 2001a). Add. Occ. Ref.: Mai (2001b).

  17. 17.

    Itea europaea, Pliocene, Belfeld (Limburg, The Netherlands). Mai (1985).

  18. 18.

    Itea europaea, Pliocene, Boschi di Barbania (Piedmont, Italy). Martinetto (2001). Add. Occ. Ref.: Martinetto (1999).

  19. 19.

    Itea europaea, Pliocene, Dunarobba (Umbria, Italy). Martinetto (2001).

  20. 20.

    Itea europaea, Pliocene, Front Canavese (Piedmont, Italy). Martinetto (1999), Martinetto et al. (2007).

  21. 21.

    Itea europaea, Pliocene, Oberzella (Thuringia, Germany). Gümbel and Mai (2004).

  22. 22.

    Itea europaea, Pliocene, Pranzalito (Piedmont, Italy). Martinetto (2001).

  23. 23.

    Itea europaea, Pliocene, Stura di Lanzo (Piedmont, Italy). Martinetto (1999), Martinetto et al. (2007).

  24. 24.

    Itea europaea, Pliocene, Villafranca d’Asti (Piedmont, Italy). The evidence for this occurrence is ambiguous. Martinetto (1999) states in the text that this taxon is present in this flora, but does not indicate that it is present in the accompanying table. Martinetto (2001) also seems to imply that the taxon may be present here. Not mapped.

Leaves

  1. 1.

    cf. Itea, early Eocene, Falkland (British Columbia, Canada). Smith (2011). Add. Occ. Refs.: Moss et al. (2005), Smith et al. (2009).

  2. 2.

    Itea, early Eocene, Republic (Washington, USA). Wolfe & Wehr (1987; 1991), Wehr and Hopkins (1994), Greenwood et al. (2005). Add. Occ. Ref.: Wolfe et al. (2003).

  3. 3.

    Itea faujasii, early Oligocene, Chiavon (Venetia, Italy). Massalongo (1859) as Cedrela faujasii, transferred to Itea faujasii by Meschinelli and Squinabol (1892). Add. Occ. Ref.: Hably et al. (2007).

  4. 4.

    Itea faujasii, late Miocene, Rochessauve (Rhône-Alpes, France). Faujas de Saint-Fond (1803) and Unger (1845) as Cedrela, Unger (1850) as Cedrela faujasii, transferred to Itea faujasii by Meschinelli and Squinabol (1892). Add. Occ. Ref.: Grangeon (1958). Holotype: Faujas de Saint-Fond (1803), pl. 56, fig. 3.

  5. 5.

    Itea transsilvanica, late Miocene, Chiuzbaia (Maramures, Romania). Petrescu and Givulescu (1986). Holotype: Petrescu and Givulescu (1986), fig. 3.

Pollen

Eocene

  1. 1.

    Itea, early Eocene, Falkland (British Columbia, Canada). Moss et al. (2005). Add. Occ. Ref.: Smith et al. (2009).

  2. 2.

    Itea, early Eocene, Hat Creek (British Columbia, Canada). Moss et al. (2005). Add. Occ. Refs.: Church (1975), Church et al. (1979).

  3. 3.

    Itea, early Eocene, McAbee (British Columbia, Canada). Moss et al. (2005). Add. Occ. Refs.: Hills and Baadsgaard (1967), Ewing (1981), Dilhoff et al. (2005).

  4. 4.

    Itea, early Eocene, Republic (Washington, USA). Moss et al. (2005). Add. Occ. Ref.: Wolfe et al. (2003).

  5. 5.

    Iteapollis angustiporatus, late Eocene, Colwell Bay (Isle of Wight, UK). Gruas-Cavagnetto (1976). Add. Occ. Refs.: King (2006), Hooker et al. (2009). Oldest figured fossil Itea pollen: Gruas-Cavagnetto (1976), pl. 9, fig. 15.

Oligocene (or Range Beginning in Oligocene)

  1. 1.

    Itea, early Oligocene, Rex Creek Coal Basin (Alaska, USA). Wahrhaftig et al. (1969). Add. Occ. Refs.: Wiggins et al. (1988), Marincovich and Wiggins (1990). Wahrhaftig et al. (1969) also reported Itea pollen in upper Miocene sediments of the Grubstake Formation from the same area, but the pollen was thought to have been reworked from older deposits; the late Miocene occurrence was thus not considered valid and is not mapped here.

  2. 2.

    Iteapollis angustiporatus, early Oligocene, Tămaşa (Sălag, Romania). Petrescu and Givulescu (1986) as Iteapollenites angustiporatus.

  3. 3.

    Iteapollis angustiporatus, late Oligocene, Enspel (Rhineland-Palatinate, Germany). Herrmann (2007) and Poschmann et al. (2010) as Iteapollis angustiporus. Add. Occ. Ref.: Uhl and Herrmann (2010).

  4. 4.

    Itea, late Oligocene, Gressk (Minsk, Belarus). Rylova (Rylova 1991, 1993, Rylova 1996).

  5. 5.

    Itea, late Oligocene, Mlamolovo (Kyustendil, Bulgaria). Ivanov (1996, 2004), Palamarev et al. (1998), Bozukov et al. (2009).

  6. 6.

    Itea spp., late Oligocene & middle Miocene, Smolyarka (Brest, Belarus). Late Oligocene: Murashko et al. (1998) as Itea. Middle Miocene: Rylova (1989) as Itea bielorussica, I. menkei, and I. miocenica. Holotype: I. bielorussica, Rylova (1989), pl. 1, fig. 16 & pl. 2, fig. 9. Holotype: I. menkei, Rylova (1989), pl. 1, fig. 15 & pl. 2, fig. 8. Holotype: I. miocenica, Rylova (1989), pl. 1, fig. 18 & pl. 2, fig. 10.

Miocene, Subdivision Unknown

  1. 1.

    Iteapollis angustiporatus, Miocene, Olszyna (Lubusz, Poland). Sadowska (1973) as Itea, synonymized with Iteapollis angustiporatus by Thiele-Pfeiffer (1980). Not Mapped.

  • early Miocene (or range beginning in early Miocene)

    1. 1.

      Iteapollis angustiporatus, early Miocene, Brandon Lignite (Vermont, USA). Traverse (1955) as Corylus?, Traverse (1994) as Iteapollis angustiporatus. Pollen illustrated by Traverse (1955, fig. 9(35)) recognized as cf. Itea by Ziembińska and Niklewski (1966), recognized as Itea by Petrov and Drazheva-Stamatova (1973) and Sadowska (1973), recognized as equivalent to Iteapollis angustiporatus by Ziembińska-Tworzydło (1974), synonymized with Iteapollis angustiporatus by Thiele-Pfeiffer (1980). Add. Occ. Ref.: Tiffney (1994). Oldest figured fossil Itea pollen from North America: Traverse (1955), fig. 9(35); Traverse (1994), pl. 2, fig. 17.

    2. 2.

      Iteapollis angustiporatus, early-middle Miocene, Gierlachowo (Greater Poland, Poland). Ziembińska-Tworzydło (1974). Add Occ. Refs.: Piwocki and Ziembińska-Tworzydło (1997).

    3. 3.

      Iteapollis angustiporatus, early-middle Miocene, Gołębin Stary (Greater Poland, Poland). Ziembińska-Tworzydło (1974), Ziembińska-Tworzydło & Ważyńska (1981), Ziembińska-Tworzydło et al. (1994). Add. Occ. Ref.: Piwocki and Ziembińska-Tworzydło (1997).

    4. 4.

      Itea-type, early Miocene, Nenana Coal Field (Alaska, USA). Leopold and Liu (1994).

    5. 5.

      cf. Itea, early-middle Miocene, Niedźwiedzice (Lower Silesia, Poland). Ziembińska and Niklewski (1966).

    6. 6.

      Itea, early Miocene, Nizhino (Primorsky, Russia). Pavlyutkin and Petrenko (2010). Add. Occ. Ref.: Pavlyutkin and Chekryzhov (2007).

    7. 7.

      Iteapollis angustiporatus, early-middle Miocene, Nowa Wieś (Lower Silesia, Poland). Ziembińska-Tworzydło (1974). Add. Occ. Ref.: Piwocki and Ziembińska-Tworzydło (1997).

    8. 8.

      Iteapollis angustiporatus, early Miocene, Oder II (Bavaria, Germany). Thiele-Pfeiffer (1980).

    9. 9.

      Itea, early-middle Miocene, Pacific Northwest, localities unknown (California, Idaho, Oregon, & Washington, USA). Gray (1964), Wolfe (1969).

    10. 10.

      Itea, early-middle Miocene, Rittsteig (Bavaria, Germany). Seitner (2004b).

    11. 11.

      Iteapollis angustiporatus, early-middle Miocene, Ślepuchowo (Greater Poland, Poland). Ziembińska-Tworzydło (1974). Add. Occ. Ref.: Piwocki and Ziembińska-Tworzydło (1997).

    12. 12.

      Iteapollis angustiporatus, early-middle Miocene, Trwyn y Parc (Wales, UK). Walsh et al. (1996). Add. Occ. Refs.: Morawiecka et al. (1996), Walsh et al. (1999).

    13. 13.

      Iteapollis angustiporatus, early-middle Miocene, Ustronie (Lower Silesia, Poland). Ziembińska and Niklewski (1966) as cf. Itea, Ziembińska-Tworzydło (1974) as Iteapollis angustiporatus. Pollen figured by Ziembińska and Niklewski (1966, pl. 7, fig. 22) synonymized with Iteapollis angustiporatus by Thiele-Pfeiffer (1980). Add. Occ. Ref.: Piwocki and Ziembińska-Tworzydło (1997). First figured fossil Itea pollen identified as Itea : Ziembińska and Niklewski (1966), pl. 7, fig. 22.

    14. 14.

      cf. Itea, early-middle Miocene, Wirczyn (Lower Silesia, Poland). Ziembińska and Niklewski (1966).

    15. 15.

      Iteapollis angustiporatus, early Miocene, Zăghid-Hida (Sălag, Romania). Petrescu and Givulescu (1986) as Iteapollenites angustiporatus.

  • middle Miocene (or range beginning in the middle Miocene)

    1. 1.

      Itea bielorussica, middle Miocene, Bukcha (Homyel’, Belarus). Rylova (2004). Add. Occ. Ref.: Yakubovskaya et al. (2005).

    2. 2.

      Itea, middle Miocene, Deleina (Vidin, Bulgaria). Ivanov et al. (2002a), Ivanov (2004). Add. Occ. Ref.: Ivanov et al. (2002b).

    3. 3.

      Itea, middle-late Miocene, Drenovets (Vidin, Bulgaria). Ivanov et al. (2002a), Ivanov (2004). Add. Occ. Ref.: Ivanov et al. (2002c).

    4. 4.

      Itea, middle Miocene, GSW/3 (Lubusz, Poland). Sadowska (1992). Add. Occ. Ref.: Dyjor (1992).

    5. 5.

      Itea, middle Miocene, GSW/6 (Lubusz, Poland). Sadowska (1992). Add. Occ. Ref.: Dyjor (1992).

    6. 6.

      Iteapollis angustiporatus, middle Miocene, Incescu-Zambal (Çorum, Turkey). Kayseri and Akgün (2008).

    7. 7.

      Itea, middle Miocene, Jaroszów (Lower Silesia, Poland). Sadowska (1977). Add. Occ. Ref.: Dyjor and Sadowska (1986).

    8. 8.

      Itea, middle Miocene, Jerzmanowa (Lubusz, Poland). Sadowska (1977). Add. Occ. Ref.: Dyjor and Sadowska (1986).

    9. 9.

      Iteapollis angustiporatus, middle Miocene, Karolewo-Dąbki 3 (Greater Poland, Poland). Kohlman-Adamska (1993).

    10. 10.

      Iteapollis angustiporatus, middle Miocene, Karolewo-Dąbki 4 (Greater Poland, Poland). Kohlman-Adamska (1993).

    11. 11.

      Iteapollis angustiporatus, middle Miocene, Komorniki (Lower Silesia, Poland). Worobiec (2009).

    12. 12.

      Iteapollis angustiporatus, middle Miocene, Konin (Greater Poland, Poland). Ziembińska-Tworzydło et al. (1994).

    13. 13.

      Iteapollis angustiporatus, middle Miocene, Kosztowo (Greater Poland, Poland). Kohlman-Adamska (1993).

    14. 14.

      Itea, middle Miocene, Kreuzau (North Rhine-Westphalia, Germany). Ferguson et al. (1998).

    15. 15.

      Iteapollis angustiporatus, middle Miocene, Krosinko (Greater Poland, Poland). Ziembińska-Tworzydło (1974). Add. Occ. Ref.: Piwocki and Ziembińska-Tworzydło (1997).

    16. 16.

      Iteapollis angustiporatus, middle Miocene, Legnica (Lower Silesia, Poland). Sadowska (1977) as Itea, Wacnik and Worobiec (2001) and Worobiec (2009) as Iteapollis angustiporatus. Add. Occ. Ref.: Dyjor and Sadowska (1986).

    17. 17.

      Iteapollis angustiporatus, middle Miocene, Liszkowo (Greater Poland, Poland). Kohlman-Adamska (1993).

    18. 18.

      Itea, middle Miocene, Łojowice (Lower Silesia, Poland). Sadowska (1977). Add. Occ. Ref.: Dyjor and Sadowska (1986).

    19. 19.

      Iteapollis, middle Miocene, Lubstów (Greater Poland, Poland). Durska (2008).

    20. 20.

      Itea, middle-late Miocene, Lukasberg (Upper Austria, Austria). Masselter and Hofmann (2005).

    21. 21.

      Itea, middle Miocene, Makresh (Vidin, Bulgaria). Ivanov et al. (2002a), Ivanov (2004). Add. Occ. Ref.: Ivanov et al. (2002d).

    22. 22.

      Itea, middle Miocene, Mirostowice (Lubusz, Poland). Sadowska (1973, 1977) as Itea. Add. Occ. Ref.: Dyjor and Sadowska (1986).

    23. 23.

      Iteapollis angustiporatus, middle Miocene, Mosina (Greater Poland, Poland). Ziembińska-Tworzydło (1974). Add. Occ. Ref.: Piwocki and Ziembińska-Tworzydło (1997).

    24. 24.

      Iteapollis angustiporatus, middle Miocene, Oczkowice (Greater Poland, Poland). Ziembińska-Tworzydło (1974). Add. Occ. Ref.: Piwocki and Ziembińska-Tworzydło (1997).

    25. 25.

      Iteapollis angustiporatus, middle Miocene, Oder I (Bavaria, Germany). Thiele-Pfeiffer (1980).

    26. 26.

      Itea, middle Miocene, Pątnów (Greater Poland, Poland). Sadowska and Giża (1991).

    27. 27.

      Itea, middle Miocene, Ruszów (Lower Silesia, Poland). Sadowska (1977). Add. Occ. Ref.: Dyjor and Sadowska (1986).

    28. 28.

      Iteapollis angustiporatus, middle Miocene, Rypin (Kuyavia-Pomerania, Poland). Stuchlik (1964) as Corylus americana after figure of Iteapollis labeled Corylus? by Traverse (1955), recognized as Itea by Petrov and Drazheva-Stamatova (1973) and Sadowska (1973), recognized as equivalent to Iteapollis angustiporatus by Ziembińska-Tworzydło (1974), synonymized with Iteapollis angustiporatus by Thiele-Pfeiffer (1980). See also Ziembińska-Tworzydło et al. (1994, pl. 8, fig. 12), pollen refigured from Stuchlik (1964, pl. 11, fig. 8) assigned to Iteapollis angustiporatus.

    29. 29.

      Iteapollis angustiporatus, middle Miocene, Spreetal (Saxony, Germany). Schneider (1965) as Psilodiporites angustiporatus, synonymized with Iteapollis angustiporatus by Ziembińska-Tworzydło (1974). Holotype: Schneider (1965), pl. 1, fig. 10.

    30. 30.

      Iteapollis angustiporatus, middle Miocene, Tuplice (Lubusz, Poland). Sadowska (1973, 1977) Pollen figured by Sadowska (1973, pl. 2, fig. 1–6) synonymized with Iteapollis angustiporatus by Thiele-Pfeiffer (1980). Add. Occ. Ref.: Dyjor and Sadowska (1986).

    31. 31.

      Iteapollis angustiporatus, middle Miocene, Vad-Borod Basin (Bihor, Romania). Petrescu and Givulescu (1986) as Iteapollenites angustiporatus.

    32. 32.

      Itea menkei, middle Miocene, Wakendarf-II (Schleswig-Holstein, Germany). Menke (1976) as Itea, assigned to Itea menkei by Rylova (1989).

    33. 33.

      Itea, middle Miocene, Zarów (Lower Silesia, Poland). Sadowska (1977). Add. Occ. Ref.: Dyjor and Sadowska (1986).

  • late Miocene (or range beginning in the late Miocene)

    1. 1.

      Iteapollis angustiporatus, late Miocene, Achldorf (Bavaria, Germany). Seitner (2004a).

    2. 2.

      Itea, late Miocene, Badersdorf (Burgenland, Austria). Hofmann and Zetter (2005).

    3. 3.

      Iteapollis angustiporatus, late Miocene-Pliocene, Beli Breg Mine (Sofia, Bulgaria). Petrov and Drazheva-Stamatova (1973), Ivanov (2004) and Ivanov et al. (2007) as Itea. Pollen figured by Petrov and Drazheva-Stamatova (1973, figs. 1–2) synonymized with Iteapollis angustiporatus by Thiele-Pfeiffer (1980).

    4. 4.

      Itea, late Miocene, Cherasco (Piedmont, Italy). Bertini and Martinetto (2011).

    5. 5.

      Itea, late Miocene, Czarny Dunajec (Lesser Poland, Poland). Oszast and Stuchlik (1977). Add. Occ. Refs.: Stuchlik (1980), Kovar-Eder (1987).

    6. 6.

      Iteapollis angustiporatus, late Miocene, Darova-Lugoj (Timis, Romania). Petrescu and Kolovos (1982). Also, Petrescu and Givulescu (1986) as Iteapollenites angustiporatus.

    7. 7.

      Itea, late Miocene, DSDP Site 380A (Black Sea). Traverse (1978), Traverse et al. (2005). Add. Occ. Ref.: Popescu (2006).

    8. 8.

      Itea, late Miocene-Pliocene, Gozdnica (Lubusz, Poland). Sadowska (1977). Add. Occ. Refs.: Dyjor and Sadowska (1986), Dyjor (1992).

    9. 9.

      Itea, late Miocene, Gozdnica-Stanisław (Lubusz, Poland). Sadowska (1973, 1992). Add. Occ. Ref.: Dyjor (1992).

    10. 10.

      Itea, late Miocene-Pliocene, Katina (Blagoevgrad, Bulgaria). Hristova and Ivanov (2009a, b).

    11. 11.

      Itea, late Miocene, Maccarone (Marche, Italy). Bertini (2006).

    12. 12.

      Iteapollis angustiporatus, late Miocene, Melnik (Blagoevgrad, Bulgaria). Petrov and Drazheva-Stamatova (1973) and Ivanov (2001a, 2004) as Itea, Ivanov (2001b) as Iteapollis angustiporatus. Pollen figured by Petrov and Drazheva-Stamatova (1973, figs. 3–4) synonymized with Iteapollis angustiporatus by Thiele-Pfeiffer (1980).

    13. 13.

      Itea, late Miocene, Nysa (Opolskie, Poland). Sadowska (1977). Add. Occ. Ref.: Dyjor and Sadowska (1986).

    14. 14.

      Itea, late Miocene-Pliocene, Ohoopee Dune Field (Georgia, USA). Rich et al. (2002). Add. Occ. Ref.: Hansen et al. (2001).

    15. 15.

      Itea, late Miocene, Opole (Opolskie, Poland). Sadowska (1977). Add. Occ. Ref.: Dyjor and Sadowska (1986).

    16. 16.

      Itea, late Miocene-Pliocene, Paczków (Opolskie, Poland). Sadowska (1977). Add. Occ. Ref.: Dyjor and Sadowska (1986).

    17. 17.

      Itea, late Miocene-Pliocene, Poznań (Greater Poland, Poland). Troć and Sadowska (2006).

    18. 18.

      Iteapollis angustiporatus, late Miocene, Roşiori-Oradea (Bihor, Romania). Petrescu and Givulescu (1986) as Iteapollenites angustiporatus.

    19. 19.

      Itea, late Miocene-Pliocene, Sarzana (Liguria, Italy). Bertoldi et al. (1994), Bertoldi (1997), Bertini and Martinetto (2008).

    20. 20.

      Iteapollis angustiporatus, late Miocene, Vasiltepe (Sivas, Turkey). Kayseri and Akgün (2008).

    21. 21.

      Itea, late Miocene, Wrocław (Lower Silesia, Poland). Sadowska (1977). Add. Occ. Ref.: Dyjor and Sadowska (1986).

Pliocene

  1. 1.

    Itea, Pliocene, Ahrensburg (Schleswig-Holstein, Germany). Menke (1975).

  2. 2.

    Iteapollis angustiporatus, pollen, Pliocene, Hausruck brown coal (Austria). Klaus (1953) as Diorites sp. (V: Symplocaceae), synonymized with Iteapollis angustiporatus by Thiele-Pfeiffer (1980). First figured fossil Itea pollen: Klaus (1953), pl. 1, fig. 7, as Diorites sp. (V: Symplocaceae). Not mapped.

  3. 3.

    Itea, Pliocene, Horn Island (Mississippi, USA). Gohn et al. (1996).

  4. 4.

    Iteapollis angustiporatus, Pliocene, Oldenswort (Schleswig-Holstein, Germany). Menke (1975, 1976) as Itea. Pollen figured by Menke (1976, pl. 16, figs. 12–15) synonymized with Iteapollis angustiporatus by Thiele-Pfeiffer (1980).

  5. 5.

    Itea, Pliocene, Santa Barbara (Tuscany, Italy). Bertini and Roiron (1997), Bertini (2002; 2010), Bertini and Martinetto (2011).

  6. 6.

    Itea, Pliocene, Olten (Romania). Verbal communication of Ms. St. Roman to Petrescu and Givulescu (1986). Not mapped.

  7. 7.

    Itea, Pliocene, Sośnica (Lower Silesia, Poland). Stachurska et al. (1973). Add. Occ. Ref.: Dyjor and Sadowska (1986).

  8. 8.

    Itea, Pliocene, Wólka Ligezowska (Masovia, Poland). Popescu et al. (2010).

  9. 9.

    Itea, pollen, Pliocene, locality unknown (The Netherlands). Personal communication of Zagwijn to Muller (1981). Not mapped.

Pleistocene (or Range Beginning in Pleistocene)

  1. 1.

    Itea, Pleistocene, Green Pond (Georgia, USA). Watts (1973).

  2. 2.

    Itea, Pleistocene-Holocene, Lake Louise (Georgia, USA). Watts (1971, 1980).

  3. 3.

    Itea virginica, Pleistocene, Wilcox Bluff (Louisiana, USA). Delcourt and Delcourt (1977).

Holocene (Less Than 11,700 Years Old, Not Mapped)

  1. 1.

    Itea virginica, Holocene (ca. 10,000 years b.p. to recent), Anderson Pond (Tennessee, USA). Delcourt (1979).

  2. 2.

    Itea, Holocene (ca. 3,500–7,300 years b.p.), B.L. Bigbee (Mississippi, USA). Whitehead and Sheehan (1985).

  3. 3.

    Itea, Holocene, Bob Black Pond (Georgia, USA). Watts (1970).

  4. 4.

    Itea virginica, Holocene (ca. 7,300 years b.p. to recent), Jackson Pond (Kentucky, USA). Wilkins et al. (1991).

  5. 5.

    Itea, Holocene (<2,000 years b.p. to recent), Minnie’s Lake (Georgia, USA). Fair-Page and Cohen (1990).

  6. 6.

    Itea, Holocene (4,065–6,340 years b.p.), Mitchell River 1 (Florida, USA). Saunders et al. (2009).

  7. 7.

    Itea, Holocene (Georgia, USA), Quicksand Pond. Watts (1970).

  8. 8.

    Itea, Holocene (5,000 years b.p. to recent), Rockyhock Bay (North Caroline, USA). Whitehead (1981).

Appendix 2

Table 2 List of localities and coordinates used in mapping. Abbreviations for organs: fl, flower; fs, fruit/seed; l, leaf; p, pollen. Abbreviations for age intervals: eEo early Eocene, lEo late Eocene, eOl early Oligocene, lOl late Oligocene, eMio early Miocene, mMio middle Miocene, lMio late Miocene, Plio Pliocene, Plei Pleistocene, Hol Holocene
Full size table

Appendix 3

Samples of Itea europaea examined at the Museum für Naturkunde, Berlin, Germany.

BHUPM: Berzdorf Hgd. 8824; Gozdnica 1999/365; Hambach 7E-F 1998/601; Hambach 9B 1999/63; Hambach Fp. 1 2001/125; Hambach Fp. 2 2001/209; Hartau 1048; Klara II Kausche 4695; Kleinleipisch 2 3979; Klettwitz VFI/OFI 1993/877; Klettwitz 2 1236; Klettwitz 2 2036; Klettwitz 3 3834; Klettwitz 4 3926; Klettwitz 5 3642; Plessa Oberbank 5056; Reichwalde 5151; Reuverton/Belfeld 7830; Römerkeller 6163; Welzow 4388.

Appendix 4

Specimens of putative Itea leaves from Republic, Washington, examined for this study.

UWBM: UWBM 31262, UWBM 31268, UWBM 36812, UWBM 36813, UWBM 36814, UWBM 36815, UWBM 55069, UWBM 56689, UWBM 57466 (2 leaves), UWBM 76355, UWBM 76359, UWBM 76363 (loc. B4131), UWBM 76365, UWBM 76366, UWBM 76369, UWBM 76372, UWBM 76377, UWBM 76379, UWBM 76383, UWBM 76385, UWBM 77440, UWBM 77441, UWBM 95509, UWBM 97090, UWBM 97579, UWBM 97580, UWBM 97582. Part/Counterpart specimens: UWBM 31260A, B; UWBM 36871A, B; UWBM 39725, UWBM 76391; UWBM 39732A, B; UWBM 56593A, B; UWBM 76356, UWBM 76360; UWBM 76357, UWBM 76361; UWBM 76358, UWBM 76374; UWBM 76362, UWBM 97578; UWBM 76363 (loc. A0307), UWBM 76382; UWBM 76367A, B; UWBM 76371, UWBM 76381; UWBM 76373, 76376; UWBM 76375, UWBM 76378; UWBM 76581, UWBM 97581.

SR: SR 00-04-26, SR 00-07-05, SR 01-7-41, SR 01-12-2, SR 01-12-03, SR 01-12-04, SR 02-19-04, SR 02-27-03, SR 02-30-43, SR 90-5-7, SR 90-12-15, SR 92-13-4, SR 92-17-6, SR 92-17-9, SR 93-1-3, SR 95-25-60, SR 95-27-17, SR 99-9-01. Part/counterpart specimens: SR 01-07-09A, B; SR 93-9-4A, B; SR 97-5-2A, B; SR 99-18-4A, B.

USNM: Unnumbered (Loc. USGS 8428).

Part/Counterpart specimens from different collections: SR 93-4-1, UWBM 76370; UAPC-ALTA S13290, UWBM 76380.

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Hermsen, E.J. A Review of the Fossil Record of the Genus Itea (Iteaceae, Saxifragales) with Comments on its Historical Biogeography. Bot. Rev. 79, 1–47 (2013). https://doi.org/10.1007/s12229-012-9114-3

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