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A review of the fossil record of the genus Itea (Iteaceae, Saxifragales) with comments on its historical biogeography.

Introduction

Itea L. (Iteaceae J. Agardh, Saxifragales sensu APG III 2009) is a small genus of about 20 species (Table 1) of shrubs and trees of tropical to temperate forest, forest margin, or wet (e.g., swamp and riparian) habitats ranging in elevation from sea level up to about 3,000 m. The genus includes plants having simple, pinnately-veined leaves, with usually hermaphroditic flowers. Each flower is characterized by an actinomorphic, pentamerous perianth composed of a whorl of sepals and a whorl of petals, a haplostemonous androecium with antesepalous stamens, and a bicarpellate gynoecium; the styles of the flowers are fused to free, and the single stigma is capitate. The flowers mature into septicidal capsules bearing multiple seeds (e.g., Spongberg, 1972; Verdcourt, 1973; Jin & Ohba, 2001; Ge et al., 2002; Siddiqi, 2005; Kubitzki, 2007a; Morin, 2009). Phylogenetic analyses based on molecular sequence datasets clearly indicate that Itea belongs within the Saxifragales sensu APG III (relevant research summarized in Soltis et al., 2005; see most recently Soltis et al. 2011), and recent analyses have consistently suggested that Itea is part of a subclade within Saxifragales called the "Saxifragaceae Alliance" (Fishbein et al., 2001; Soltis et al., 2007, Soltis et al. 2011; Jian et al., 2008). This clade also includes Pterostemon S. Schauer, the sister genus to Itea, which was recently transferred into Iteaceae from the monotypic family Pterostemonaceae Small (APG III, 2009); as well as the family Saxifragaceae Juss. and its sister genus Ribes L. (Grossulariaceae DC sensu APG III), which together form the sister clade to Iteaceae (Fig. 1). While the flowers of Itea are morphologically and anatomically similar to those of the closely related species within Saxifragaceae and Ribes (Bensel & Palser, 1975a, b), Itea is easily distinguished by its unusual bilaterally symmetrical, diporate pollen grains (e.g., Erdtman, 1952; Bensel & Palser, 1975b; Hideux & Ferguson, 1976).

The monophyly of the genus Itea as traditionally circumscribed, while not yet formally tested in a phylogenetic analysis, is not considered in question. Kubitzki (2007a), however, recently argued that the putative sister taxon to Itea, the morphologically similar African shrub Choristylis rhamnoides Harv., should be subsumed within Itea. While in the past the two genera have consistently been recognized as separate (e.g., Bentham & Hooker 1862-1867; Engler, 1890; Takhtajan, 1997), Kubitzki argued that the expansion of Itea to include more species over time has resulted in morphological overlap of Itea with Choristylis Harv. where floral and inflorescence characters were previously distinct. He further pointed out that the remaining significant morphological difference--number of ovule integuments--putatively justifying a separation between Itea (bitegmic) and Choristylis (previously reported as unitegmic) is based on a misinterpretation of the number of ovule integuments in Choristylis (Kubitzki, 2007a). He thus suggested the new combination Itea rhamnoides (Harv.) Kubitzki for the African taxon. His conclusion is in concordance with phylogenetic studies based on molecular markers that strongly suggest that Itea and Choristylis form a monophyletic group (Fishbein et al., 2001; Soltis et al., 2007, Soltis et al. 2011; Jian et al., 2008). Thus, expansion of Itea to include Choristylis maintains monophyly of the genus under recent estimates of phylogeny and yields a morphologically consistent taxon where all members share a unique pollen morphology that distinguishes them from plants in closely related genera.

The significance of this minor expansion of the generic limits of Itea has interesting implications for understanding the historical biogeography of the genus. Itea has generally been viewed as a classic eastern North American-eastern Asian disjunct taxon (Li, 1952; Wu, 1983; Wen, 1999; Guo & Ricklefs, 2000), with evidence from fossils suggesting that its geographical distribution was likely relictual from a Paleogene to Neogene Northern Hemisphere distribution that included at least Europe and eastern North America (e.g., Petrov & Drazheva-Stamatova, 1973; Mai, 1985). Today, one species (I. virginica) occurs widely in wet lowland habitats and in association with ponds or streams at higher elevations in the southeastern United States (Spongberg, 1972; Morin, 2009). About 18 species generally occur in montane habitats (100 to 3,000 m) in eastern Asia, ranging from the island of Honshu (Japan) in the northeast, west along the Himalayas into Pakistan in the northwest, and as far south as the island of Java (Indonesia); the center of diversity for the genus is in southeastern China (e.g., Merrill, 1921; Parker, 1924; Spongberg, 1972; Jin & Ohba, 2001; Ohba & Niu 2001; Siddiqi, 2005; Kubitzld, 2007a). The addition of I. rhamnoides expands the distribution of the genus to include a disjunct region of eastern Africa, extending from Uganda to South Africa (Verdcourt, 1973). Thus, rather than appearing only in three of five regions (present in southeastern North America, northeastern Asia, and southeastern Asia; absent in western North American and southwestern Eurasia) considered refugia for subtropical to warm temperate Northern Hemisphere "Tertiary relict genera" (Milne & Abbott, 2002; Milne, 2006), Itea displays an eastern North American-eastern Asian-eastern African disjunct pattern (Fig. 2). This distribution pattern is unusual among genera displaying an otherwise eastern North America-eastern Asia disjunct pattern (Guo & Ricklefs, 2000). In the present study, I will review the extensive record of fossils attributed to Itea and explore how the genus might have achieved its present range based on the evidence obtained from fossils.

Materials and Methods

Material Examined

Specimens of most material discussed in this study were not examined. Collections of the fruit and seed taxon Itea europaea Mai held at the Museum fur Naturkunde (BHUPM) in Berlin, Germany, were, however, examined (Appendix 3). While no new conclusions were reached concerning the morphology of I. europaea, the locality information stored with the specimens supplemented locality information for L europaea provided in published papers.

Fossils representing 68 leaves (from 67 specimens, part-counterpart specimens considered single specimens) thought to represent Itea sp. from the Republic flora were also examined (Appendix 4). These include fossils held at the Stonerose Interpretive Center and Eocene Fossil Site (SR) in Republic, Washington, USA; the University of Washington Burke Museum of Natural History and Culture (UWBM) in Seattle, Washington, USA; the United States National Museum of Natural History (USNM), in Washington, DC, USA; and the University of Alberta Paleobotanical Collection (UAPC-ALTA) in Edmonton, Alberta, Canada. The terminology and characteristics outlined by Ellis et al. (2009) were used as the guide for the informal description of the Republic leaves, and Ellis et al. (2009) were followed in describing leaf venation in this paper. A small selection of fossil leaves from the Republic flora is shown in Figs. 3, 4, 5, 6, 7, 8 and 9.

Herbarium specimens of Itea were examined for leaf characteristics at the following collections: the United States National Herbarium (US) Washington, DC, USA; the Wiegand Herbarium (BH), Cornell University, Ithaca, New York, USA; and the herbarium of the Botanischer Garten und Botanisches Museum Berlin-Dahlem (B), Freie Universitat, Berlin, Germany. Leaves of some extant taxa were cleared and mounted for comparison to the fossil leaves from the Republic flora using methods adapted from Buechler (2004a, b). Each leaf was treated with a solution of 10 % NaOH followed by solution of 20 % bleach to remove as much pigment as possible while still maintaining the integrity of the lamina. Following treatment by bleach, the leaves were soaked in distilled water, then 50 % EtOH; each soak lasted 24 h. Afterward, the leaves were stained in a solution of safianin dissolved in 50 % EtOH. Excess stain was removed by soaking the leaves in 50 % EtOH, then 100 % EtOH. Leaves were mounted between two sheets of HighlandMC Transparency Film for Copiers using cedar oil (Texas Fragrances, Leakey, Texas, USA) as a mounting medium. Cleared leaves are shown in Figs. 10, 11, 12, 13 and 14.

Imaging

Photographs of fossil specimens were taken with a Sony Cyber-Shot DSC-F717 digital camera. The original backgrounds surrounding the rock matrix of the fossil specimens was digitally removed using Adobe[R] Photoshop[R] CS4 Extended Ver. 11 ([C]1999-2008 Adobe Systems Inc.). Cleared leaf specimens were scanned using a CanoScan LiDE 100 flatbed scanner. All plates and figures were assembled in Photoshop[R]. Linear adjustments to brightness and contrast were made on the images of the cleared leaf specimens in Photoshop[R].

Mapping

Spatiotemporal records of the occurrences of fossils thought to represent stem- or crown-group Itea were identified and mapped primarily using published sources. In several instances, where unpublished sources (dissertations, preliminary government reports) were the only or best documentation for an occurrence, they have been cited. Mapped occurrences include both figured and unfigured material. Google[TM] Language Tools were used to translate or transliterate text in some sources, where necessary. Each locality or flora was mapped in Google[TM] Earth 5.2.1.1588 in order to generate coordinates in decimal degrees. Although published coordinates were sometimes available, the position of most occurrences was estimated using the coordinates of nearby towns or geographic features. In order to produce distribution maps (Figs. 15, 16, 17 and 18), occurrences were mapped in ArcMap ver. 9.3.1 (Environmental Systems Research, Inc., Redlands, California) using World Continents (ESRI, Source: ArcWorld Supplement) as a base map; points were plotted using the geographic coordinate system WGS 1984, corresponding to the system utilized by Google[TM] Earth. A list of fossils thought to represent stem- or crown-group Itea with temporal and geographic information is given in Appendix 1; a list of occurrences (localities and floras) and their coordinates used in producing distribution maps for fossils are given in Appendix 2.

Because there are few Eocene records, all were included in the same distribution map, although there is a significant temporal gap separating the initial and later Eocene occurrences. Other occurrences were mapped in roughly equal-age segments of time: Oligocene (ca. 11 million years), early-middle Miocene (ca. 11.4 million years), and late Miocene-Pleistocene (ca. 11.6 million years). The final time interval coincides with a steep drop in global temperature (Zachos et al., 2001). Numerical ages for time interval boundaries are based on Ogg (2009).

Fossils Attributed to Itea

As is true of most plant fossils, fossils attributed to, thought to represent, or thought to be closely related to Itea are preserved as dispersed organs rather than as whole plants. Among these fossils are dispersed pollen, capsulate fruits (sometimes containing seeds), and dispersed seeds, leaves, and flowers. Each fossil type is briefly introduced below, along with a discussion of why each is considered to have affinities to Itea. Fossil types are ordered in what I believe to reflect the degree of certainty (from most to least) that can be attached to their inclusion within the Itea crown clade or its stem group.

Pollen

Fossil pollen attributed to Itea is often simply assigned to the genus with no specific epithet attached; however, the dispersed pollen taxon for pollen grains with morphology consistent with those of Itea is Iteapollis angustiporatus (Schneider) Ziembifiska-Tworzydlo (Ziembifiska-Tworzydto, 1974). Dispersed pollen has also been assigned the epithets Itea bielorussica Rylova, Itea menkei Rylova, and Itea miocenica Rylova (Rylova, 1989), and is sometimes assigned to the extant species I. virginica in Quaternary sediments of the southeastern United States (Delcourt & Delcourt, 1977; Delcourt, 1979; Wilkins et al., 1991). Fossil Itea pollen was first connected to the extant genus in the 1960's (Gray, 1964; Ziembinska & Niklewski, 1966; Wahrhaftig et al., 1969; Wolfe, 1969), and first illustrated as cf. Itea in 1966 (Ziembifiska & Niklewski, 1966, pl. 7, fig. 22). Prior to that, Klaus (1953) had illustrated fossil Itea pollen as Diorites sp. (perhaps a misspelling of the form genus name Diporites van der Hammen), and attributed it to Symplocaceae Desf. Traverse (1955) and Stuchlik (1964) illustrated it as pollen of the extant genus Corylus L. Schneider (1965) later assigned it to the form genus Psilodiporites Varma & Rawat and coined the species epithet angustiporatus, which is retained in the current binomial for the dispersed pollen species. The fossil form of the pollen did not receive direct comparative morphological study with extant Itea pollen until 1973 (Petrov & Drazheva-Stamatova, 1973; Sadowska, 1973). Dispersed pollen showing similarities to pollen of Itea was given its own generic name, Iteapollis, by Ziembinska-Tworzydto in 1974. The holotype of Iteapollis angustiporatus is from the Spreetal open-pit mine in the state of Saxony, Germany, and was illustrated by Schneider (1965, pl. 1, fig. 10); the age of this specimen is middle Miocene (Schneider, 1965). Thiele-Pfeiffer (1980) provides a synonymy list for figured Iteapollis angustiporatus pollen through 1980; for full details and references for synonymy of fossil Itea pollen grains, see Appendix 1.

Fossil Itea or Iteapollis pollen is diporate with the pores aligned on the equatorial axis. It is bilaterally symmetrical and more or less oblate in polar view; it is biconvex and subisopolar, with one pronouncedly convex pole and one more flattened pole in equatorial view. The exine is largely unomamented externally (psilate to scabrate) and has two layers that are visible using light microscopy (LM). In equatorial view, the pores are displaced toward the flatter pole of the grain. The size of the grains is slightly variable, reportedly about 10 to at least 22.2 [micro]m high along the polar axis and about 17 to 34.6 [micro]m long along the longer equatorial axis (Traverse, 1955; Schneider, 1965; Petrov & Drazheva-Stamatova, 1973; Sadowska, 1973; Ziembinska-Tworzydlo, 1974; Menke, 1976; Rylova, 1989). Thiele-Pfeiffer (1980) reported that the average grain size is about 12 by 21 [micro]m. This pollen type is reported from the early Eocene to recent in North America, the late Eocene to Pliocene in a region encompassing Europe and Anatolia, and from the early Miocene of eastern Asia (Appendix 1). The pollen morphology of extant Itea has been described, illustrated, and/or discussed by many investigators (e.g., Erdtman, 1952, 1955; Agababian, 1960; Wakabayashi, 1970; Pastre & Pons, 1973; Petrov & Drazbeva-Stamatova, 1973; Sadowska, 1973; Hideux, 1974; Hideux & Ferguson, 1976; Lieux, 1982; Nowicke & Skvarla, 1983; Frederiksen et al., 1985; Rylova, 1989). Overall, the extant and fossil pollen types are similar in shape, size, lack of exine sculpture, number of layers of exine visible under LM, and number and shape of the apertures.

One particularly interesting feature of fossil and extant Itea pollen is the appearance of the exine around the pores. Traverse (1955: 46) described the region around each pore in fossil Itea pollen from the Miocene Brandon lignite of Vermont, USA, as appearing "pebbly" when examined using LM. Observation of a textured area around each pore was also made and clearly illustrated by Petrov & Drazeheva-Stamatova (1973, esp. fig. 3) for Miocene pollen grains from Bulgaria and by Menke (1976, esp. plate 16, Figs. 13 and 14) for Pliocene pollen grains from Germany. Published light micrographs of L virginica (e.g., Hideux & Ferguson, 1976, plate 2, figs. 31-33) and L japonica (Wakabayashi, 1970, plate 3, esp. fig. y3) clearly show a distinctly textured area around the pores, and some authors have mentioned that there is a "roughened" zone (Lieux, 1982: 336) or a region of "roughening" (Frederiksen et al., 1985: 50) of the inner exine around the pores of Itea pollen when observed with a light microscope or from light photo micrographs. Using transmission electron microscopy (TEM), Nowicke and Skvarla (1983: 181) noted that "[t]he endexine appears loose and fragmented near the apertures" in extant Itea pollen; if the same is true of the fossil grains, this may explain the textured appearance of the exine surrounding the pores under light microscopy.

Erdtman (1955: 407), who studied and described the pollen of many different angiosperm taxa, wrote that "[t]he pollen type found in Itea is exceedingly rare." A recent phylogenetic analysis suggests that the diporate pollen type may be a synapomorphy for the Itea clade among the Saxifragales (Hermsen et al., 2006). Of the extant taxa most closely related to Itea, Saxifragaceae tend to have tricolpate or tricolporoidate pollen; Pterostemon has tricolporate pollen; and Ribes species generally have polyporate pollen, often with elongate to irregularly-shaped ectoapertural regions (e.g., Erdtman, 1952; Agababian, 1963; Ferguson & Webb, 1970; Hideux & Ferguson, 1976; Verbeek-Reuvers, 1977a, b; Weigend, 2007). Some unrelated angiosperms, however, do have pollen that morphologically resembles that of Itea, and Frederiksen et al. (1985) have reviewed and summarized the morphological similarities and differences among diporate, subisopolar, more or less psilate pollen types produced by extant and extinct plants. The pollen nearest that of Itea in morphology might be produced by the extant gentianaceous saprophyte genera Leiphaimos Schltdl. & Cham. and Voyria Aubl. (Spongberg, 1972; Frederiksen et al., 1985). Pollen of these taxa has a variable number of apertures (1-6), although diporate pollen forms can look remarkably like pollen of Itea (see photo micrographs in Nilsson & Skvarla, 1969). Itea pollen differs from the gentianaceous forms, however, in having two visible layers in the exine; layers in the gentianaceous forms are indistinct under LM, and also often with TEM (Nilsson & Skvarla, 1969; Frederiksen et al., 1985). Additionally, no textured zone is described surrounding the apertures of the gentianaceous taxa.

Pollen with the combined characteristics of Itea--diporate, psilate, subisopolar, bilaterally symmetrical, with a two-layered exine and a textured area around the apertures--may be unique among the angiosperms. Because it characterizes extant Itea, this pollen type arose somewhere on the stem lineage of the clade (Fig. 1). Thus, the occurrence of pollen consistent with that of extant Itea is an unambiguous indication of the Itea group in the fossil record, but not necessarily of the crown clade.

Most of the published occurrences of fossil Itea pollen are well documented with regard to geographic location and age (Appendix 1), although the pollen is often not figured. Reports pertaining to occurrences of fossil Itea pollen in the Neogene of the Pacific northwestern region of the United States are, however, vague. This is problematic because western North America is integral to understanding the historical biogeography of Itea, and there are few documented post-Eocene occurrences of Itea fossils from this region. At least two investigators, Gray (1964) and Wolfe (1969), have independently indicated that Itea pollen is present in Miocene sediments on the west coast of North America. Gray (1964) indicated that Itea pollen occurs in Miocene sediments of northern California, Idaho, Oregon, and Washington. Unfortunately, she did not identify the specific floras she studied. Wolfe (1969) indicated that Itea pollen occurs in early to middle Miocene sediments of the northwestern United States, somewhere in a region encompassing northern Nevada, Oregon, and Washington. Although Wolfe listed the floras he studied, it is not clear which yielded the fossil Itea pollen; all fifteen early to middle Miocene floras from this region had a megafossil component and only two definitely had an examined palynological component. One of the two, called "Rockville" by Wolfe (Succor or Sucker Creek) has a published palynoflora, but to date no record of fossil Itea pollen is documented (Graham, 1965, 1999). Among unpublished floras, Wolfe (1969) indicated only that Collawash, from the early Miocene of Oregon, had a pollen flora that had been examined. Therefore, it is also possible, but not definitively conclusive, that the Itea pollen documented by Wolfe (1969) was from Collawash. A "Pacific Northwest" placeholder datapoint has been included in the early-middle Miocene map roughly in the region of the Collawash flora to represent the occurrence of Itea pollen in that region during that time (Fig. 17, Appendix 2).

Fruits & Seeds

Only one taxon, Itea europaea Mai, represents fossil capsules and seeds morphologically consistent with those of Itea. Specimens of this taxon are known from European carpofloras, or deposits of fruits and seeds displaying three-dimensional preservation (Gee, 2005). Itea europaea is known from the early Miocene to late Pliocene, having been reported from Belarus, Bulgaria, Germany, Italy, Poland, and western Russia (Appendix 1). Itea europaea was erected by Mai in 1985, but was afterward figured by Pingen (1987) as an ericaceous capsule and seeds and by Dorofeev (1988) as Carpolithus sp. 1 capsules and seeds. Pingen (1996) later synonymized these fossils with L europaea. The holotype of Itea europaea was figured by Mai (1985, pl. 6, fig. 1) from lower Miocene sediments of the Berzdorf open-pit mine in the state of Saxony, Germany (Mai, 1999, 2000a, b; Czaja, 2003). It is currently held at the Museum fur Naturkunde in Berlin, Germany. Itea europaea is documented to co-occur with fossil Itea pollen in several floras during the same time interval, such as Kreuzau in the middle Miocene of Germany (Pingen, 1987; Ferguson et al., 1998), Melnik in the late Miocene of Bulgaria (Petrov & Drazheva-Stamatova, 1973; Mai & Palamarev, 1997; Ivanov, 2001a, b), and in the Gozdnica flora in the late Miocene to Pliocene of Poland (Sadowska, 1973, 1977; Dyjor & Sadowska, 1986; Mai, 2001a, b). In general, the geographic and stratigraphic range of L europaea shows substantial overlap with known occurrences of fossil Itea pollen in Europe (Appendix 1).

Capsules of I. europaea are 3.5 to 6 mm long and 1.4 to 5 mm wide, pedicellate, with a persistent calyx of five sepals and gynoecium of two superior carpels; the styles, where preserved, are persistent and appressed to one another or free. Dehiscence is septicidal. Seeds occur in two longitudinal rows per carpel and are about 1.3 to 2 mm long and 0.8 to 1.2 nun wide; they are ovoid to oblong and have a glossy testa covered by small polygonal cells (Mai, 1985, 2000a; Pingen, 1996; Mai & Palamarev 1997; Ferguson et al., 1998; Gumbel & Mai, 2004). Capsules of L europaea are consistent with capsules of extant Itea in being pedicellate; in having a persistent pentamerous calyx, two carpels, and two free styles upon dehiscence; in dehiscing septicidally; and in bearing multiple seeds (e.g., Spongberg, 1972; Jin & Ohba, 2001; Kubitzki, 2007a). Several authors have also reported that flowers of extant Itea have two rows of ovules per carpel (Spongberg, 1972; Jin & Ohba, 2001). This characteristic has been observed (Palmatier, 1942) and illustrated (Spongberg, 1972, fig. 8c) specifically for the North American species L virginica, which is probably the most thoroughly studied of the extant species. It is not, however, a universal feature within the genus. A drawing of the ovary of I. rhamnoides shows multiple ovules attached to the placenta in transverse section (Verdcourt, 1973, fig. 9; reproduced in Kubitzki, 2007a, fig. 70f), and figures of L yunnanensis show at least three rows of ovules in dissected carpels (Ge et al., 2002, figs. 27, 28).

Itea virginica is generally considered the closest living morphological and ecological proxy for L europaea (Mai, 1985, 2000a; Pingen, 1996; Mai & Palamarev 1997; Ferguson et al., 1998; Gumbel & Mai, 2004). In addition to the arrangement of the ovules in the carpels, the obvious morphological similarities between L europaea and L virginica include that both have hypogenous flowers bearing glossy ovoid to oblong seeds ornamented with numerous tiny, polygonal cells (Spongberg, 1972; Mai, 1985; Pingen, 1996). The texture of the testa is qualitatively more similar in L europaea and L virginica when compared to the ovoid seeds of L japonica (Mai, 1985) and I. rhamnoides (illustrated by Verdcourt, 1973, fig. 1.13).

Engler (1890, 1928) considered seed shape an important infrageneric character in grouping extant species of Itea. He subdivided Itea into two sections, Deciduae Engler and Sempervirentes Engler. Deciduae, also called section Itea (Spongberg, 1972), included the ovoid-seeded, deciduous species L virginica and I. japonica; Sempervirentes included the evergreen Asian species with fusiform seeds (Engler, 1890; 1928; see also Spongberg, 1972; Kubitzki, 2007a). This division was consistent until L rhamnoides was transferred into Itea, causing the classification to break down because Itea rhamnoides has more or less ovoid seeds but is evergreen, thus combining distinguishing traits of both of Engler's sections (Kubitzki, 2007a). Seed shape is, however, still a feature that is possibly phylogenetically significant. Seed morphology characteristics may help us to better understand the position of L europaea within the genus when an infrageneric phylogeny of extant Itea is available.

Flowers

Conwentz (1886) described the only floral taxon entirely morphologically consistent with the genus Itea, Adenanthemum iteoides Conwentz, from two specimens in Baltic amber; he only figured one of the specimens (Conwentz, 1886, plate 9, figs. 15-24). He thought the flowers comparable to those of Itea chinensis and I. virginica. Similarities between flowers of Adenanthemum Conwentz and Itea include that the flowers are hermaphroditic, pedicellate, have a pentamerous and actinomorphic perianth with distinct whorls of sepals and petals, a valvate corolla, a haplostemonous androecium with antesepalous stamens, dorsifixed anthers with introrse dehiscence, a superior ovary (ovary position is variable in Itea), two fused styles, and a single, capitate stigma (Conwentz, 1886; Kubitzki, 2007a). Conwentz (1886) may have been referring to the trichomes of Adenanthemum when he mentioned slight morphological differences distinguishing the two genera. As noted above, however, Itea has become a somewhat more morphologically variable taxon than it originally was as new species have been discovered and described (Kubitzki, 2007a). For instance, Adenanthemum has stalked glandular trichomes on its pedicel, calyx, and corolla (Conwentz, 1886); Itea glutinosa, first described in 1921, also has sessile or stalked glandular trichomes on its pedicel and calyx (Jin & Ohba, 2001) and I. rhamnoides has stalked glandular trichomes on its calyx (A1-Shammary & Gornall, 1994). Adenanthemum also has branched trichomes on its petals (Conwentz, 1886). While branched trichomes have not been reported on extant species, trichome anatomy should arguably be considered a distinguishing character at the specific rather than generic level for Itea. Thus, Adenanthemum is morphologically completely consistent with the genus Itea.

Some ambiguity exists, however, in the possible relationships between Adenanthemum and Itea. Pollen, the most diagnostic characteristic of Itea, is unknown for Adenanthemum. Dispersed fossil Itea pollen is documented from only one locality in the Eocene of Europe, from the upper Eocene of the Isle of Wight, UK (Gruas-Cavagnetto, 1976; Hooker et al., 2009). Furthermore, some Saxifragaceae and Ribes have flowers that are morphologically similar to those of Itea (e.g., Bensel & Palser, 1975a, b). Flowers of Saxifragaceae are most easily distinguished from those of Itea by the position of the anther attachment, basifixed with the filament inserted into a pit in Saxifragaceae, dorsifixed without a pit in Itea; and degree of fusion of the carpels, generally distally free with unfused styles and stigmas in Saxifragaceae, generally fused with consistently fused stigmas in Itea (Kubitzki, 2007a; Soltis, 2007). Adenanthemum is similar to Itea and differs from Saxifragaceae in the position of its anther attachment and degree of carpel fusion. Ribes flowers have a partially to fully inferior ovary and free stigmas (Weigend, 2007). Adenanthemum clearly differs from Ribes in having hypogenous flowers that have a single (or fused) stigma.

Baltic amber is found in the amber-bearing marine "blue earth" deposit of the Samland Peninsula and in blue-earth-derived accumulations in various areas on the edge of the Baltic Sea (Schlfiter, 1990). The in situ blue earth that is the most prolific producer of amber is assigned to the Upper Blue Earth Member of the Prussian Suite; it bears amber well-known for its organic inclusions, particularly of insects (Schluter, 1990; Kasinski & Kramarska, 2008). This sedimentary unit is thought to be a deltaic deposit derived from terrestrial amber-beating sediments eroded from the region of Fennoscandia (Schluter, 1990; Kasinski & Kramarska, 2008; Ritzkowski, 2008). The hypothesized source region for the blue earth sediments has been corroborated by K/ Ar dating of mica (Ritzkowski, 2008). The inclusion of Adenanthemum on the occurrence maps presents some difficulties because its source area is not precisely known. I have thus estimated its geographic position of origination on the Scandinavian Peninsula (Fig. 15, Appendix 2).

Dating the Baltic amber is also problematic. The Upper Blue Earth Member in which most of the Baltic amber occurs is now thought to be mid-to-upper Eocene in age. K/Ar dating of glauconite found within the blue earth has yielded an estimated age of 44.1 [+ or -] 1.1 Ma (Lutetian, lower middle Eocene) for the sediments (Ritzkowski, 1997). Perkovsky et al. (2007) have recently argued on the basis of other published evidence from glauconite dating and microfossil biostratigraphy that the Prussian Suite is upper Eocene in age. This interpretation is also preferred in a recent correlation study by Kasifiski and Kramarska (2008), who considered the Prussian Suite to straddle the Eocene-Oligocene boundary with the blue earth occurring just below the boundary. Because the amber found in blue earth deposits is redeposited from the region in which it originally formed, it is thought to be at least slightly older than the blue earth itself (Schluter, 1990; Weitschat & Wichard, 1998). Perkovsky et al. (2007) argued that the biota of the Baltic amber is transitional between known Northern Hemisphere early middle Eocene and Eocene-Oligocene boundary insect assemblages, thus suggesting that it belongs between these assemblages temporally. The combined evidence from these sources clearly suggests that the amber is no younger than late Eocene in age, although its maximum age is not well established. For the purposes of mapping, I have assigned it to the late Eocene (Fig. 15).

The specimens representing A. iteoides were originally held at the Westpreussische Provinzial Museum (WPM) (today, the Muzeum Archeologiczne Gdanska) in Danzig, Germany (today, Gdansk, Poland) (Kosmowska-Ceranowicz, 2001, 2008; Szadziewski & Sontag, 2008). All amber specimens once held at the museum--estimated minimally to number 13,151--were moved during World War II (Kosmowska-Cranowicz 2001; 2008). Most of these specimens have since been lost to science, with fewer than 2,500 pieces of the original collection currently recognized in the collections of the Muzeum Ziemi (Warsaw, Poland) and the Westpreussisches Landesmuseum MOnster (Munster, Germany) (Kosmowska-Ceranowicz, 2001; Hoffeins, 2008); several are also held at the Muzeum Archeologiczne Gdanska (Szadziewski & Sontag, 2008). Based on what has been written recently about the biography of Hugo Conwentz and the history of the amber collections of WPM (Kosmowska-Ceranowicz, 2001, 2008; Pielinska, 2008; Szadziewski & Sontag, 2008), it is probable that the Adenanthemum specimens were part of a large collection of amber assembled by Franz Anton Menge and left to the museum in 1880, the year WPM opened. This is because the Adenanthemum specimens belonged to the museum at the time they were originally published (Conwentz, 1886). Unfortunately, they have not been documented among specimens of the former WPM amber collection currently held in Warsaw (Kosmowska-Ceranowicz, 2001), and the MOnster and Gdansk collections are known to hold only specimens originally donated to WPM by Otto Helm in 1902 (Hoffeins, 2008; Szadziewski & Sontag, 2008). The Adenanthemum specimens are thus apparently not available for further study.

Conwentz (1886) did not designate either of the two specimens he mentioned as the type of A. iteoides, although he only illustrated what he considered to be the better specimen. If the specimens were still known to exist, they could be considered syntypes (International Code of Botanical Nomeclature 2006, Vienna Code, Article 9.4; McNeill et al., 2006). Because the available evidence indicates the original material has been lost, however, it is necessary to designate a neotype for this taxon, and illustrations are permissible neotypes (ICBN 2006, Articles 9.6 & 9.14; McNeill et al., 2006). The neotype is designated in Appendix 1.

Leaves

Two species representing fossil leaves and one additional collection of leaves have been attributed to Itea. The first of these to be described and published was I. faujassii (Unger) Meschinelli & Squinabol, reported from two floras: lower Oligocene sediments of Chiavon, Italy (Massalongo, 1859; Meschinelli & Squinabol, 1892), and upper Miocene sediments of Rochessauve, France (Faujas de Saint-Fond, 1803; Unger, 1845, 1850; Meschinelli & Squinabol, 1892; Grangeon, 1958). This taxon was illustrated by Faujas de Saint-Fond (1803, plate 56, fig. 3) from its type locality of Roehessauve; he considered it similar to extant Cedrela odorata L. Unger (1850) later assigned fossils of this type the binomial Cedrela faujasii Unger. When Meschinelli and Squinabol (1892) transferred this taxon from Cedrela P. Browne to Itea, they did not provide discussion to illuminate their reasons for doing so. The original specimen illustrated by Faujas de Saint-Fond (1803) is a simple, entire-margined, pinnately veined leaf; it may have brochidodromous secondary venation. The leaf as depicted appears to show at least three orders of veins. Leaves of extant species of Itea with entire margins have eucamptodromous to brochidodromous venation (Figs. 10 and 12), but not enough information is available about I. Jaujasii to make a meaningful comparison.

The second leaf taxon ascribed to Itea is L transsilvanica Petrescu & Givulescu, published in 1986 from Chuizbaia, Romania, and assigned a late Miocene age (Petrescu & Givulescu, 1986). The leaf is about 5 cm long, simple, pinnately veined, roughly ovate in shape, and has spinose teeth; only two orders of venation are preserved (Petrescu & Givuleseu, 1986). Originally assigned to Mahonia Nutt., the leaf was removed to Itea because it has eraspedodromous secondary venation. Petrescu and Givulescu (1986) considered it comparable to L ilicifolia Oliver, an extant species with spinose teeth (Jin & Ohba, 2001). Extant species of Itea may have a mixed venation pattern including craspedodromous veins; Itea species are, however, more generally characterized by semicraspedodromous, eucamptodromous, or brochidodromous secondary venation, or some mixture of these types of venation in the same lamina (Figs. 10, 11, 12, 13 and 14). Itea ilicifolia in particular has eucamptodromous to semicraspedodromous venation (Fig. 13). Thus, reevaluation of the affinities of this taxon is warranted.

The final collection of leaves attributed to Itea comes from the Republic flora of Washington, USA, part of the early Eocene Okanogan (alternative spelling, Okanagan) Highlands complex of floras from Washington and adjacent British Columbia, Canada (Greenwood et al., 2005). These leaves were originally briefly described by Wolfe and Wehr (1987) on the basis of three pieces representing two leaf apices from USGS locality 8428. Many more putative Itea leaves have since been discovered in the flora (Figs. 3, 4, 5, 6, 7, 8 and 9), numbering at least 68 specimens (Appendix 4) from the Boot Hill, Comer Lot, and Knob Hill Mine localities. These are unlobed, simple, petiolate leaves with pinnate venation; they vary in shape from elliptic to oblong, ovate, or obovate. The leaves are generally characterized by acuminate apices (although they can be straight, convex, or retuse) and convex to rounded to slightly cordate bases. Each has four to fifteen secondary veins, which are semicrapedodromous; sometimes, however, they are weakly so and appear nearly eucamptodromous. Tertiary venation is generally opposite percurrent. The leaves have five to six orders of venation and areolation is good. The putative Itea leaves at Republic are highly variable in their size, measuring from about 1.2 to 13 cm in length and 0.9 to 6.5 cm in width, with a length:width ratio of 1.09 to 2.95. All, however, share a characteristic serrulate margin with 8 to 15 (to 21) teeth per cm of leaf margin; the teeth are simple (one order of teeth present), regularly spaced, and possibly glandular.

The putative Itea leaves from Republic appear to be a good match for leaves belonging to some extant species of Itea. The serrulate margin in particular is similar to that of Itea virginica (Fig. 14), and was one of three primary characteristics Wolfe and Wehr (1987) originally cited in attributing the fossil leaves to Itea. It should be noted, however, that isolated leaves attributed to Itea are probably the most ambiguous form of evidence for the presence of Itea in the fossil record, since 1) many dicotyledonous plants bear simple, unlobed, pinnately-veined leaves, and 2) thus far, no single leaf character or suite of leaf characters have been suggested or discovered that are absolutely distinctive to the genus Itea when considered in the absence of floral, fruit, or pollen characters. The leaves of species within the genus Itea as a whole are, in fact, rather variable in size, shape, characteristics of the margin (entire or toothed, with teeth of various types), and angle of orientation of the secondary veins. Some examples are shown in (Figs. 10, 11, 12, 13 and 14).

Thus, the Republic leaves require additional morphometric study to further clarify the number of species they represent and additional morphological comparison with taxa in other families to confirm their affinities. Notably, Itea pollen is found in the Republic sediments in proportions of up to 5 % of the palynoflora (Moss et al., 2005); this suggests that Itea plants were growing on the site based on comparison to samples of pollen from modem habitats in or near which Itea occurs (Cohen, 1975). Smith (2011) recently reported four additional specimens of leaves cf. Itea from Falkland, another Okanogan Highlands flora. Itea pollen is also reported from Falkland, but in lower proportions than at Republic (Moss et al., 2005).

Ancient Distributions of Itea Based on its Fossil Record

A summary of the data in Appendices 1 and 2 and Figs. 15, 16, 17 and 18 is given below, with the most pertinent references cited. For complete references supporting fossil distributions, see Appendix 1. For floras/localities and coordinates, see Appendix 2. Worldwide distribution maps of fossil Itea occurrences have been published previously by Petrov and Drazheva-Stamatova (1973) and Mai (1985, fig. 4).

Eocene, 56 Ma-34 Ma (Fig. 15)

The Eocene distribution of iteoid fossils is both statigraphically and geographically disjointed. The first appearance of fossils attributed to Itea is from the lower Eocene (Ypresian) of the Pacific coast of North America, in the Okanogan Highlands floras of the northwestern United States and adjacent southwestern Canada, ranging from about 49-52 Ma (e.g., Greenwood et al., 2005; Moss et al., 2005). Following this cluster of occurrences, there is a significant temporal and geographic gap in the fossil record of Itea. The next oldest occurrences are in the late Eocene (Priabonian) of northwestern Europe (Conwentz, 1886; Gruas-Cavagnetto, 1976), over 10 million years later. As noted above, however, the lower limit on the age of the Baltic amber is not well constrained, so it may be more ancient than late Eocene.

Oligocene, 34 Ma-23 Ma (Fig. 16)

Itea has few occurrences in the Oligocene. In the early Oligocene (Rupelian), only three occurrences are known, one from Alaska, USA (Wahrhaflig et al. 1969), and two from central to southern Europe, including one record of the dubious leaf taxon Itea faujasii in Italy (Massalongo, 1859; Meschinelli & Squinabol, 1892) and one record of Iteapollis in Romania (Petrescu & Givulescu, 1986). In the late Oligocene (Chattian), occurrences are restricted to continental Europe.

Early-Middle Miocene, 23 Ma-11.6 Ma (Fig. 17)

The Miocene is perhaps the time of greatest geographic expansion of Itea in the fossil record, and of the greatest number of occurrences. In the early Miocene (Aquitanian and Burdigalian, 23-16 Ma), occurrences of fossil Itea are widespread and nearly circumboreal in distribution, although only five are from outside of continental Europe. Fossil Itea pollen is reported, once again, in the western part of North America, definitively from Alaska (Leopold & Liu, 1994) and more tentatively from the Pacific Northwest region of North America (Gray, 1964; Wolfe, 1969). Fossil 1tea pollen is also again documented from the British Isles; it is known from one locality on the Isle of Anglesey, although the age of this record is poorly established (Morawiecka et al., 1996; Walsh et al., 1996, 1999). For the first time, fossil pollen is reported from eastern North America (Traverse, 1955, 1994), and the only known report of fossil Itea pollen in eastern Asia is documented from the Russian province of Primorsky (Pavlyutkin & Petrenko, 2010). The remainder of occurrences are from continental Europe, where the fruit and seed taxon Bea europaea is first documented from several localities in Germany (Mai, 1985, 2000a, b), within the geographic vicinity of the known distribution of European pollen occurrences during this time interval.

Middle Miocene (Langhian and Serravallian, 11.6-16 Ma) occurrences of Itea fossils are concentrated exclusively in Europe and Anatolia, with the vast majority of reports of fossil Itea pollen and Itea europaea from a region encompassing eastern Germany and western Poland. During this time, Itea europaea is recorded in Tambov Province in the western part of the Russia (Dorofeev, 1988; Pingen, 1996), far to the east of other European occurrences of Itea fossils. The first record of fossil Itea pollen from western Asia also comes from this time interval, during which Iteapollis is identified from Turkey (Kayseri & Akgun, 2008).

Late Miocene-Pleistocene, 11.6 Ma-11.7 ka (Fig. 18)

During the late Miocene (Tortonian and Messinian, 11.6-5.5 Ma), there are no records of Itea fossils from eastern Asia. Records from Europe are again numerous, largely overlapping the latitudinal range of middle Miocene reports. At the southern limits of its range, there is one record of fossil Itea pollen from the Black Sea (Traverse, 1978; Traverse et al., 2005) and one report of Iteapollis from Turkey (Kayseri & Akgiin, 2008). The latter is the southernmost occurrence of an Itea fossil in the Eastern Hemisphere. Longitudinally, the distribution of fossil Itea occurrences shows a westward shift. In the east, there are no further reports from western Russia or Belarus, and in the west, fossil Itea pollen is first reported from Italy. There is also a report of the dubious leaf taxon Itea faujasii from France.

Pliocene records of Itea fossils are restricted to Europe and southeastern North America. In the southeastern United States, pollen has been found on Horn Island, off the southern coast of Mississippi in the Gulf of Mexico (Gohn et al., 1996). Pollen from the state of Georgia, USA, is also recorded from late Miocene or Pliocene sediments (Rich et al., 2002). In Europe, pollen, fruits, and seeds show a north-south geographic range similar to that in the late Miocene, although Itea apparently does not persist in the southeastern portion of its late Miocene range (Anatolia, Bulgaria, and the Black Sea) for very long into the Pliocene, if at all. Several reports of fossil Itea (pollen and Itea europaea) from Italy and Germany demonstrate that Itea was still present in Europe in the late Pliocene (Piacenzian) (Giimbel & Mai 2004; Bertini, 2002, 2010; Martinetto et al., 2007; Bertini & Martinetto, 2011). Martinetto (1999) and Martinetto et al. (2007) have suggested that Itea europaea disappears at the Piacenzian-Gelasian boundary in Italy, which is currently considered the boundary separating the Pliocene from the Pleistocene (Ogg, 2009).

Pollen records from the Pleistocene are restricted to the southeastern United States, overlapping with the range of extant Itea virginica (Figs. 2 and 18; Appendix 1). These reports include Itea pollen not assigned to a species and L virginica pollen documented from two sites in Georgia and one in Louisiana (Watts, 1971; 1973; Delcourt & Delcourt, 1977). No records were found for eastern Asia or Africa, despite occurrences of Itea in these regions today (Fig. 1).

Discussion

Origin and Age of Itea

The first appearance of Itea fossils is in the early Eocene Okanogon Highlands floras of western North America (e.g., Wolfe & Wehr, 1987; Greenwood et al., 2005; Moss et al., 2005). The Okanogan Highlands floras represent a well-known fossil Lagerstatte bearing multiple genera with connections to the modem floras of eastern North America and/or eastern Asia (Johnson, 1996; DeVore et al., 2005). The first documented occurrences of a number of extant genera--for instance, Carpinus L. (Betulaceae Gray), Corylus L. (Betulaceae), Fothergilla L. (Hamamelidaceae R. Br.), Neviusia A. Gray (Rosaceae Juss.), and Trochodendron Siebold & Zucc. (Trochodendraceae Eichler)--are also known from these floras (DeVore et al., 2005). Angiosperms are thought to have enjoyed their highest species origination and diversification rates during the early Eocene, possibly coinciding with a postCretaceous spike in the origination of new and evolutionarily significant floral structures (Crepet & Niklas, 2009; see also Crepet, 2008). This may have been driven in part by favorable conditions created by high global temperatures during the Early Eocene Climatic Optimum (Zachos et al., 2001). It is plausible, then, that the presence of Itea pollen--although the functional significance of its morphology, if any, is unknown--in these floras documents a very early occurrence of the taxon, perhaps correctly indicating its approximate time and geographic region of first occurrence.

There is a common historical biogeographic pattern wherein some angiosperm taxa endemic to or present in eastern Asia today have a more ancient record of occurrence in elsewhere in the Northern Hemisphere (Manchester, 1999; Tiffney & Manchester, 2001; Manchester et al., 2009). Based on this pattern, it has been suggested that eastern Asia may be a refugium for some formerly more widespread genera rather than a center of origin (Tiffney & Manchester, 2001; Manchester et al., 2009). Itea fits this pattern, appearing much later in eastern Asia than in North America or Europe. Because the fossil record is incomplete, however, there is always a chance that Itea originated elsewhere and its first appearance is simply indicative of the taxon occurring in the right setting to be preserved in the fossil record at that time. Itea has a record that is globally discontinuous temporally (especially in the Paleogene) and geographically, and new fossil discoveries could easily change our perspective on its origins and direction of dispersal. For instance, prior to 1987 when Wolfe and Wehr first suggested that Itea leaves were part of the Republic flora, the earliest occurrence of a fossil that might represent Itea was known from Europe (Conwentz, 1886). The record provided by the Baltic amber floral taxon Adenanthemum iteoides was, in fact, the oldest recognized for the majority of the twentieth century (e.g., Petrov & Drazheva-Stamatova, 1973; Mai, 1985).

Pollen is the most frequently recognized and diagnostic type of fossil representing Itea. Where it is recorded, Itea or Iteapollis fossil pollen is usually present in relatively low abundance of less than one percent to several percent of the palynoflora (e.g., Stuchlik, 1964; Traverse, 1955; Sadowska & Gila, 1991; Sadowska, 1992; Kohlman-Adamska, 1993). It can, however, reach or exceed proportions of 5 % (e.g., Bertoldi et al., 1994; Sadowska, 1977; Ivanov, 1996; Wacnik & Worobiec, 2001; Moss et al., 2005), with a maximum recorded abundance of 15 to 20 % in late Miocene sediments from Rosiori-Oradea, Romania (Petrescu & Givulescu, 1986). Empirical studies from modem floras, although few, can indicate roughly how likely it is that 1tea pollen will be present in the fossil record where Itea plants were present. For example, in their study of pollen rain in North Carolina, USA, Whitehead and Tan (1969) determined that the percentage of ltea pollen in lake-bottom sediments corresponded to the estimated basal area of Itea shrubs in the vegetation of the surrounding region. In that study, Itea ranged between 0.2 and 0.3 % of the pollen sampled when it was present. A study of airborne pollen in Taiwan also showed a low prevalence of Itea pollen. Less than 0.25 % of the airborne pollen counted from a trap placed near a forest in which 1tea parviflora occured was identified as Itea; Itea pollen was also only found in one of the 12 months during which sampling was performed (Chen& Huang, 2000). Higher proportions of pollen, 1 to 12 % of the palynoflora, were found by Cohen (1975) in peats underlying Nyssa-dominated areas within the Okefenokee Swamp (Georgia, USA); according to Cohen (1975: 126), Itea virginica is "occasional" in these areas. Peats sampled from other vegetation types represented in the Okefenokee Swamp had a proportion of less than 1% Itea pollen when this pollen type was present (Cohen, 1975). Itea is not a notable component of most of these other vegetation types (Cohen, 1973, 1975).

Taken together, these studies suggest that 1tea pollen is likely to be distinctly larger proportion of the palynoflora in deposits formed directly beneath vegetation in which Itea is a component and a much smaller proportion in sediments from adjacent areas, even when Itea plants are growing nearby. This is perhaps unsurprising, as Itea is thought to be insect-pollinated (Lieux, 1982). Thus, primary factors that likely determine the quality of the fossil record of Itea are preservation of deposits of the. proper age and habitat type to support 1tea. Within sediments that are temporally consistent with the possible presence of 1tea, sampling sufficient numbers of pollen grains in or near the proper environments is likely to be critical to detecting the presence of Itea. Since there are clearly sampling differences in the Cenozoic pollen record around the world, the degree of confidence that we can reasonably have in the fossil record of Itea is not equal everywhere. For instance, continental Europe is heavily sampled throughout much of the Cenozoic, and the bulk of reports of Itea fossils come from this region, Poland in particular (Appendix 1). Thus, the record of Itea in Europe may be considered reasonably complete, and it might be somewhat surprising to discover fossils that significantly extend the known stratigraphic range of the genus in that region. In contrast, the record in Asia, Africa, and North America is insufficiently documented. Lack of temporally and geographically suitable deposits (for instance, in eastern North America; Tiffney, 1985, 1994; Tiffney & Manchester, 2001), as well as possibly insufficient sampling (for instance, in eastern Asia; Manchester et al., 2009), are responsible for this.

Possibly supporting the hypothesis that Itea has a substantial missing record is the recent chronogram published by Jian et al. (2008, fig. 4) based on 28 taxa in Saxifragales and using a maximum age constraint and four minimum age constraints based on the saxifragalean fossil record. This chronogram suggests that crown-group Itea (represented by two terminals, Itea and Choristylis, in their analysis) originated in the midEocene, slightly post-dating the first appearance of fossil Itea pollen in the stratigraphic record. The split between Itea and Pterostemon, its sister genus, is dated to the latest Late Cretaceous. If these dates are accurate, they suggest that further fossils on the stem lineage between the Itea-Pterostemon node and the crown-group Itea node have yet to be discovered. It is, however, uncertain where along the stem lineage of Itea the morphological features that now characterize the genus and render it identifiable in the fossil record, such as its diagnostic diporate pollen, might have originated.

Biogeography of Itea

Itea has historically been considered one among many plant genera displaying an eastern North American-eastern Asian disjunct pattern (e.g., Li, 1952; Wu, 1983; Mai, 1985, 1995; Wen, 1999; Guo & Ricklefs, 2000; Ivanov, 2004). Many genera displaying this common disjunction are thought to be Northern Hemisphere "Tertiary relict genera" (Milne & Abbott 2002; Milne, 2006). These genera are believed to have been formerly distributed more widely across the Northern Hemisphere when the latitudinal temperature gradient was lower and high-latitude zones had warmer, more equable climates; they crossed between continents over the North Atlantic Land Bridge (NALB) in the northern Atlantic and the Bering Land Bridge (BLB) in the northern Pacific (e.g., Tiffney, 1985, 2008; Manchester, 1999; Wen, 1999; Tiffney & Manchester, 2001; Milne & Abbott, 2002; Milne, 2006). They exist today only in remnant ranges generally encompassing one or more of five noncontiguous regions: southeastern North America, western North America, northeastern Asia, southeastern Asia, and southwestern Eurasia (Milne & Abbott, 2002; Milne, 2006). Although Itea deviates from this pattern in occurring in a disjunct region in eastern Africa, its fossil record is congruent with that of a typical Northern Hemisphere relict. Itea currently inhabits or once inhabited all regions that are part of a Northern Hemisphere Tertiary relict pattern and evidence provided by fossils suggests that populations of Itea once had intercontinental connections over the Bering Land Bridge and North Atlantic Land Bridge for some duration of time between the Eocene and Miocene.

The fossil record of Itea on either side of Beringia is poor. The first appearance of Itea in the fossil record is in the early Eocene of western North America (Wolfe & Wehr, 1987; Moss et al., 2005), and fossil Itea pollen is reportedly widely distributed in the Pacific Northwest of North America later in the Cenozoic (Gray, 1964; Wahrhaftig et al., 1969; Wolfe, 1969; Leopold & Liu, 1994), although precise locality information for Neogene occurrences outside of Alaska is lacking. The diversity of Itea in eastern Asia--18 (Jin & Ohba, 2001; Liu, 2001; Ohba & Niu 2001) of the estimated 20 extant species live there--might suggest that this region is an ancient area of diversification for the genus (see discussion of this general topic in Tiffney & Manchester, 2001). There is, however, only one reported eastern Asian occurrence of Itea fossil pollen, from middle Miocene sediments of Primorsky, in a flora found near Vladivostok, Russia (Pavlyutkin & Petrenko, 2010).

It is clear that the shortest route between eastern Asia and North America for Itea would have been over the Bering Land Bridge, which served as a terrestrial connection between these regions until the latter part of the Neogene (Tiffney & Manchester, 2001). It has been argued, however, that some taxa may have migrated between North America and eastern Asia via Europe, since the high-latitude BLB may not have been a viable intercontinental route for evergreen broadleaved plants and thermophilic plants: winter light limitation and progressively cooler climatic conditions around the BLB beginning in the mid-Eocene may have created a hostile environment for these types of plants (Tiffney & Manchester, 2001; Manchester et al., 2009). Manchester et al. (2009), for instance, suggested that the deciduous thermophiles Sargentodoxa Rehder & E.H. Wilson (which occurs with Itea in the Brandon Lignite flora; Tiffney, 1993, 1994) and Tapiscia Oliv. may have invaded eastern Asia from North America through Europe, crossing continents over the NALB. Both genera are known from the Eocene of westem North America and have a Cenozoic fossil record in Europe (Manchester et al., 2009), paralleling the situation of Itea. In contrast to these genera, however, there is evidence that Itea reached high latitudes near the BLB: fossil Itea pollen has been documented in lower Oligocene and upper Miocene deposits in Alaska (Wahrhaftig et al., 1969; Leopold & Liu, 1994). Its presence in the early Miocene of eastern Asia (Pavlyutkin & Petrenko, 2010) further lends support to the hypothesis of a Beringial crossing. Thus, the presence of fossil Itea pollen in the Oligo-Miocene of Alaska and early Miocene of eastern Russia suggests it may have crossed the BLB from North America to eastern Asia in the latter Paleogene to early Neogene.

In the North Atlantic, fossil Itea pollen and iteoid flowers are first documented from upper Eocene sediments of the eastern Atlantic, in northwestern Europe (Conwentz, 1886; Gruas-Cavagnetto, 1976). The most logical inference is that Itea spread to Europe from North America over the NALB, because the first occurrences of Itea pollen in North America predate those in Europe. This hypothesis is bolstered by the directionality of invasion of Itea in Europe. After appearing in the northwest (Fig. 15), near the eastern end of the NALB where it is hypothesized to have connected to Europe (Tiffney, 1985), early Oligocene to early Miocene occurrences of Itea are found to the southeast in continental Europe (Figs. 16 and 17; Appendix 1). Itea finally reaches its most easterly and southerly occurrences in Europe and Anatolia in the middle to late Miocene (Figs. 17 and 18, Appendix 1), followed by range contraction and eventual extirpation, probably at the Pliocene-Pleistocene boundary (Martinetto, 1999; Martinetto et al., 2007; see also Appendix 1). The first appearance of fossil Itea pollen on the western side of the Atlantic is in the early Miocene Brandon Lignite of Vermont (Traverse, 1955, 1994). Because this is an isolated datapoint on the eastern coast of North America, where the Cenozoic record is poor (Tiffney, 1985, 1994; Tiffney & Manchester, 2001), it is difficult to incorporate into a biogeographic scenario. It is thus unknown when Itea may have first reached the more southerly portions of the western Atlantic region, although it must have reached the northern part of this region in the Eocene if it made the crossing from North America to Europe.

The present climate of Vermont, USA, is too cool to support Itea and other thermophilic genera found within the Brandon Lignite, many of which inhabit suitable areas of southeastern North America and eastern Asia today (Tiffney, 1994). It is thus logical that Itea achieved its present distribution in southeastern North America simply by migrating down the Atlantic coast during the Neogene (Figs. 17 and 18), when global climate cooled and, finally, the Northern Hemisphere experienced glaciation beginning in the Pliocene (Zachos et al., 2001). Watts (1970) suggested that the extant taxon Itea virginica is of a group of plants represented best on the southeastern Coastal Plain--the "Coastal Plain disjuncts"--that achieved a wide but sporadic inland distribution throughout southeastern North America by invading patches of suitable habitat to the north and west following glacial retreat in the Quaternary. This distribution, where L virginica is generally found in moist to saturated substrates to standing water in wooded areas throughout the southeastern United States (e.g., Beaven & Oosting, 1939; Spongberg, 1972; Cohen, 1973; Cranfill, 1991; Brooks et al., 1993; Anderson et al., 2009; Morin, 2009), indicates that L virginica is tracking a suitable habitat type under a suitable climatic regime. This may prove useful in understanding the spread and eventual extinction ofltea in Europe, since L virginica is likely not only a morphological but also an ecological proxy for Itea europaea and the plant or plants represented by fossil Itea pollen in the Cenozoic of Europe. The fossils are thought to represent plants that grew in habitats with regularly or continuously saturated substrates, such as swamps and floodplains (Mai, 2001b; Ivanov, 2001a; Hofmann & Zetter, 2005). Itea virginica is also known at least sometimes to be an understory shrub in Taxodium Rich.-dominated swamp forests (e.g., Beaven & Oosting, 1939; Cohen, 1973; Anderson et al., 2009); likewise, 1tea europaea and fossil 1tea pollen from Europe are also known to occur in floras where the taxodiaceous genera Taxodium or Glyptostrobus Endl. are well-represented in the paleoflora (e.g., Bertoldi et al., 1994; Ivanov, 2001a; Mai, 2001b; Hofmann & Zetter, 2005; Martinetto et al., 2007). There is an especially interesting occurrence in a sample from the Fornace Filippi locality (in Appendices 1 & 2, cited as part of the Sarzana flora) in the late Miocene of northwestern Italy, where 1tea pollen makes up over 12 % of a palynoflora dominated by Taxodium-type pollen (>34 %). In this instance, Bertoldi et al. (1994: 166) suggested that Itea was "an in-loco plant" given the proportion of its pollen in the flora, a hypothesis supported by studies of modern habitats that support L virginica in southeastern North America (Cohen, 1975).

Verdcourt (1973), presumably hypothesizing that the African species L rhamnoides originated in southern Africa, suggested that L rhamnoides spread south-to-north following mountain ranges to achieve its present distribution in eastern Africa. Given the geographic proximity of its present distribution to Europe, L rhamnoides is likely to have arisen from the Cenozoic European populations of Itea; the fossil record thus suggests that the likely migration pattern was north-to-south. This inferred Cenozoic European-recent African connection is interesting in light of recent studies suggesting that New World-African disjunct patterns between species or clades in some extant taxa probably arose from the ancient connection between the two regions via the NALB and Europe (Lavin et al., 2000; Davis et al., 2002; Xiang et al., 2005, 2006; Milne, 2006; Erkens et al., 2009). Given the African-New World pattern emerging in recent studies, it might be predicted that L virginica and L rhamnoides are sister taxa. This would fit the evidence provided by the morphological and ecological similarities between L virginica and the extinct L europaea, perhaps the precursor to L rhamnoides.

The timing of this possible North American-African split in Itea is ambiguous on the basis of the fossil evidence. Occurrence data indicate that Itea likely first crossed the NALB in the Eocene (Fig. 15), and reports of fossil Itea pollen on either side of the proposed route of the NALB occur at least as recently as the early Miocene (Traverse, 1955, 1994; Walsh et al., 1996). Our evolving understanding of the operation of the NALB now suggests it may have functioned as a transatlantic connection relatively late into the Neogene for some taxa (Tiffney, 2008; Denk et al., 2010), making the possibility of a Miocene transatlantic connection between populations of Itea plausible. Only further study of the phylogenetic relationships of the extant taxa within the genus will help to clarify the timing and order of isolation of populations of Itea occupying different refugia today.

A Note on Using Itea Fossils as Calibration Points

Parham et al. (2012) recently outlined a series of "best practices" for identifying fossils to use as calibration points in divergence dating studies and are here cited as a guide for the evaluation of the suitableness of Itea fossils for use as calibration points. Given the "exceedingly rare" (Erdtman, 1955: 407) pollen morphology of Itea, as well as its relatively good fossil record, fossil Itea pollen should be well-suited for use as a calibration point in calculating divergence times on phylogenetic trees generated from molecular sequence data. Because the evolution of the pollen is a transformation optimized on the stem of the Itea clade (Hermsen et al., 2006), and because it may be unique to (autapomorphic for) Itea among the angiosperms, it is best used to calibrate the node representing the most recent common ancestor of Itea and Pterostemon on phylogenies of extant taxa (Fig. 1). The oldest occurrences of Itea pollen are in the early Eocene Okanogan Highlands floras of the Pacific Northwest of North America (Moss et al., 2005). The Okanogan Highlands floras in which fossil Itea pollen is documented all have associated radiometric dates from Ar/Ar, K/Ar, or U/Pb analyses, placing them between about 49 and 52 Ma (Church et al., 1979, recalculated from Church, 1975, using an updated constant; Ewing, 1981, recalculated from Hills & Baadsgaard, 1967, using an updated constant; Wolfe et al., 2003; Dilhoffet al., 2005; Moss et al., 2005). None of this pollen, however, is figured, so the diagnostic characters of the Okanogan Highlands material cannot be confirmed. The characters of fossil Itea pollen in general are, however, well-documented (see above).

The oldest figured pollen grain is upper Eocene in age (Gruas-Cavagnetto, 1976, pl. 9, fig. 15). Originally assigned to the Middle Headon Beds (Gruas-Cavagnetto, 1976), more recent stratigraphic nomenclature places this fossil in the Colwell Bay Member of the Headon Hill Formation (King, 2006). This stratigraphic unit is upper Eocene in age, its upper boundary being about 35.7 Ma based on biostratigraphy, magnetostratigraphy, and correlation (Hooker et al., 2009). Given that it is a great deal younger than the Okanogan Highlands occurrences, however, the age associated with the Colwell Bay Member fossil is not ideally suited for use as a calibration.

Because current evidence suggests that the original specimens are no longer available for study and given the uncertainty surrounding the age of the Baltic amber (discussed above), Adenanthemum iteoides should not be used as a calibration point. Itea europaea may prove suitable as a calibration point following further investigation of its likely phylogenetic position within the genus Itea. Application of fossil leaves attributed to 1tea as calibration points should be avoided unless or until diagnostic characters are discovered for the leaves of the genus.

Conclusions

The fossil record of Itea begins about 50 million years ago in the Pacific Northwest of North America, and is sporadic throughout the Paleogene, when fossil occurrences are recorded only from North America and Europe. The fossil record suggests that Itea invaded Europe over the North Atlantic Land Bridge no later than the late Eocene and spread from the northwest toward the south and east. The genus reached a peak of abundance in the middlelate Miocene of Europe, coinciding with a relatively warm period during the Cenozoic. Itea declined in Europe during the Pliocene, becoming extirpated from the European-western Asian region by the Pleistocene. Fossil Itea pollen is found in Alaska in the early Oligocene and early Miocene and in eastern Asia in the early Miocene, providing evidence Itea could have crossed the Bering Land Bridge from North America to eastern Asia. Evidence provided by the distribution of Itea fossils in Europe suggests that Itea reached eastern Africa from Europe, exhibiting a recently recognized New World-African disjunction pattern that is thought to have arisen via Cenozoic connections between these two regions through the NALB and Europe. Given the ecological and morphological similarities of I. virginica and European Itea fossils and the evidence suggesting that I. rhamnoides most likely arose from extinct Cenozoic European Itea, one might predict that I. rhamnoides and I. virginica are sister taxa showing a transatlantic disjunction. This hypothesis, however, remains to be tested phylogenetically.

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. Adenanthemum iteoides, late Eocene or older, Baltic Amber (thought to have originated in Fennoscandia). Conwentz (1886). Add. Occ. Refs.: Schluter (1990), Ritzkowski (1997, 2008), Perkovsky et al. (2007), Kasinski 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. Itea europaea, Miocene, locality unknown (Belarus). Mai (1985), Dorofeev (1988), Pingen (1996). Not mapped.

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

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

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

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

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

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

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

11. 1tea europaea, late Miocene, Baldevo (Blagoevgrad, Bulgaria). Mai & Palamarev (1997), Palamarev et al. (2005).

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

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

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

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

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

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

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

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

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

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

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

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

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

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

3. Iteafaujasii, 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. Itea faujasii, late Miocene, Rochessauve (Rhrne-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. Oct. Ref.: Grangeon (1958). Holotype: Faujas de Saint-Fond (1803), pl. 56, fig. 3.

5. Itea transsilvaniea, late Miocene, Chiuzbaia (Maramures, Romania). Petrescu and Givulescu (1986). l-Iolotype: Petrescu and Givulescu (1986), fig. 3.

Pollen

Eocene

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

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

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

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. 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. Iteapollis angustiporatus, early Oligocene, Tamasa (Salag, Romania). Petrescu and Givulescu (1986) as Iteapollenites angustiporatus.

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

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

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

Miocene, Subdivision Unknown

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. 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 Ziembinska and Niklewski (1966), recognized as Itea by Petrov and Drazheva-Stamatova (1973) and Sadowska (1973), recognized as equivalent to Iteapollis angustiporatus by Ziembinska-Tworzydlo (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. Iteapollis angustiporatus, early-middle Miocene, Gierlachowo (Greater Poland, Poland). Ziembinska-Tworzydlo (1974). Add Occ. Refs.: Piwocki and Ziembinska-Tworzydlo (1997).

3. Iteapollis angustiporatus, early-middle Miocene, Goiqbin Stary (Greater Poland, Poland). Ziembiflska-Tworzydlo (1974), Ziembinska-Tworzydlo & Wa2yflska (1981), Ziembinska-Tworzydlo et al. (1994). Add. Occ. Ref.: Piwocki and Ziembiflska-Tworzydlo (1997).

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

5. cf. Rea, early-middle Miocene, Niedzwiedzice (Lower Silesia, Poland). Ziembinska and Niklewski (1966).

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

7. Iteapollis angustiporatus, early-middle Miocene, Nowa Wies (Lower Silesia, Poland). Ziembinska-Tworzydlo (1974). Add. Occ. Ref.: Piwocki and Ziembinska-Tworzydlo (1997).

8. Iteapollis angustiporatus, early Miocene, Oder II (Bavaria, Germany). ThielePfeiffer (1980).

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

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

11. Iteapollis angustiporatus, early-middle Miocene, Slepuchowo (Greater Poland, Poland). Ziembinska-Tworzydlo (1974). Add. Occ. Ref.: Piwocki and Ziembiflska-Tworzydlo (1997).

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

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

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

15. Iteapollis angustiporatus, early Miocene, Zaghid-Hida (Salag, Romania). Petrescu and Givulescu (1986) as Iteapollenites angustiporatus.

middle Miocene (or range beginning in the middle Miocene)

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

2. 1tea, middle Miocene, Deleina (Vidin, Bulgaria). Ivanov et al. (2002a), Ivanov (2004). Add. Occ. Ref.: Ivanov et al. (2002b).

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

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

5. 1tea, middle Miocene, GSW/6 (Lubusz, Poland). Sadowska (1992). Add. Occ. Ref.: Dyjor (1992).

6. Iteapollis angustiporatus, middle Miocene, Incescu-Zambal (Corum, Turkey). Kayseri and Akgun (2008).

7. Itea, middle Miocene, Jarosz6w (Lower Silesia, Poland). Sadowska (1977). Add. Occ. Ref.: Dyjor and Sadowska (1986).

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

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

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

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

12. Iteapollis angustiporatus, middle Miocene, Konin (Greater Poland, Poland). Ziembinska-Tworzydlo et al. (1994).

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

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

15. Iteapollis angustiporatus, middle Miocene, Krosinko (Greater Poland, Poland). Ziembinska-Yworzydto (1974). Add. Occ. Ref.: Piwocki and ZiembinskaYworzydto (1997).

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

18. Itea, middle Miocene, gojowice (Lower Silesia, Poland). Sadowska (1977). Add. Oec. Ref.: Dyjor and Sadowska (1986).

19. Iteapollis, middle Miocene, Lubst6w (Greater Poland, Poland). Durska (2008).

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

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

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

23. Iteapollis angustiporatus, middle Miocene, Mosina (Greater Poland, Poland). Ziembinska-Tworzydlo (1974). Add. Oce. Ref.: Piwocki and ZiembinskaTworzydlo (1997).

24. Iteapollis angustiporatus, middle Miocene, Oczkowice (Greater Poland, Poland). Ziembinska-Tworzydlo (1974). Add. Oce. Ref.: Piwocki and Ziembinska-Tworzydlo (1997).

25. Iteapollis angustiporatus, middle Miocene, Oder I (Bavaria, Germany). ThielePfeiffer (1980).

26. Itea, middle Miocene, Patnow (Greater Poland, Poland). Sadowska and Giza (1991).

27. Itea, middle Miocene, Rusz6w (Lower Silesia, Poland). Sadowska (1977). Add. Occ. Ref.: Dyjor and Sadowska (1986).

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 Ziembinska-Tworzydlo (1974), synonymized with Iteapollis angustiporatus by Thiele-Pfeiffer (1980). See also Ziembinska-Tworzydlo et al. (1994, pl. 8, fig. 12), pollen refigured from Stuchlik (1964, pl. 11, fig. 8) assigned to Iteapollis angustiporatus.

29. Iteapollis angustiporatus, middle Miocene, Spreetal (Saxony, Germany). Schneider (1965) as Psilodiporites angustiporatus, synonymized with Iteapollis angustiporatus by Ziembinska-Tworzyd]o (1974). Holotype: Schneider (1965), pl. 1, fig. 10.

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

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

33. Itea, middle Miocene, Zar6w (Lower Silesia, Poland). Sadowska (1977). Add. Occ. Ref.: Dyjor and Sadowska (1986).

late Miocene (or range beginning in the late Miocene)

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

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

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 DrazhevaStamatova (1973, figs. 1-2) synonymized with Iteapollis angustiporatus by Thiele-Pfeiffer (1980).

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

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

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

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

8. Itea, late Miocene-Pliocene, Gozdnica (Lubusz, Poland). Sadowska (1997). Add. Occ. Refs.: Dyjor and Sadowska (1986), Dyjor (1992).

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

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

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

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 Reapollis angustiporatus by Thiele-Pfeiffer (1980).

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

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

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

16. Itea, late Miocene-Pliocene, Paczk6w (Opolskie, Poland). Sadowska (1977). Add. Occ. Ref.: Dyjor and Sadowska (1986).

17. Itea, late Miocene-Pliocene, Poznafl (Greater Poland, Poland). Trod and Sadowska (2006).

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

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

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

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

Pliocene

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

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

4. Iteapollis angustiporatus, Pliocene, Oldenswort (Schleswig-Holstein, Germany). Menke (1975, 1976) as 1tea. Pollen figured by Menke (1976, pl. 16, figs. 12-15) synonymized with Iteapollis angustiporatus by ThielePfeiffer (1980).

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

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

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

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

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

Pleistocene (or Range Beginning in Pleistocene)

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

2. 1tea, Pleistocene-Holocene, Lake Louise (Georgia, USA). Watts (1971, 1980).

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

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

1. 1tea virginica, Holocene (ca. 10,000 years b.p. to recent), Anderson Pond (Tennessee, USA). Delcourt (1979).

2. 1tea, Holocene (ca. 3,500--7,300 years b.p.), B.L. Bigbee (Mississippi, USA). Whitehead and Sheehan (1985).

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

4. 1tea virginica, Holocene (ca. 7,300 years b.p. to recent), Jackson Pond (Kentucky, USA). Wilkins et al. (1991).

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

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

7. 1tea, Holocene (Georgia, USA), Quicksand Pond. Watts (1970).

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,
IEo late Eocene, e0l early Oligocene, 101 late Oligocene,
eMio early Miocene, inMio middle Miocene, IMio late Miocene,
Plio Pliocene, Plei Pleistocene, Hol Holocene

Locality             Organs   Age         Longitude     Latitude

Achldorf             p        IMio        12.367000     48.433000
Ahrensburg           p        Plio        10.239803     53.675712
Badcrsdorf           p        IMio        16.373244     47.201788
Baldevo              I's      IMio        23.770104     41.628960
Baltic Amber         fl       lEo         16.663265     62.084635
Belfeld              fs       Plio        6.120920      51.313652
Bcli Breg Mine       p        IMio-Plio   22.853242     42.868756
Berzdorf             fs       eMio        14.959409     51.090627
Boschi di Barbania   fs       Plio        7.606624      45.287281
Brandon Lignite      p        chio        -73.050000    43.833333
Bukcha               p        mMio        27.666667     51.766667
Chelnovskih          fs       mMio        40.949657     52.864795
Cherasco             p        IMio        7.858252      44.651654
Chiavon              1        c01         11.547988     45.740279
Chiuzbaia            I        IMio        23.684532     47.702100
Colwell Bay          p        lEo         -1.537032     50.689820
Czamy Dunajcc        p        IMio        19.848337     49.439202
Darova-Lugoj         p        IMio        21.916100     45.688011
Deleina              p        mMio        22.633000     44.050000
Drenovets            p        mMio-IMio   22.983000     43.700000
DSDP Site 380A       p        IMio        29.613700     42.099000
Dunarobba            fs       Plio        12.452055     42.671722
Enspel               p        101         7.895882      50.613230
Falkland             1, p     cEo         -119.628000   50.516000
Front Canavese       fs       Plio        7.677222      45.273333
Gierlachowo          p        emio-mMio   16.860544     51.983837
Golgbin Stary        p        emio-mMio   16.792892     52.081437
Gozdnica             fs, p    IMio-Plio   15.068595     51.440308
Gozdnica-Stanislaw   p        IMio        15.061816     51.443134
Green Pond           p        Plei        -84.916976    34.269499
Gressk               p        101         27.493051     53.169486
GSW/3                p        mMio        15.109665     51.440475
GSW/o                p        mMio        15.110938     51.430756
Hambach              fs       IMio        6.502027      50.910703
Hartau               fs       cmio        14.814386     50.862104
Hat Creek            p        cEo         -121.629986   50.746197
Hom Island           p        Plio        -88.668848    30.235070
Incescu-Zambal       p        mMio        34.802158     40.974060
Jaroszow             p        mMio        16.419308     50.987868
Jerzmanowa           p        mMio        16.044937     51.596210
Karolewo-Dabki 3     p        mMio        17.292765     53.137532
Karolewo-Dabki 4     p        mMio        17.331612     53.138630
Katina               p        IMio-Plio   23.236944     42.833611
Kauschc-Klara II     fs       IMio        14.207310     51.606524
Kleinleipisch        fs       mMio        13.750000     51.517000
Klettwitz            fs       mMio        13.900000     51.550000
Komomiki             p        mMio        16.412095     51.173190
Konin                p        mMio        18.251073     52.223033
Kosztowo             p        mMio        17.194059     53.160567
Kreuzau              fs, p    mMio        6.505627      50.747826
Krosinko             p        mMio        16.801286     52.231243
Lake Louise          p        Plci-Hol    -83.258333    30.725000
Lavrovo              fs       mMio        40.939190     52.339161
Legnica              p        mMio        16.155323     51.207007
Liszkowo             p        mMio        17.321185     53.229546
Lojowice             p        mMio        17.211985     50.736850
Lubstow              p        mMio        18.477237     52.343789
Lukasberg            p        mMio-IMio   13.545470     48.089090
Maccarone            p        IMio        13.111164     43.404234
Makresh              p        mMio        22.667000     43.767000
McAbee               p        cEo         -121.142800   50.796967
Melnik               fs, p    IMio        23.392757     41.522761
Mirostowicc          p        mMio        15.097869     51.575862
Mlamolovo            p        101         23.024122     42.343132
Mosina               p        mMio        16.847054     52.245407
Nenana Coal Field    p        eMio        -149.000000   64.000000
Niedzwiedzice        p        eMio-mMio   16.021661     51.285443
Nizhino              p        eMio        131.783333    43.483333
Nowa Wio             p        eMio-mMio   16.444180     51.781136
Nysa                 p        IMio        17.335461     50.480956
Oberzella            fs       Plio        10.038722     50.838635
Oczkowicc            p        mMio        17.038281     51.692216
Oder I               p        mMio        12.171054     49.282757
Oder II              p        eMio        12.165713     49.284688
Ohoopee Dune Field   p        IMio-Plio   -82.381952    32.440668
Oldenswort           p        Plio        8.859316      54.394329
Opole                p        IMio        17.924438     50.670428
Pacific Northwest    p        eMio-mMio   -122.329020   45.199652
Paczkow              p        IMio-Plio   17.005726     50.463565
P4tnow               p        mMio        18.250000     52.316667
Plessa               fs       mMio        13.619050     51.464372
Poznah               p        IMio-Plio   16.925168     52.406374
Pranzalito           fs       Plio        7.810783      45.408785
Reichwalde           fs       eMio        14.663012     51.380992
Republic             1, p     cEo         -118.737807   48.648218
Rex Creek            p        0.00E+00    -148.666667   64.083333
Rittstcig            p        eMio-mMio   13.367000     48.583000
Rochcssauve          I        IMio        4.626456      44.679669
Rosiori-Oradea       p        IMio        21.941509     47.057919
Ruszow               p        mMio        15.182834     51.400763
Rypin                p        mMio        19.409410     53.065711
Santa Barbara        p        Plio        11.484135     43.567139
Sarzana              p        IMio-Plio   9.959704      44.112518
Slepuchowo           p        eMio-mMio   16.722155     52.595199
Smolyarka            p        101, mMio   24.979293     52.530864
Sosnica              p        Plio        16.799711     51.026241
Sprectal             p        mMio        14.371498     51.490779
Stura di Lanzo       fs       Plio        7.563611      45.215278
Tamasa               p        col         23.189271     46.945639
Trwyn y Parc         p        cmio-mMio   -4.451288     53.418682
Tuplicc              p        mMio        14.828686     51.675721
Ustronic             p        emio-mMio   16.285223     51.432668
Vad-Borod Basin      p        mMio        22.625000     47.000001
Vasiltcpc            p        IMio        36.075031     39.185200
Wakcndarf-II         p        mMio        10.081172     53.782161
Wclzow               fs       IMio        14.172836     51.574483
Wilcox Bluff         p        Plci        -91.387105    30.875278
Wirczyn              p        emio-mMio   16.113436     51.302021
Wo1ka Ligczowska     p        Plio        20.583333     51.583333
Wroclaw              p        IMio        17.038538     51.107885
Zaghid-Hida          p        cmio        23.313709     47.062597
Zarow                p        mMio        16.494740     50.940760


Appendix 3

Samples of ]tea europaea examined at the Museum fur 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; R6merkeller 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 (lot. 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 36871 A, 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 (lot. 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 01-1204, 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 99-184A, B. USNM: Unnumbered (Loc. USGS 8428).

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

DOI 10.1007/s12229-012-9114-3

Acknowledgments

This project includes information from the second chapter of my disscrtation (Hermsen, 2005) as well as much new research done since 2005.1 would especially like to rccognize 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 l 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[R] was used as a source for extant taxon authorities.

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Elizabeth J. Hermsen (1,2)

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

(2) Author for correspondence; e-mail: hcrrnscn@ohio.edu

Table 1 Extant species of flea based on recently published
descriptions and revisions by Jin and Ohba  (2001), Liu (2001),
Siddiqi (2005), Kubitzki (2007a) and Morin (2009)

Species                          Distribution

Ilea amoena Chun                 China (Guangdong, Guangxi)
Ilea chinensis Hook. & Am.       Bhutan, southeastern China,
                                 northeastern India, Laos, Myanmar,
                                 Thailand, Vietnam
Ilea coriacea Y.C. Wu            southeastern China (incl. Hainan)
Ilea glutinosa Hand.-Mazz,       southeastern China
Ilea ilicilolia Oliv.            southeastern China
Ilea indochinensis Mcrr.         southeastern China, Vietnam
Itea japonica Oliv.              Japan (Honshu, Shikoku, Kyushu)
Ilea kiukiangensis C.C. Huang    China (Tibet/Xizang, Yunnan)
& S.C. Huang ex H. Chuang
Ilea kwangsiensis H.T. Chang     China (Guangxi)
Ilea macroplrylla Wall.          Bhutan, China (Guangxi, Hainan,
                                 Yunnan), northeastern India, Indonesia
                                 (Borneo, Java, Sumatra), Laos,
                                 Myanmar, the Philippines (Luzon),
                                 Thailand, Vietnam
Ilea nutans Roylc                northern India, northern Pakistan
Ilea oldhandi C.K. Schncid.      Japan (Ryukyu Islands), Taiwan
Ilea omeiensis C.K. Schncid.     southeastern China
Ilea parviflora Hemsl.           Taiwan

Ilea rharnnoides (Harv.)         Burundi, Democratic Republic of the
Kubitzki                         Congo, Lesotho, Malawi, Mozambique,
                                 South Africa, Swaziland, Tanzania,
                                 Uganda, Zambia, Zimbabwe
Ilea riparia Collett & Hemsl.    China (Yunnan), Myanmar, Thailand
Ilea tenuinervia S.Y. Liu        China (Guangxi)
Ilea vhginica L.                 southeastern USA
Itea yangchuensis S.Y. Jin       China (Guangdong)
Ilea yunnanensis Franch.         southeastern China

Sources for distribution data: Africa, Verdcourt (1973),
Dowsctt-Lemaire (1985); China, Japan, and Taiwan, Jin and Ohba
(2001), Liu (2001), Ohba and Niu (2001); India, Clarke (1879),
Merrill (1906, 1921), Jamir et al. (2006); Indonesia, Merrill
(1921); Pakistan, Parker (1924), Siddiqi (2005); The Philippines,
Merrill (1906); USA (Morin, 2009). 1. macroplrylla in Laos
documented by Newman ct al. (2007). Remaining countries listed for
l. chinensis and 1. rnacrophylla, An and Ohba (2001). Chinese
provinces listed if three or less for a given species
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Date:Mar 1, 2013
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