Pl. Syst. Evol. 266: 45–57 (2007) Plant Systematics DOI 10.1007/s00606-007-0540-3 and Evolution Printed in The Netherlands

A brief review of the fossil history of the family Rosaceae with a focus on the Eocene Okanogan Highlands of eastern Washington State, USA, and British Columbia, Canada

M. L. DeVore1 and K. B. Pigg2

1Department of Biological and Environmental Sciences, Georgia College & State University, Milledgeville, GA, USA 2School of Life Sciences, Arizona State University, Tempe, AZ, USA

Received January 16, 2006; accepted August 17, 2006 Published online: June 28, 2007 Springer-Verlag 2007

Abstract. Many of the oldest definitive members of temperate regions (Heywood 1993). Members the Rosaceae are present in the Eocene upland floras of the Rosaceae have radiated into a wide of the Okanogan Highlands of northeastern Wash- variety of environments ranging from mesic to ington State and British Columbia, Canada. Over a xeric communities and are elements of boreal dozen rosaceous taxa representing extant and extinct and tundra ecosystems. No doubt one of the genera of all four traditionally recognized subfam- driving forces for the Rosaceae’s success is the ilies are known from flowers, fruits, wood, pollen, presence of agamospermy, hybridization, poly- and especially leaves. The complexity seen in Eocene Rosaceae suggests that hybridization and poly- ploidy and vegetative reproduction within the ploidy may have played a pivotal role in the early family. All of these microevolutionary pro- evolution of the family. Increased species diversity cesses contribute to generating novel genetic and the first appearance of additional modern taxa combinations capable of colonizing and per- occur during the Late Paleogene in North America sisting in new, open habitats. and Europe. The Rosaceae become increasingly It comes as no surprise that the Rosaceae important components of fossil floras during the captured the eye of G. Ledyard Stebbins, Jr. Neogene, with taxa adapted to many habitats. Stebbins believed that the subfamily Maloideae (also called the Pyroideae) are allopolyploids Key words: Eocene, fossil, Okanogan Highlands, that combined adaptive gene complexes de- Prunus , Rosaceae, temperate floras, Tertiary. rived from ancient lineages of Prunoideae with those from Spiraeoideae (Stebbins 1950, Evans and Campbell 2002). Stebbins suspected that Introduction these new complexes had high fitness in new The Rosaceae is a large family of approxi- habitats that arose during the middle and late mately 122 genera and 3,370 species of trees, Tertiary. One thing was missing in order for shrubs and herbs of worldwide distribution, Stebbins’ concept to be supported: the fossil with its maximum development in north evidence. 46 M. L. DeVore and K. B. Pigg: Fossil history of Rosaceae

In October, 1994, Stebbins responded to a Dieter Mai, Zlatko Kvacˇ ek and Harald Wal- letter sent to him by Wes Wehr that was ther for more detailed discussion (e.g. Mai and accompanied by a list of rosaceous taxa from Walther 1978; Mai 1984, 1995; Kvacˇ ek and Okanogan Highlands floras (DeVore and Pigg Walther 1998, 2001, 2004; Kvacˇ ek et al. 2004). 2005; Wesley Wehr Papers at the University of This is by no means the end of the story. Washington Libraries). Wehr, who died in Numerous rosaceous fossils are nestled within April, 2004, was Affiliate Curator of Paleobo- museum cabinets in need of further description. tany at the Burke Museum on the University of Therefore, this is the story as it stands today Washington campus. In addition to his training and our understanding of the fossil record for in music composition, contributions to the Rosaceae will no doubt become clearer once Northwest School of art, and involvement in undescribed taxa are critically examined and Seattle’s literary scene, Wehr cared for the more widely compared. Burke’s extensive collections of Tertiary plant The Eocene radiation. The most significant and insect fossils from the Pacific Northwest early representation of fossil Rosaceae occurs (Archibald et al. 2005). Stebbins commented in in the early and middle Eocene floras of central his response to Wehr that ‘‘…your information and southern British Columbia, Canada and and lists will be most useful to me should I have northeastern Washington state, USA, an occasion to publish further communications in area known as the Okanogan1 Highlands this field’’ (Stebbins, written communication (Figs. 1–16). This region contains a rich fossil 1994). In fact, Stebbins (1993) did make refer- flora that differs markedly from contempora- ence to fossil Rosaceae, in his commentary on neous floras of southeastern North America the genus Neviusia A. Gray. But, the fossil and the coastal areas of the Pacific Northwest. material itself was not formally described until The Claiborne Formation of Kentucky and more than ten years later (DeVore et al. 2004). Tennessee has a megathermal flora dominated There is no shortage of plant fossils represent- by lauraceous leaves and legumes (Dilcher ing the Rosaceae, but there is a shortage of 1973), and the Chuckanut and Puget floras of paleobotanists working on deciphering the the North Pacific Coast are warm temperate fossil history of the family. floras with sabal palms (Graham 1999). In There is indeed a rich fossil record for contrast, the Okanogan Highlands floras con- Rosaceae. The fossil record of rosaceous plants tain both megathermal and temperate ele- is extensive in North America and Europe from ments. the Eocene on. Neogene rosaceous occurrences The most diverse of the Okanogan High- are more widespread with reports known in lands floras, at Republic, Washington contains Asia, particularly Japan, the Arctic, several a prominent coniferous component (Schorn Gondwanan regions, and northern Africa. and Wehr 1986, Wolfe and Wehr 1987), as well Fruits, flowers, seeds, pollen, wood and leaves as a rich dicot representation (Wehr and are known from many localities, however, the Hopkins 1994, Johnson 1996, Wehr and Man- fossil record has not been well summarized, chester 1996, Pigg and Wehr 2002, DeVore et and therefore has not been readily available to al. 2005, Dillhoff et al. 2005, Greenwood et al. neontologists interested in the family’s evolu- 2005). Early evidence of several families and tionary history. In this paper we review the genera that later become prominent members evolutionary history of the Rosaceae as re- of the cooler temperate floras of the Neogene flected in the fossil record, with a strong are found in the Okanogan Highlands floras. emphasis on the North American record, Along with the diverse Acer L. species (Wolfe particularly from the Okanogan Highlands. and Tanai 1987) and members of Betulaceae We briefly highlight significant European records, however the interested reader should consult Kirchheimer (1973) and the papers of 1 ‘‘Okanagan’’ is the alternative Canadian spelling. M. L. DeVore and K. B. Pigg: Fossil history of Rosaceae 47

Figs. 1–7. Fossil and extant rosaceous flowers. cf. Prunus (1–3)ExtantPrunus (4)cf.Spiraea (5, 6) Extant Spiraea (7). 1 Prunus-like flower showing ovoid, asymmetric ovary on long pedicel, elongate style with broad stigmatic region, and perianth parts attached to hypanthial rim. Relative size of ovary suggests this specimen may be post-fertilization; SR 96-09-04, x 4; 2 Young Prunus-like flower showing gynecium with attached perianth and stamens, SR 96-11-47a, x 3.5; 3 Young Prunus-like flower showing gynecium surrounded by perianth parts attached to hypanthial rim; SR 05-05-06b, x 4; 4 Extant Prunus flower in longitudinal section, for comparison, x 10; 5 Spiraea-like infructescence, UWBM 56785a (B3389), x 4.3; 6 Counterpart of Spiraea-like infructescence in Figure 5, showing detail of fruit, UWBM 56785b, x 7.5 (B3389); x 7.5; 7 Extant Spiraea latifolia (Ait.) Borkh. fruits for comparison, NYBG

(Crane and Stockey 1987, Pigg et al. 2003), close to, but perhaps not exactly like, extant perhaps the most intriguing family in these genera; and (3). Extinct genera with affinities floras is the Rosaceae. at higher taxonomic levels, such as Paleorosa The rosaceous genera of the Okanogan Basinger (Basinger 1976, Cevallos-Ferriz et al. Highlands floras fall into three general cate- 1993) and Stonebergia Wolfe & Wehr (Wolfe gories: (1). Extant genera that are well estab- and Wehr 1988) which combine characters of lished by the Eocene; (2). Eocene forms are several genera or even traditionally recognized 48 M. L. DeVore and K. B. Pigg: Fossil history of Rosaceae

Figs. 8–16. Fossil rosaceous leaves. cf. Spiraea (8–10) Amelanchier (9) Prunus (12, 15, 16); Photinia pagae (13); Stonebergia 14. 8 Spiraea-like leaf with prominent stipule, UWBM 71372b (B2737), x 1.3; 9, 10 Part and counterpart specimen of Spiraea-like leaf, UWBM 54112, x 0.9; 11 Amelanchier, UWBM 54168 (3389), x 1.3; 12 Overview of Prunus leaf showing position of paired basal glands on lamina (arrows), SR 05-05-01a (4131), x 1; 13 Photinia pagae, UWBM 71291 (B2737), x 1.5; 14 Stonebergia holotype specimen UWBM 54110a, 54111, x 1.4; 15 Prunus leaf showing detail of higher order venation and glandular, marginal teeth, SR 00-05-17a, x 5.4; 16. Detail of basal portion of leaf in Fig. 12, showing laminar glands (arrows), SR 05-05-01a (4131), x 3.8 subfamilies. It is not unusual for both extant L. (including related forms; Figs. 1–3, 12, 15, and extinct taxa to be found side by side in 16). The first acceptable records of Prunus fossil floras of Eocene age (DeVore et al. 2004, come from these lacustrine deposits and 2005). In other situations, the fossils are include fruits, wood, and leaves. Additionally, strikingly similar to their extant counterparts, in this paper we illustrate flowers and young but may show a wider range of variation or fruits from Republic currently under study that have certain features rare or otherwise are remarkably similar to those of extant unknown for them. Prunus (Figs. 1–4). Brown (1962) is often The best known rosaceous genus of the credited with describing the first occurrence of Eocene Okanogan Highlands floras is Prunus Prunus endocarps based on material from M. L. DeVore and K. B. Pigg: Fossil history of Rosaceae 49 the Paleocene of Colorado. However, Steven types from Europe (Dorofeev 1963; Kirchhei- Manchester has recently reassessed these fossil mer 1973; Mai 1984, 1995). These are known fruits and has interpreted them to belong primarily from external morphology and not within the family Icacinaceae (S. Manchester, anatomical structure. Florida Museum of Natural History, personal Permineralized wood of the traditional communication). subfamily Prunoideae has been found by the Permineralized Prunus endocarps with Eocene in western North America, the Oligo- internal anatomical preservation occur in the cene of Europe, and the Oligocene-Miocene of Okanogan Highlands Princeton chert of south- Asia (reviewed by Wheeler and Manchester ern British Columbia (Cevallos-Ferriz and 2002). A unique combination of features that Stockey 1991), and in the coeval Clarno Nut are generically diagnostic of Prunus include Beds of Oregon (Manchester 1994). The endo- narrow, numerous vessels that are solitary carps are important because they provide and in multiples, simple perforations, alternate anatomical structure of reproductive organs intervessel pits, heterocellular rays, fibers with that can be compared to that of extant distinctly bordered pits, diffuse parenchyma, cherries. They also provide a fossil record that and traumatic axial canals (Wheeler and can be assessed independently from the leaf Manchester 2002). Wood referred to Prunus record. Prunus endocarps are quite rare in the gummosa (Platan.) Wheeler, Scott & Bargho- Princeton chert, with only a handful of spec- orn is known from western North America in imens known among the thousands of fossils Yellowstone Park (Wheeler et al. 1978), the recovered from this well-studied assemblage Princeton chert (Cevallos-Ferriz and Stockey (Cevallos-Ferriz and Stockey 1991). They are 1990), and the Clarno Formation (Wheeler thought to have been incorporated from ‘‘up- and Manchester 2002). land’’ sources, rather than from the autoch- A diverse assemblage of Prunus leaves is thonous sediments like the mostly aquatic in also known from the Okanogan Highlands situ plants (Cevallos-Ferriz et al. 1991, Pigg floras (Wehr and Hopkins 1994, Dillhoff et al. and Stockey 1996). Cevallos-Ferriz and Stock- 2005, Greenwood et al. 2005). In addition to ey (1991) described three forms, all of which sharing typical features of Prunus leaf mor- show the characteristic apex, endocarp and phology, venation and tooth type (Fig. 15), seed coat layers with the same organization leaves are recognizable by the characteristic seen in extant Prunus. However, these patterns presence of paired glands on the petiole or were difficult to relate in detail to extant cherry basal region of the lamina (Figs. 12, 16). endocarps since comparative data are not Among the hundreds of specimens found, known from all extant groups of this large, there is a broad range of variation in leaf cosmopolitan genus. Manchester (1994) morphology, with at least five or six general reported two species from Clarno: Prunus types present. weinsteinii Manchester and P. olsonii Man- Along with the Prunus leaves from the chester. Prunus weinsteinii is similar to the Republic site, ongoing collecting efforts of the endocarp type 1 of the Princeton chert (Ceval- Stonerose Interpretive Center at Republic, los-Ferriz and Stockey 1991), and which Washington, have resulted in the recovery of Manchester compared with extant P. avium a group of small rosaceous flowers and young (L.) L. Manchester also named Pruniticarpa fruit remains. These specimens, which are cavallosii Manchester for an endocarp from currently under study (Figs. 1–3) show many the Clarno Formation that had similarities features consistent with flowers and young with Prunus but could not be placed in the fruits of extant Prunus (Fig. 4). These flowers genus conclusively. The record of Prunus are 1–1.6 cm long, and are attached to an endocarps continues on into the Late Paleo- elongate pedicel. Gynoecia have broad, some- gene and Neogene with a range of endocarp what flattened stigmas similar to those seen in 50 M. L. DeVore and K. B. Pigg: Fossil history of Rosaceae extant Prunus (Figs. 1, 4), elongate styles this taxon are not consistent with what we and an ovoid, somewhat asymmetric ovary currently know about rosaceous flowers. The (Figs. 1, 3) which one might expect in a authors place Landeenia in the Sapindales drupaceous fruit with a single developing ovule incertae sedis. This report reminds us to (Fig. 4). Perianth parts are attached along a question the confidence with which we can hypanthial rim (Figs. 1–3) and include several identify the presence of Rosaceae on the basis whorls of petals and/or sepals. Anthers are of pollen alone. present in at least one flower (Fig. 2), and are Another rosaceous reproductive structure folded inward as in extant Prunus (Fig. 4). from the Clarno Formation is Quintacava They are being studied for in situ pollen. Manchester (Manchester 1994). Although it Another genus for which we have both cannot be assigned to a modern genus, this fertile and vegetative evidence is Spiraea anatomically preserved fruit shares similarities L. (Figs. 5, 6, 8–10). One recently discovered with the Maloideae. Wood of the Maloideae is infructescence from Republic is 2 cm long with also known at Clarno and may represent the 14 small fruits (Figs. 5, 6), each 2 mm long, that same plant (Wheeler and Manchester 2002). are remarkably like those of extant Spiraea Leaves recognized as Malus L./Pyrus L. are (Fig. 7). Beautifully preserved Spiraea-like part of the ‘‘Eocene Orchards’’ at Republic leaves are also known (Figs. 8–10). Interest- (Wehr and Hopkins 1994). Pollen of Malus/ ingly, however, these commonly have promi- Pyrus has also been reported from the Late nent stipules (Fig. 8), unlike the extant genus in Eocene Florissant locality (Leopold and Clay- which they are usually absent. Poole 2001). The most completely known reproductive The majority of Okanogan Highlands structures of Rosaceae in the Okanogan High- rosaceous remains are known not from repro- lands are the anatomically preserved flowers ductive structures but from leaves. Although assigned to Paleorosa. This flower was first there are limitations, genera with distinctive described by Basinger (1976) from the Prince- foliage features can provide significant infor- ton chert. Additional information was later mation. Rosaceous leaves are simple to highly reported by Cevallos-Ferriz et al. (1993). dissected but seldom truly compound, and Paleorosa shares features of both the tradi- typically stipulate (Wolfe and Schorn 1990). tional subfamilies Maloideae and Spiraeoideae, They typically have brochidodromous second- with its closest similarities to the firethorn ary venation, ‘‘A-O’’ tertiaries and composite Pyracantha M. Roemer (Cevallos-Ferriz et al. intersecondaries. Leaves are typically toothed, 1993), a genus which may also be represented with variably shaped single- or multiple-sized by leaf remains at Republic (W. Wehr, written teeth. Teeth typically have weakly developed communication). The Princeton chert thus ‘‘rosoid’’ venation, with a central vein and demonstrates a classic Eocene situation where weak laterals, and often small apical glands. an extinct genus, Paleorosa, occurs in the same We recently described Neviusia dunthornei strata as a modern genus, Prunus. DeVore, Moore, Pigg & Wehr, based on leaves Paleorosa is significant in containing the from the Okanogan Highlands site at One Mile oldest known in situ rosaceous pollen; isolated Creek, British Columbia (DeVore et al. 2004). pollen is also known in the Eocene of Argen- This is the only fossil record of the small tribe tina (Zetter et al. 1999). Eocene pollen with Kerriae, which contains Neviusia as its only striking rosaceous features was also described North American taxon, along with its two in situ from floral remains of the genus Asian relatives Kerria (L.) DC. and Rhodoty- Landeenia Manchester & Hermsen from wes- pos (Thunb.) Makino. The genus was origi- tern North America (Manchester and Hermsen nally described by Asa Gray from the type 2000). Interestingly, although its pollen ap- species N. alabamensis A. Gray from south- pears rosaceous, other floral features of eastern North America, where it is widespread. M. L. DeVore and K. B. Pigg: Fossil history of Rosaceae 51

Several years ago, surprisingly, a second spe- Stonebergia Wolfe & Wehr (Fig. 14), an cies, N. cliftonii Shevock, Errter & Taylor was extinct genus in the Chamaebatiaria (Porter) found as a narrowly distributed endemic at Maxim. complex is first recognized in the Mount Shasta, in northern California (Shev- Eocene (Wolfe and Wehr 1988). Additional ock et al. 1992, Ertter 1993, Stebbins 1993). extinct genera in this group (Stockeya Wolfe The fossil leaf from One Mile Creek most and Wehr, Eleopoldia Wolfe and Wehr) were greatly resembles the California species with its named from later Eocene and Oligocene floras wide leaves with broad teeth, rather than the (Wolfe and Wehr 1988). southeastern N. alabamensis which has longer, Rosaceous genera are also important com- narrower leaves with smaller teeth. ponents of Late Eocene floras in western North Photinia pagae Wolfe & Wehr (Fig. 13) was America, as characterized by the famous Flor- described from the middle Eocene Republic issant Formation of Colorado (MacGinitie flora of eastern Washington state by Wolfe 1953, Wolfe 1987, Manchester 2001, Meyer and Wehr (1987), and a second undescribed 2003). The Florissant flora includes Rubus, species is known from the same flora (Wehr, Malus, Prunus and possibly Vauquelinia. Addi- personal communication 2000). This genus, tionally, this flora has some of the earliest which is often cultivated as an ornamental has occurrences of Cercocarpus, Crataegus, Holo- a native Asian distribution today, with 40 discus (K. Koch.) Maxim. (Schorn 1998), Rosa species occurring from the Himalayas to Japan L., and leaves referred to Lomatia R. Br. and Sumatra. Thorns from this locality may also represent The oldest evidence of the genus Amelan- Rosa (Meyer 2003). chier Medikus is from the middle Eocene Both leaves and fruits of Rosa occur later Okanogan Highlands of British Columbia in the Oligocene Bridge Creek flora of Oregon (Fig. 11; W. Wehr, personal communication, (Meyer and Manchester 1997), as well as in 2001). This genus has an extensive fossil record later Neogene floras of Europe (Mai and of leaves throughout western North America. Walther 1978; Kvacˇ ek and Walther 1998, Fossil occurrences are relatively common in 2001, 2004; Hably et al. 2000; Kvacˇ ek et al. Eocene and Miocene floras of western North 2004). The European and North American leaf America, and have been reviewed by Schorn and fruit species of Rosa differ, suggesting the and Gooch (1994). Amelanchier and Ama- North American-European disjunction within lanchites have been reported from the Paleo- this genus today may be of ancient origin cene Ravenscrag flora of Saskachewan (Becker 1963, Mai and Walther 1978, Hably (McIver and Basinger 1993), however their et al. 2000, Tiffney and Manchester 2001). identification with Rosaceae has not been The Late Oligocene Creede flora of Colo- verified. rado (Axelrod 1987; Wolfe and Schorn 1989, Other rosaceous leaves with first or early 1990) has a rich rosaceous component. Among occurrences in the Okanogan Highlands in- members of Spiraeoideae are the Eurasian clude those similar to the widespread genera genus ‘‘Eleiosina Raff.’’ (=Sibiraea Maxim.), Rubus L., Crataegus L. and Sorbus L., the Holodiscus, and the extinct genera Eleopoldia eastern Asian Sorbaria (DC) A. Braun., and and Stockeya. Eleopoldia is closely related to Pyracantha today of Europe and Asia. Other extant herbaceous plants Luetkea Bong., Geum extant genera in the Okanogan Highlands L., and Sanguisorba L., while Stockeya is an floras show the transition into drier habitats. ancestral relative of the Chamaebatiaria group. These include the chaparral plants Hesperom- Other rosaceous genera include Sorbus, Cra- eles Lindley which is related to extant Central tageus, Potentilla, Cercocarpus (with possible and South American taxa, as well as Vauqu- achenes), ‘‘Osmaronia E. Greene’’ (=Oemleria elinia Baillon. and Cercocarpus Kunth. which Reichb.), and rare Prunus. Wolfe and Schorn occur today in southwestern North America. (1989) analyzed the paleoecological distribution 52 M. L. DeVore and K. B. Pigg: Fossil history of Rosaceae of these taxa in the Creede flora using sented by nine species (Reid and Reid 1915). multivariate statistical techniques and recog- Rubus, which is known as early as the Eocene nized four major plant communities. Stockeya (Chandler 1962, 1963) undergoes further radi- was an important member of the mesic ation and has been documented in many juniper woodland community at Creede, while European floras (summarized in Mai 1995). its modern relative, the shrubby desert sweet Other genera known in the European Neogene (C. millefolium) inhabits drier high-desert, include Cotoneaster Medikus., Pyrus, Mespilus chaparral sites today. Wolfe and Schorn L., Pyracantha, Stephanandra Sieb. & Zucc., (1989) concluded that a habitat shift related Physocarpus, Comarun L., Potentilla L., Agri- to modern physiological adaptations occurred monia L., and Filpendula Mill. (Kirchheimer in the Stockeya lineage prior to morpholog- 1973; Mai 1984, 1995 and references cited ical change. Several other genera at Creede therein). are closely related to modern-day herbaceous In Africa Bruce Tiffney has recognized the genera, suggesting that a shift from shrubby first documented occurrence of a Prunus to herbaceous habit also occurred (Wolfe and endocarp (subgenus Amygdalus) from the Schorn 1989). Neogene of Kenya (Tiffney, personal commu- Lyonothamnus A. Gray is a monotypic nication, 2005). In Asia Prunus, Sorbus, and evergreen shrub that is today endemic to the Rosa are known from the Neogene of Japan Channel Islands off coastal California. Based (Tanai and Onoe 1961, Miki and Kokowa on its fossil record it is clear that this genus 1962, Tanai and Suzuki 1963). Rosaceous had a broader distribution in the Neogene genera in Late Tertiary and younger strata than today, extending from Oregon to south- (early Miocene to Early Quaternary) of Arc- ern California and into west central Nevada tic and subarctic North America include (Erwin and Schorn 2000). The genus is first Dryas L., Cratageus, Fragaria L., Physocarpus, recognized in the Early Miocene (23 my) Potentilla (four species), Prunus (two species), Collawash flora of Oregon and occurs in Rubus (two species), Rosa, Sorbus, and Geum numerous Middle and Late Miocene localities, L. (Matthewes and Ovendon 1990). and, like today, in known only from far Pollen records of Rosaceae were reviewed western North America. by Muller (1981). He recognizes Parastemon The Neogene fruit and seed record of A. DC type pollen in the Oligocene and lower Europe contains many rosaceous genera (Reid Miocene of Cameroon and the upper Miocene and Reid 1915; Chandler 1962, 1963; Dorofeev of Borneo and Filipendula type and Sangui- 1963; Mai and Walther 1978; Martinetto 2001; sorba officinalis L. from the Pliocene of the Kvacˇ ek and Walther 1998, 2001, 2004). Re- Netherlands. Pollen of Adenostoma Hook & views of older rosaceous fruit and seed litera- Arn. (Chamise), an extant component of ture can be found in Kirchheimer (1973); more today’s chaparral vegetation of California, recent reviews are in Mai and Walter (1978) has been described from the Late Pleistocene and Mai (1995). Prunus endocarps are well of the Santa Barbara Basin (Heusser 1978), known and have been reviewed by Mai (1984). and the Late Pleistocene La Brea Tar Pits By the Miocene, genera such as Rosa, Cratae- (Stock 1992). Rosaceous pollen in the postgla- gus and Rubus are common elements of the cial record of western Canada, has been Bohemian Bilini leaf flora (Kvacˇ ek et al. 2004). studied by Williams and Hebda (1991). The Miocene-Pliocene of Saugbagger flora of Alsace, eastern France contains fruits of Ru- Summary bus, Crataegus, Mesopilus, Sorbus, and Prunus (seven species; Geissert et al. 1990). In the Currently, the fossil record of the Rosaceae Pliocene of the Netherlands Rosaceae were can provide some insights to dating the abundant, especially Prunus, which was repre- origin and evolution of major tribes and M. L. DeVore and K. B. Pigg: Fossil history of Rosaceae 53 genera within the family. Reciprocially, molec- evidence of the evolutionary history of rosa- ular data provides a context for interpreting ceous plant occurrence. what vegetative, fruit and floral characters are Biogeographical relationships within Ros- stable enough to be used in unraveling the aceae can also be illuminated partly with the array of fossil leaves that clearly have affinities fossil record. For example, a recently described within the Rosaceae. fossil species of Neviusia from One Mile Creek In the fossil record, plant remains are has helped to interpret the disjunct relation- typically found as disarticulated organs rather ship of N. alabamensis and N. cliftonii (DeVore than whole plants. It is even more important in et al. 2004). Other members of the rose family, this type of situation to be able to recognize the for example, Photinia, which is documented in salient features that define a given taxon. both Republic and Princeton floras, is only Rosaceous fruits and flowers are readily iden- naturally distributed in Sumatra, Japan, and tifiable by their characteristic features, and the Himalayan Mountains today (Wolfe and indeed, fruit characters are the primary features Wehr 1987). Fossil rosaceous taxa from of each traditional subfamily. Some rosaceous western North America may also give clues leaves also have distinctive characters, as do to the genera with South American ties. For pollen grains and woods. It is possible to example, Hesperomeles, found at Republic, is identify fossils to the family, and to a genus or currently distributed in Central America and subgeneric level when the right characters are Peru (Wehr and Hopkins 1994). However, the preserved. Fortunately, the fossil record favors use of fossil taxa in discerning biogeographic preservation of leaves, fruits and seeds and has patterns within the family will have to wait the potential to provide insights to character until additional taxa are formally described evolution within the family. and affinities of these fossil members of Ros- It is often difficult to determine the phylo- aceae can be assessed. genetic relationships of these fossil occurrences The family clearly radiated in response to to modern genera. Fossils with rosaceous the active tectonic margin of western North affinities sometimes demonstrate a mosaic of America, generating directly, or indirectly, new characters of several extant taxa and are habitats as suggested by Stebbins. European difficult to place systematically. Other rosa- occurrences are widely documented, especially ceous fossils lack sufficient diagnostic charac- for the Neogene, and additional Asian, and ters to place in extant genera, while still others rare African representatives are known. As show a wider range of variability than that current studies on different continents pro- found within individual genera today. Some of gress, it will soon be essential to examine the these larger groups may represent fossil evi- global distribution of the family to better dence of polyploid or hybrid complexes, such understand the details of its complex evolu- as occur in the Rosaceae today. Because of tionary history and paleobiogeography. these problems, paleobotanists have often been reluctant to assign some fosssils to established This review is dedicated to the late Wes Wehr, genera (see Becker 1963). As a result the fossil Burke Museum of Natural History & Culture, record, particularly of the Paleogene has not Seattle, who inspired and encouraged our interest in the fossil Rosaceae, and the late Jack A. been readily accessible to neontologists inter- Wolfe, who made many of the original identifi- ested in rosaceous phylogeny. Neogene occur- cations of Republic Rosaceae and recognized the rences, particularly of fruits and seeds, are paleoecological significance of leaf physiognomy. better known in Europe, however these too We thank Drs. Volker Wissemann and C. S. have not been exhaustively studied. While we Campbell for inviting our participation in this readily acknowledge that there are limitations symposium volume; Wes Wehr and Ron Eng, to what can be learned from the fossil record, Burke Museum of Natural History & Culture, we also recognize its unique potential as direct Thomas and Richard M Dillhoff, Evolving Earth 54 M. L. DeVore and K. B. Pigg: Fossil history of Rosaceae

Foundation, and Lisa Barkesdale and Catherine Cevallos-Ferriz S. R. S., Stockey R. A., Pigg K. B. Brown, Stonerose Interpretive Center for access (1991) The Princeton chert: Evidence for in situ to fossils for study; Steven R. Manchester for aquatic plants. Rev. Palaeobot. Palynol. 70: 173– photographs of fossil Spiraea-like fruits and 185. extant Spiraea; Bruce Tiffney for literature and Chandler M. E. J. (1962) The Lower Tertiary floras helpful comments on a draft of the manuscript; of southern England 2. Flora of the Pipe-Clay Harald Walther and an anonymous reviewer for Series of Dorset (Lower Bagshot). British helpful comments on the manuscript; Gary Lun- Museum Natural History, London. dell, University of Washington Libraries, for Chandler M. E. J. (1963) The Lower Tertiary floras library assistance; John C. Benedict for technical of southern England. 3. Flora of the Bourne- assistance; and the late G. Ledyard Stebbins, Jr. mouth Beds; the Bouscombe, and the Highcliff for his insights on the significance of the fossil Sands. British Museum Natural History, Lon- record to rosaceous evolution. This research was don. supported in part by National Science Founda- Crane P. R., Stockey R. A. (1987) Betula leaves and tion grants EAR-0345569 (to MLD) and EAR- reproductive structures from the Middle Eocene 0345838 (to KBP). of British Columbia, Canada. Canad. J. Bot. 65: 2490–2500. DeVore M. L., Moore S. M., Pigg K. B., Wehr W. References C. (2004) Fossil Neviusia leaves (Rosaceae: Archibald S. B., Pigg K. B., Greenwood D. R., Kerriae) from the lower-middle Eocene of Manchester S. R., Barkesdale L., Johnson K. R., southern British Columbia. Rhodora 106: 197– Sternberg M., Stockey R. A., DeVore M. L., 209. Rothwell G. W. (2005) Wes Wehr dedication. DeVore M. L, Pigg K. B. (2005) Rosaceous Canad. J. Earth Sci. 42: 115–117. evolution in the Okanogan Highlands, Wash- Axelrod D. I. (1987) The Late Oligocene Creede ington, USA and British Columbia, Canada. Flora, Colorado. Univ. Calif. Publ. Geol. Sci. XVII International Botanical Congress, Vienna, 130: 1–235. Austria, July 18–23, 2005 (poster). Basinger J. F. (1976) Paleorosa similkameenensis DeVore M. L., Pigg K. B., Wehr W. C. (2005) gen. et sp. nov., permineralized flowers (Rosa- Systematics and phytogeography of selected ceae) from the Eocene of British Columbia. Eocene Okanagan Highlands plants. Canad. J. Canad. J. Bot. 54: 2293–2305. Earth Sci. 42: 205–214. Becker H. F. (1963) The fossil record of the genus Dilcher D. L. (1973) A paleoclimatic interpretation Rosa. Bull. Torrey Bot. Club 90: 99–110. of the Eocene floras of southeastern North Brown R. W. (1962) Paleocene flora of the Rocky America. In: Graham A. (ed.) Vegetation and Mountains and Great Plains. US Geological vegetational history of northern Latin America. Survey Prof. Paper 375: 1–119. Elsevier Science Publishers, Amsterdam, pp. 39– Cevallos-Ferriz S. R. S., Erwin D. M., Stockey R. 59. A. (1993) Further observations on Paleorosa Dillhoff R. M., Leopold E. B., Manchester S. R. similkameenensis (Rosaceae) from the Middle (2005) The McAbee flora of British Columbia Eocene Princeton chert of British Columbia. and its relation to the early-Middle Eocene Rev. Palaeobot. Palynol. 78: 277–292. Okanagan Highlands flora of the Pacific North- Cevallos-Ferris S. R. S., Stockey R. A. (1990) west. Canad. J. Earth Sci. 42: 151–166. Vegetative remains of the Rosaceae from the Dorofeev P. I. (1963) Tertiary floras of western Princeton chert (Middle Eocene) of British Siberia. Komorov Botanical Institute Academy Columbia. IAWA (International Association of Nauk SSSR, Leningrad, USSR. Wood Anatomists) Bull. New Series 11: 261– Ertter B. (1993) What is snow-wreath doing in 280. California? Fremontia 22: 4–7. Cevallos-Ferriz S. R. S., Stockey R. A. (1991) Erwin D. M., Schorn H. E. (2000) Revision of Fruits and seeds from the Princeton chert Lyonothamnus A. Gray (Rosaceae) from the (Middle Eocene) of British Columbia: Rosaceae Neogene of western North America. Int. J. Pl. (Prunoideae). Bot. Gaz. 152: 369–379. Sci. 161: 197–193. M. L. DeVore and K. B. Pigg: Fossil history of Rosaceae 55

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Cenozoic of western North America: Journal of Argentina. XVI International Botanic Congress, the Faculty of Science, Hokkaido University, St. Louis. Abstract Number: 2556; Poster No. = Series IV 22: 1–246. 952. Wolfe J. A., Wehr W. (1987) Middle Eocene dicot- yledonous plants from Republic, northeastern Washington. US Geol. Survey Bull. 1597: 1– 25. Addresses of the authors: Melanie L. DeVore Wolfe J. A., Wehr W. (1988) Rosaceous Cham- (e-mail: [email protected]), Dept. Biologi- aebatiaria-like foliage from the Paleogene of cal and Environmental Sciences, Georgia College & western North America. Aliso 12: 177–200. State University, Milledgeville, GA 31061 USA. Zetter R., Hofmann C.-C., Draxler I., Ferguson D. Kathleen B. Pigg (e-mail: [email protected]), School K., Durango de Cabrera J., Del Vergel M. (1999) of Life Sciences, PO Box 874501, Arizona State Innovations in Eocene vegetation of southern University, Tempe, AZ 85287-4501 USA.