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Biogeographical Relationships of North American Tertiary Floras PDF

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Preview Biogeographical Relationships of North American Tertiary Floras

BIOGEOGRAPHICAL NORTH RELATIONSHIPS OF AMERICAN TERTIARY FLORAS' Europe document lllo-r .Hill \sl.l lull (.1 .1 1 selected conifer and angiospcnii geueia ll I The ami The origin of the exianl Mora vegetation of purpose review this highlight Tertian- is <>l l<> North America has heen the subject of much in- records of selected conifer and angiosperm genera terest and debate ever since the similarities ma\ be considered soundly and flori-lie that identified between North America and Asia were docu- which are significant understanding die hiogeo- first in mented. Many of the woody genera comprising graphic affinities of North American Tertiary floras. North America's present-day forests have excellent present a review of about 90 genera with reliable I fossil records that can be traced through the Ter- Tertiary records in North America and other con- tiary in North America and other continents of ihe tinents. Examination the straligraphic ranges of of Northern Hemisphere (Wolfe, 1975; Graham, 1993; different taxa in North America. Europe, and Asia many Tiffney, 1985a, b; Mai, 1995). In addition, provides the basis for assessing ph\ logeograpluo genera that are no longer native to North \uierua patterns and pathways biota dispersal through of have well-documented Tertian records. Patterns of the Tertiary. South American and African records among geographic disjunction extinct and extant are mentioned when known, but the emphasis on is genera provide important clues to the history of affinities within the Northern Hemisphere. For a North American and former flora continuity of review of relationships between North and South lh. Tertiary forests in the Northern Hemisphere. By America, see Burnham and Graham (1999, this is- eomparing the stratigraphic records of genera sue). among shared two or more continents possible Because of our familiar vantage point of the pres- is if to consider the pathways and timing of plant inter- ent day. neobi botanists alike tend change through the Tertiary. to regard the modern flora as an endpoint showing Many ideas have been published on the paleo- the "true" floristic patterns of extant genera and botanical origins and development of extant flora species. Thus a genus such as Ginkgo, with a wide and vegetation of the Northern Hemisphere (e.g.. paleogeographic distribution (Tralau, 1968), mav be Engler, 1879; Chaney, 1940, 1947; Wolfe, 1975; classified as an "East Asian element." Clearly, such & & Raven Latham Axelrod, 1974; Ricklefs, 1993; "elements" have more to do with extinctions else- Mai, 1995; Akhmetiev, 1996). In the attempt to be where than they do to the natural geographic affin- comprehensive as as possible, investigators have ity or origin of the genus (Wolfe. 1975). Fossils can sometimes relied uncritically upon genera reported document former geographic' distribution patterns many in the literature. Closer scrutiny reveals er- of both extinct and extant genera and thus provide roneous generic determinations (Dilcher. 1974). a means of tracking the changing floristic relation- ollii I W. Wehr, Wen, and V V. J. ','," .Nln^e,,. Ilongshan War h translation of Chinese Mali ill. lil 1 1 m!, >v R. Stockey, B. ' Iiiln Kvaoek, and editing by ^ I. erendeen and Can.- Peter ,,ln. II for , 1 EAR This rese ai.li was supported in part h\ grants 9506727 and l\l'')722lll 1 499 Mu .resents no. in the Contribiations to Paleobiology fron the Florida i History. Gainesville, Florida 3261 U.S.A. iral 1, Gaud. 472-522. 86: 1999. Volume Number 2 86, Biogeographical Relationships landmasses through Tertian genera Kirchheiincr diffei is thai of (l'>->7). mm Re. siudies have emphasized pholngu ml ml iphasizes the European fossil record but .-nl ; molecular phylogenetics as a framework to con- also repol'ls selected reliable occurrences nf genera Many log. graphic In-lorv iii^jian la\a ir.- in North America and Asia. of the age as- - ' |' ; r. i 1 1 j • . « I I icuo.l hv Wen. in pi. ss _i signments in Kirchheimers catalog have since been \ 1! .g i i II i I [i ses .lev eloped from phylogenetie and molecular revised, but Kirchheimer was careful to give local- studies can be tested by observations from the fossil current age assignments by attention more recent to It might be argued that the fossil record is too literature. Helpful, but less comprehensively doc- ihonm; f-agmeiiiarv in piov ((!.' a i.m lor-tauditi- ol umented, reviews ta\a icpoiicl the paleobo- li of in however, ..graph the alternative of ic v tanical literature are provided by Taylor (1990), i I . i ; i using only model, - i il uti >n I * i > I Collinson et al. (1993), and Mai (1995). Useful potheses del to li i - I l.i-i guides to Asian literature on Cenozoic Paleobotany i | i ' ' leads In an even more incomplete picture. If forced include Tanai (1992a, 1994) and Liu et al. (1996). we to rely onlv on extant generic distributions, Published reviews of the record of critical fossil would be Carya had unlikely to predict that its particular phylogenetie groups, for example, that of Europe greatest diversification Tertiary of in tin- the magnoliid angiosperms (Friis et ah, 1997), and (Mai, 1981; Manchester, 1987a) despite absence its individual families Retulaceae, Crane. 1989) (e.g., from the extatil European flora, or that Ensete and or genera Nyssa, Eyde, 1997) are also very (e.g., (both native Africa and Asia itha to to- day) would be present in the early Tertiary of North The decision whether accept generic records to America (Manchester, 1994b). summary was based on whether the organ(s) for this American In this article, review the North Ter- I and morphological/anatomical features preserved and described can be considered diagnostic truly tmm comparison gmi. w -Iraligm: -j.,'-!ii ia in ill: la< genus mod- the indicated. \- the taxomuiiv of ol in Northern Hemisphere. other parts of the l make go ern plants, often difficult to reliable is it based upon Rare paleobotanical specimens showing and/or fruits floi foliage have revealed "mosaic" plants, in which the modern leaves closely resemble those of a genus, while the or llowers icveal novel characters limits indicate belong an extinct genus (Man- that lliev to well-known problem paleobotanical a that It is chester, 1989a; Manchester et al., 1998). In the number includes a large of generic as- literature Vrl dubious Mgrunorils \uici ol '.all lilv. I:i ic; : 'r unavailable productive parts arc studies of fertile much problen applies of the literature prior to to and material, particularly flowers fruits, prov ide lh. the 1970s and actually persists to the present day. best systematic resolution. ppK This hecau.se iiivcsiigatnrs <iinVi is diif.'-tvnl :i Reports based on fossil foliage are also impor- what necessary idenliS ni -1 h ilards of is to more commonly tant, particularly since leaves are may nus. also -n It l>. _ I. I I I I i many preserved than depositional environ- fruits in names ohtam geologists quick, tentative for fos- to ments, hut the) require careful seruthn to deter- may may that not be well preserved or that not sils more provide characters a secure de- sufficient for Leaves some families arc so distinctive as tions. in main assigmin .> par In nisi in. n! to a to be reliable in diagnosing a genus, as in Rlalan- us iiiscnssiiig feu gci lied is |iis! l>\ n ! Herb arc >etween the fossil species and the mod- a. < i. in. I < -1 Iv lh. i< convergence and instances of parallelism in foliage from !isliagi.is!i ei iteii.i r|,.it ii- lli, i! me that cause to be less confident of generic as- ss hie candidates are not considered, and signments. In the Betulaceae, Juglandaceae, inadvisable accept reports ion, to is it family using leaf morphology relatively easy, but is secure generic determinations icquire In fruits. may good these families the leaves provide a "best estimalc" genera are present, but identifi- ol lilt 1 1 i ' that the south. -i emphasize megafossil reports of fruits, flowers, stratigraphic charts. The Tertiary record for South I and leaves, because .mi more confident the Vmeriea and \lina rb known relative \silli is still I | aUn and e\aliiiiln>ii ..I lli.se ..rga ib. \\ |><>ll«'n I pro\ide biogeogiaphicallv uiipni laul records, direct exchange of plant species between North and lull South America (luring the early to Lite Tertian re- pollen the de< leased systematic resolution mains weak. For more detailed accounts of rela- <>l is i I aneed h\ the relatively -I at igraphic resolution tionships inferred between the fossil record of itz.ii 1 1 (wind-dispersed (Milieu ma\ he lound throughout a North \meriea and the living genera of Africa and sedimentary sequence, whereas megafossils are South America, see Raven and Axelrod (1974). Bumham eonliiied speeihe laeies). lor eritieal systematic Taylor and and Graham |,. (1990), (1999). The million-year chronology epochs of Tertiary refer to Midler (1981). follows Berggren et al. (1995). The Brandon Lignite In addition, before accepting records for inclu- flora of Vermont significant as one of the few is sion summary, was necessary in this it thai I li«\ I" in ii in I i i il I is ii the north- documented fully in the literature with convincing eastern niled Slates. nlortunately. there are no I I photographs.)] had that the opportunity to observe associated datable rocks or annual fossils to pro- I means ide an independent determining the age. v ,,l am to stu.K eritieal specimens. indebted en though manv voars considered be to \ lor likely to I iatois ol the following collections: niversitv of Oligoeene, the current conscn-u- is that mav be I il California, Berkeley (specimen numbers cited have Miocene lW4a; early (Tiffnev. Traverse, 1994). I UCMP); Moscow; the prefix University of Idaho, adopt this age the ussioiis involving Bran- foi .lis. Museum Mn D.nxei of Natural History; IVaho.h don fossils. There are also problems with the pre- seum of Natural History, Yale University (YPM); cise age of many floras in eastern \sia. have usu- I Museum, Hinted States National Washington, D.C. ally accepted the assignments given by the most SNM): Museum Burke of Natural History and eases there continue (I >< !>i - -i ui« to < i 'i' 1 I i i (UWBM); Culture, Seattle Department Biolog,. be disagreements among and al different investigators ..I Sciences, University of Alberta; Natural llis|or\ mav be expected that the ages of manv -lies will il Museum, London (BM-V); Museum. National continue to be revised in the future. Senekenhcg Museum Plague; of Natural History. have attempted to be conservative in the po- I Frankfurt |SM): Botanical Department. Hungarian sitioning of stratigraphic i.uigcs. hu- the ranges 1 Museum, Natural History Budapest; Geological presented here In- are often shorter (begin than later! stitute and Geological Museum, Moscow: kniuaiov those indicated by other authors, Mai e.g., (1995). Botanical Institute, St. Petersburg; National Sci- lot example, is possible to find reports of En it ence Museum, Tokyo; Nanjing Institute of Geology gelhardia based on pollen from the Paleocene. and Academia Paleontology, Siniea, Nanjing; Insti- tute of Botany, \cadeinia Siniea. Beijing. Speci- occur until the en. .and <>, I mens Ih cited with the preli\ are from the Florida len (usually calf house) known is 1 . Museum of Natural History, Cainesville. Extant have been produced genera to bv othei of Juglan- eom|)arative material was studied herbaria daceae at in- in the Tertiary (Manchester, 1989b). Like- MO, cluding A, FLAS, and PE. wise, there are reports nf ,1/hh.v from the Crela ceous, but without convincing infruclesceiices. may the paper be found Mabberley The Because scope in (1997). the ol this treatment limited to i- known genera the Tertian and/or Becenl ('torn flora hav are specified at their first mention in the text. of North America. omitted some interesting I Stratigraphic charts presented in tin- paper aic examples of taxa shared only between Europe and simplistic in thai the\ condense diverse records Asia, many of which are reviewed bv Mai '><>.">). ( I from each different part- of northern continent into column. However, single enables quick a this a vi- summary some sual that provides constraints on the liming ol intercontinental exchange events for each The following section highlights records that I taxon. More detail- foi ea. h genu- are provided in consider useful in evaluating the biogeographic af- the family narratives of the .Mowing section, lead- finities of North American Tertiary These floras. I. more ing to the informative literature for the genera paragraphs cite the references upon which the Pliocene found ranges I'urope after the Krlclrrriu. to -tra .die in : 21-25 (Fig. 19) and in Figures are based. This sec- day only in the broad-leaved evergreen forests of known arrange China and Taiwan, based on distinctive tion is is I Eocene the (leadings iiih and \ngiosperms, with seeds from the of the CUiilchena flora of (,-, •.in. .-.(>« Columbia Mathewes, unpublished angiosperms mi h treated un- British (R. data), ol last, .1 - i i der die (leading lneertae Sedis. Ciiramon patterns and Oligocene of Oregon (Me\ei Mancheslei. e> are reviewed the subsequent section. 1997). in Taxaceae. Amentotaxus, with four extant spe- cies China and southeast \sia. has an excellent in record North Vineiica and Europe. fossil in It is U|M-ess;Hi-ae. Dislillclive leaves of the ex- iva.lil\ recognized b\ its broad needle-like leaves < » Hi < and .ismici- with a pair of prominent stomatal bands and dis- in ,. \1< .\ i 1 ,' I i i i ated cones occur in the Paleocene of Wyoming, (Ferguson et al., 1978). f & Saskatchewan, Alberta (Mclver Basinger. extends from the ppet Cretaceous «)<•(); I I Mclver, 1992) and in the Paleocene of Altai, Xinj- (Santonian) to Miocene of North America and from ing. China (Quo et al., 1984, as Ditaxocladus Guo), the Paleocene to ppcr Miocene ol Kurope (Per I indicating trausberingial distribution. guson 1978; Jahniehen. 1990). Other genera a et al., and and Tetraclinis, with one species living in the w c-t.au of the Taxaceae, including 7h.vn.s- Torreya, Mediterranean today, has a g Tertian record in at least one extinct genus, arc well represented in I Europe (Kvaeek, 1989; Mai, 1995). In addition, one the Tertiary of western North America (Manchester, & species known tmir goc-:i |o \1io 1994b; Meyer Manchester, 1997) and central is I'M <\ii l\ < >l cue western North America based upon cones, Europe (Kvaeek, 1982), but their history best is of . & As Mesozoic seeds, and foliage (Meyer Manchester, 1997). explored through attention to floras. has not been observed in the east Asian Tertian. Taxodiaceae. This family includes several gen- it appears likely that this genu- traversed the North eta with relietual extant distribution. Metasequoia it Eocene. has become famous as an example genus once Atlantic in the late ol a Hemisphere now *.iiik»oac«'ae. The record of Ginkgo in the widespread in the Northern thai is Northern Hemisphere extends back to the Jurassic. native only to China. do not see the need to review I biogeographie but record the Tertiary of its in is & Meyer interest, documenting the relatively late confine- (most recently in Manchester, 1997). ment Asia. The Tertian records in North Amer- Cunninghamia grows today in mixed mesophytic to ica extend from the Paleocene of the Rocky Moun- and broad-leaved evergreen forests of China and tains (Brown, 1962) to the Miocene of Oregon Taiwan. Its fossil record includes cones, seeds, and (Chancy. 1920). A worldwide review of the distri- foliage from the Eocene to Miocene of western bution genus through space and time was North America (reviewed in Meyer Manchester. ol this t\ Europe and presented by Tralau (1968). 1997), the Tertiary of (Mai, 1995), the m Pinaceae. Various geneia ol iho I'm ...a. Eocene to Miocene of Japan (Matsuo, 1967; Hori- well represented in the Cretaceous and Tertiary of uchi, 1996). the Northern Hemisphere, including, for example, Glyptostrobus is native today only in southeastern & & id' Abies (Schorn Wehr, 1986), Keteleeria (Meyer Tina. but. h ' L ( 1 & Manchester, Larix (LePage Basinger, was widespread the Tertian. extends from the 1997), in It 1991; Schom. 1994). Picea (Crabtree, 1983), Pinus Paleocene (Boulter ex Kvaeek. 1989) to the Plio- (Miller K Mal.nkv. 1986). and Pseudolarix (LePage cene in Europe (Mai, 1995: Martinetto, 1998). & cones Basinger, 1993). Most of these genera are wide- </n europaeus twigs with attached spread in the Northern Hemisphere today, but Ke- occur in the Eocene to Pliocene of Japan Taiuu. ( ',,.'. re limited lo eastern \sia 1961; Malsumotoet al.. 1997b). In North \„,erica. /, ., iid ! ', . in their ru tiis « m! I!asmg< Glyptostrobus is well represented in the Paleocene i I. I I ' provided a comprehensive analysis of the Eocene of the books Mountain region (brown. (1991) to phytogeographic history of Pseudolarix in the 1962; Hoffman, 1996), and in the Miocene of Ida- Chaney Northern Hemisphere based on distinctive ho, Oregon, and Washington (Brown, 1936; its & seeds, ((in.s. and foliage, with earliest records in Axelrod, 1959; Fields, 1996). Cretaceous Asia and North America followed Sequoia and Taxodium, although restricted in the of by Oligocene establishment Europe. They con- then modern distribution to Pacific Coastal North in cluded that the genus became extinct in North America and eastern North \nieiK a. respectively. Hemisphere America by the middle to late early Miocene, and were widespread in the Northern in the . 1995: pi. 9, 2l to the Middle Miocene of Swit- fig. zerland. Araliaeeae. western and Torirellia of eastern \sia. sometimes placed own laimb. was in its eoii- some sidered by taxonomists to be allied to the Cor- Aetinidiaeeae. Artimdia is distributed in In- riaceae. but rec.nl molecular work indicates po- ., and cistern was (lnin,llr-i.i \si;l lodav. lull present sition within the Araliaeeae (Plunkett et 1996). al., North America and Europe in during the IV-i tiar Torirellia has distinctive fruits (Fig. 1A, B) with \ known In North \nicrir.i. mi seeds from three single-seeded i- lia><-il locules: a central locule with a i! mm Eocene formation Otegoti (Manclms- tile (II. o| terminal germination valve and two enlarged, blad- ter. 1994b). In Europe, fossil seeds occur from the der-like lateral chambers that are infertile. The en- Eocene to the Pliocene (Tralau, 1963; Friis. 1985; docaip tissue composed isodiametric is of sclei'eids. Martinetto, 1998). These same characters occur Eocene in fruits (Fig. Anaeardiaceae. Pentoperculum Maneliester ribed as the fossil dm an extinct genus of belonging Spon- is fruits to (1994b). There- dieae shared between die middle Eocene of Oregon fore, now offer the new combination. Tortrrllia I (Manchester, 1994b) and the lower Eocene of Eng- honrsii (Manchester) Manchester comb. nov. (see & land (Reid Chandler, 1933, as Draronton,rlon\. \ppendix Tmiii'llm honrsii occurs the middle in ). I The calyces called Astronium truncatum by Mac- Eocene of the Clarno Formation, Oregon 1C- (Fig. Ginitie (1953) from the Eocene and Oligocene of Roslyn Formation, Washington and F), (Fig. 1G), the western lilted Static are \ei\ different ve- Messel, Germain in (Collinson. 1988: I pi. 1, fig. 11). and nation fruit iimrphologv from the extant genus Fruits are also present the lower Miocene Ob- in and no are longer believed represent \nacardi- to cidorl localih of Austria (Meller, 1996, and pers. & aceae (Manchester Wang, The comm. 1998). leaves that 1998). These occurrences indicate that 7b- MacOinilie placed in the -anie species ma\ in fact iitrilio was shared between Europe and North represent Anacardiaceae, but they are at least as America during the Eo< cue. although un- is still it Rhus similar to as they are to Astronium. Rhus is known from the Asian Tertian. The timing of its known from anatomically preserved from fruits the Eocene Oregon of (Manchester, 1994b). Apocynaeeae. Elongate seeds with a terminal Catalpa lives today eastern in Asia, eastern North America, and the Canbli.'an length of the seed body are found in many extant legion. Small biwinged seeds Calalpa have u beeti f genera Apocvnaceae, of e.g.. Anoilcndron, Clegh- recognized from EaiU Oligocene Oregon the of & ornia, Echites, Forsteronia. Holarrhena, Kibatalia, (Meyer Manchester, 1997). Similarly small Ca- Odontadenia. Pottsia. Prestonia, Strophantus, and talpa seeds (Catalpa mirrosperma Saportai cur o, in \poc\naceous Wrighlia. seeds can be recognized the Oligocene France and Germain late of (Saporla. with ease, but die distinction of genera based onl\ 1889; Weyland, 1937), suggesting a possible North on seed characters is a difficult challenge. Thus, antii inkage during oi pri.n to the Migocenc \ i ( I (= the fossil generic nam.' Krhitoniuin I'nger 1850 Berberidaceae. Mahonia distributed is in = Apocynospcrmum X Chan C\psrlitrs leer lb-id \sia, Malcsia. and North and Central America to- I dler. ( >2<)) is applied to los^l remains. In the North da\. Mihough soiu. iimes subsumed within Herberts I m \rii.iie.in Tertian, such seeds occur the middle iWhittcumre. Mahonia 1997), is easily distin- Eocene tireen Ki\ei formation (Apocynospermum compound guished by genus foliage. This its is Brown coloradensis in MacGinitie, 1969: pi. 18, fig. readib recognized b\ the distinctive an '.lure of Int. the Clarno shales Oregon (Manchester, un- imparipinnately compound and 1). >f its leaves spiny leaf- ( published), and the Eocene late of Florissant, Col- lets, without the need for associated flowers or orado (Manchester, unpublished). They are not and thus fruits, is a g candidal.' for recognition I known in North America after the Eocene, but in in the fossil record. Mahonia well represented m is Europe they extend from the Middle Eocene (M.s- western North an Tertian extending \m.-rii floras, sel, Germany: Senckenberg Collection-ME 7624), from Eocene the to Pleistocene (Schorn, 1966). Al- & the Late Eocene of England (Reid Chandler. though not native in Europe today, occurs the in it 1926), Early Oligocene of Budapcst-Obuda, Hun- Oligocene France and of (Saporla. North Bo- Iii<>5) gai\ (Botanical tepartment, Hungarian Natural hemia (Buzek Neogene Hun- et al., 1990), in the of I Museum BP lliston Collection, and 63.1039), the gary (Andreanszky, and Miocene 1959), in late tin- Ccske slredohoff Mountains (Kvacek Walthcr. Abkhasia Mahonia of (see Takhtajan, 1974). Ox hil- — MO same showing Yunnan, China, 52556: A Henri 11907. B. Transverse section of tin- fruit. tilinclolia I (Manchester! comb. chambers; luo large and cmptv. one smaller ami containing a seed. C-F. Toricellia bonesii — Clnmo \ul Beds. Oregon Iransvers. section showing sinall central chamber and larg< late In I'i F..—..-.-I1 ( — UF 9578. D. Delail ihc p specimen in enlarged In show isodiametric sclereids. K. Iransvei > i>l -^ <>l ( .. 1 1 1 1 1 1 - seelioned s—pecimen Intnl.. e .iniiiig a seed, and two larger lateral chambers filled with sedin In. v. -,; -.i.!,.ll i , UF stripped awav showing two symmetri, 9577. Dorsal surface of a silicified fruit with the pericar]. partially F. placed chambers, and the median chamber with a small facet corresponding to the germination valve, holot lateral Sedin F')2;!!i. -(;. I A -alb ng.-M kva<". k k Mi..,-. n,« ,',,/,-,/ i! I'.!./., 1'r.nii M. of Bilina, western Czech Republic, is similar to impl nold from the Oligocene of western North \i & Middle locale (The coal southwestern America (Kvacek Bfizek, 1994). field ill & Mahonia Honshu: Huzioka Takahashi. and from Although most diverse in Asia today, is 1970), Kamchatka Budanfsev m.sid- poorly represented in the Asian fossil record. Tauai strata in that 1<)<>7| | Mahonia and Suzuki (1963) described lanveojolia lale first I i Eocene Bournemouth lug from the Miocene of southwestern Hokkaido, corded from the lied- of bill land (Chandler, 1963). Mahonia may he questioned. Basel on Betula, which widespread in the Northern is affinity to Mahonia Hemisphere has commonly been reported appears todav. in the available fossil record, that it because spread from North America Europe in the early the fossil record on the basis of leaves, but to between Betula and Tertiary and to Asia in the late Tertiary. Its arrival of overlap in venation pall, ins such remain Asia may have been either from Europe or North other genera of the Betulaceae, reports in speculative unless accompanied by the diagnostic America. Betulaceae. The birch family has an excel- trilobed infructesoeiiee brails. North \nicno.i Iii Hemisphere Betula documented by such bracts lent fossil record in the Northern the earliest is & Wehr Eocene (Crane, 1989). Alnus, which distributed today B. leopoldae Wolfe from the middle is & around the Northern b mispl ere d -oub lb< of One Mile Creek, British Columbia (Crane ai '.< I Subsequently, another species with Andes, oufii ied on of inirael Stockey, 1987). is is 'I ( 1 i V Wy- / docu- and associated leaves in the early Eocene in -wherry, is I I i . • . oming and in the middle Eocene of Oregon (Crane, mented in the Bridge Creek flora of Oregon (Meyer o . & common Manchester, l<>97). In Vs,a Hrtula aheadv helnlacei'iis tms, d. !».!,, col \\\- is ,u « confirmed by bracts in the Paleocene of the Tak- omiug with elongate, cone-like infriiclescences hobe flora in Sikhote-Alin (Akhmetiev, pers. coiiini. bearing small wingless mil-. \s this germs has not ( >'»7). In Europe. earliest example confirmed been recovered from regions outside North Ameri- |||«- I by bracts is in tin- early Oligocene of Markvartice, ca, may have been endemic to North America. it Czech Republic (Buzek et al., 1978). Ostrya, which occurs in North Temperate areas Carpinus occurs in North Temperate areas today, today, readily recognized by the persistent blad- is and is easily identified by the enlarged asymmet- derlike involucre which surrounds the nutlet. It is rical leaflike bract attached to the ribbed nutlet. first observed in the early Oligocene of central Eu- The genus common & the Tertiary of Asia be- rope (Engelhardt, 1885: 320; Kvacek Walther, i- 111 lemma ginning in the late Eocene (Tana 1972; 1998, pi. 12, figs. 6, 7) and western North America i, & & Tanai, 1993). In Europe the genus confirmed (Meyer Manchester, 1997) and Miocene is in the by two kinds of bracts from the Middle Eocene of China (WGCPC, 1978) and Japan (Tanai, 1972). & Eckfeld flora (Wilde Frankenhauser, 1998) and absent from northwestern North \merica \ m common continues element Oligocene and as a today, the genus occurred as late as the middle & m vo later floras (Berger, 1953; Roiron Vernet, 1978; Miocene h.mcv \\elrod. >u )i c\ ( i ! much Mai, 1995). It is more rare in the North 1959; Fields, 1996). In Asia, Ostrya fruits occur in \nieiiean Tertiary. Possibly the oldest record is an the fossil record by the middle Miocene (llu/ioka. WGCPC, undescribed species from the middle Kocene of lb 1963; 1978; Tanai, 1961). Washington public, (Wehr, 1995: Crane pi. 3, fig. 3). It I'nl, ic<>( aijiinus. is an extinct genus of fruit cotilorms Carpinus and to in the size rtioi ;di. with bract characters similar to Corylus and nutlet characters similar to Carpinus. was circumboreal It ration of the bract, but tin brad differs from mod- during the Paleocene, with occurrences England in ern species in the obovate, rather than ovate, out- (Crane, 1981), France- (Crane. 1989). China (Man- & line. Curiously, Carpinus appears to be lacking chester Guo, 1996), and North America (Crane & & from the subseqn. ill n n >rd of North Amer- et al., 1990; Sun Stockey, 1992; Manchester I. No Chen, 1996). Palaeocarpinus persisted into the Ko- I \ineiiea lo,la\ indicate-, eilhei thai the genus per- cene in western North America (Republic flora: sisted on this continent after the Eocene without Wehr, 1995: and eastern Asia (Buoy pi. 3, fig. 4) leaving a record, or that the Eocene pomdaiiM Sikhote Alin; Akhmetiev & Manchester, tin in perished, followed by a later Tertiary or Quaternary progress). recoloni/alion born Kurope or \sia. Kea\ es for mci- Caprifoliaceae. Diplodipelta from the Eocene ly referred to Carpinus from the late Eocene and to Miocene of the western ruled States an ex- is I Oligocene western North America now are ol & placed in Parararpinus, and are considered the (Manchester Donoghue, 1995). Both genera have probable candidate lor the foliage of ihe extinct gale fruits with persistent epigynniis sepals, but I i fruit Asterocarpinns (Manchester *\ Crane, 1987; the dispersal units differ. Dispersal units ol );>*'• I I & Meyer Crane, 1989; Manchester, 1997). consist of a single fruit subtended by three wings Corylus, which occurs North America, Europe, K in |.i from the inflorescence lna< ' I ' and \sia today, is first confirmed on the basis of n M \e three bract-derived is - mi- Is,, i- i, i | nuts with multilobed, both foliar and spiny bracts wings, but the involucre subtends pan a ol fruits in the Middle Eocene of Republic, Washington rather than just one. The occurrence Dipclta ol (Wehr, known 1995). Corylus-like leaves are from based upon fruits in the late Eocene of England & many Paleocene and Kocene around sites the (Reid Chandler, 1926) indicates that the genus Northern Hemisphere, but is virtually impossible was not always confined to Asia and apparently it determine whether to thev are the leaves of Corylus, crossed diredb between Europe and Asia. In ad- or I'alaro.arptnus. Hence, they are best placed in dition, the morpholnmi similarity between Diplo & genus Gardner the fossil leaf Corylites (Boulter dipelta and Dipclta suggests early Tertian geo- & Kvacek, 1989; Kvacek 1994; Manchester et al., mtinuity. Chen, Man- 1996). Silicified nuts of Coryloides Cerei.lipliyllaeeae. Cercidiphyllaceae are chester from the Kocene Oregon resemble of Cor- represented by one living genus with two species ylus in morphology and anatomy, but differ from native to eastern Asia. The family has an excellent modern species b\ being perfect 1\ spherical (Man- fossil record, with leaves and fruits extending back chester, 1994b). the Cretaceous. Cretaceous and to late early Tertiary & Cranea Manchester Chen (1998) is an extinct leaves are usually placed in the fossil ». us ' ' >, i Volume Number 2 86, Biogeographical Relationships ndroi les Berry (Crane, 1984). Fruits of Nyssi- P) ( )(.|. In the \si Teil ar\. llie genus is confinned i dium Heer (s\n h by clustered fruits and associated foliage in the \ Miocene Kamchatka of Zaliv Korfa, eastern (Che- droides leaves, are pod-like follicle- sim- lebaeva, 1978). Cornaceae. The ilar to those of Cercidiphyllum, but they are Cornaceae sensu lato (includ- obliquely striated, and are borne in elongate ra- ing N\ssa< Ma-M id.'a uigiaeeae] have "• . cemes Hemi- ralher llian clusters (Crane, 1984). Nyssi- an e\c<'llcnt fossil record in the Northern common (i;un! - die l;ilc avlaeeous. P.d. ic.-m sphere dial uielm!. Poll] e\laul an e n, .-„ ( i.i I . : , i i , ami Pocenc North \mciic.a. tin- late Cretaceous Alangium distributed today from China to <if is Paleocene Palo, and Alangium lo of \~ ic ei .astern Australia in tropical \frica. i i i i i of Europe, Greenland, and Spitsbergen. The south- leaves are striking!) convergent shape and in llie ernmost occurrence North America from priman. secondary, and venation in is tertiary to the UF Hope, Arkansas (UF loc. 18607). leaves of some genera in the Malvales. and there At one Paleocene Can- locality in the of Alberta, ada, Nyssidium fruits, Trochodendroides leaves, and lure. However, ihe eridocarps are vers .liagno-he asso. seeds seedling Here placed |og< \ittngium foss remains lai.'.l ;j)hI ill liai- u. iical Iriiil in i, \ ( [ 1 1 1 | er as a single species in as. the Tertiary of North America, Europe, and Asia . & Crane Stock- .vhere, the more (Eyde et al., 1969). In North America species are I eonser\ati\e known from Eocene Oregon (Manchester. pi -. n n n the of _ t i I i to the fruits and leaves continues. 1994b) and Miocene of Vermont (Eyde et 1969). al., Detailed multiple organ investigations on Nyssi- Europe, ranges from the Eocene of England In it r/n/m-related plants from Paleocene localities in (Chandler, 1961) and Germany (Mai, 1970) to the & anudn. ug ami, and Pat Pastern Russia uveal Pliocene in Alsace, France (Geissert Gregor, « I' that these plmils were verse in phvllolaxv, -hoot 1981). In Asia, fruits are known from the Pliocene ,\ & m growth, and u -r. n n No. and Pleistocene of Japan (Miki, 1956; Miki Ko- .\ fl< ( i i 1985b). from Alberta had oppo- kawa, 1962; Eyde 1969). The ey, Joffrea speirsii et al., fossil fruits all - |e ,,:a ;pc)s> |»P also alaa aatci on long u d ml to section Marlea, which occurs in east- ;,| - ., ,;. I I short shoots with inflorescences in the leaf axils of ern Asia and Indonesia today (Eyde et 1969). al., UK.riopodial -hoit -liool-. \ v.v>,'. /».://, ,trr!,. n::: liom Infructescences and fruits of the extinct genus & the Paleocem long shoots with Amersinia Manchester, Crane Golovneva nil.ii (1999), i.i pseudnw whorled. or hoik -d. leaves arid lejtnuin n relate.! lo ihe exlnni ihinese ei deiiu, - !)<tt uji.i and « The florescences. plant referred to a- 7 -- mi n ,\iin a\e- for- ' I. i i & drocarpus amicus (Krassilov, 1976; Crane Stock- merly called lioth North in I - 1985b) has alternate phyllotaxy and bears America and Asia. The infructescences are heads ey, nee- in the leaf axils of long shoots, \i of tricarpellate fruits with four or five prominent though is clear that infructescences and fruits liraet sears on ihe peduncle below the head. Per- it corre-potidi:i<; lo V i.x<,,v/m»> were widespread in die mineralized fruits have been sectioned lo show iri- d Northern Icmisphcre, more work need. ioeular endocarps lacking an is lo ile -. -d of jiheis, I . ( , f , i ] lermiiM the geographic ranges of die different -p< axial iHllldle. u d n nig -u git -.-. .led locales Willi c. examph cies. For nai ie.a h. ipieal valves, fhese fruits, and I i ,i i 1 1 , i I - I II Ii Manches- plants with the Joffrea speirsii type of growth ar- the associated foliage, Beringiaphyllum & chitecture occurred outside North America. ter, Crane Golovneva (1999), occur together at < Rocky Mountains ,li\!l>i:!:. i,ati\. in Inn n>l lapan. eight localities in the of the U.S. ( ,<i: :. >: r. < i t oguized on the basis of chisl. and Canada, in northeastern China, and in southern r< . nd i— the extant spe- Primorye, Koryak Highland, and western Kam- io-< of 1.( i The cies in the lower lioth n chatka. Ifnssia. apparent absence of Amersinia mi i >l i i< I >i ii I & central Europe (Meyer Manchester, 1997). In from the Tertian Europe, Greenland, and Spitz- ol North America continues through the Middle bergen suggests thai ihe genu- dispersed across it & Miocene of Idaho (Smiley Rember, 1985). In Eu- Beringia but did not populate the North \llantic oi known leaves are Turgai region. < i from the Early Oligocene Pliocene (Jahnichen Cornus occurs North Temperate regions, and to et in \mcnca The al., 1980; Kovar-Eder et al., 1998). Staminate flow- e\tei d- into Soiill and \lri. a today. found with the and leaves the Miocene Pah en North America fruits in ins i- -ii a i'l's if < i i i i m of Ii, :n di ,r In in tanl species by the de- both from leaves (e.g., Cornus Inpnhorea Heer. I i i • & velopment of a perianth (Kvacek Konzalova, Hickey, 1977: 144, pi. 47, fig. 1) and fruits (Crane Garden Botanical i et al., 1990: fig. 31G-I) in the Paleocene of North onomic levels lo be accorded different laxa wilhin Dakota. Leaves are readily identified to the genus the maslixioi.ls are not agreed upon. Murrell -I 11 i because recommended (sciisii latoi (In- -sinootliK curving aerod- (1993) that Diplopanax be treated as <>f roninus -<-c.. ml. tries, lliin. widolv spaced, percur- a section within Mastixia. this proposal were to II rent tertiary veins, and entire margin, hut discrim- be accepted, then most the laxa now treated as ol genera (Mai, 1993) would need cue subsumed rcprodiielive male-rial. Paleo, Citrnn.-, fruits horn lo be within the extant genus Mastixia. North Dakota, and those described from the Early \\ Inchever approach might be taken in the future. & Eocene England of (Reid Chandler, 1933, as is clear that the fruit morphological diversity of it Dunstanea) arc auatomiealK pi-served, showing this complex was much grealei the than in I'citiarv resin cavities in the wall thai nidi, ate affinities w illi Ohem the Cornelian group dogwoods. Thus, Langtonia one ol of the il,, is i a.rnelian iherry group s These hilo.ailar ellipsoid fruits ( ( . North Atlantic during the a K l'ertiar\. I'm.I.h. conform to the Mastixioideae by having an endo- < i . dogwoods composed of the hig-hracted provided the earp of tortuous single-seeded is fruit fibers, loc- 1>\ of Cornus clarnensis from the Eocene of Oregon, ales, and elongate germination valves. |'he fruit dif- which resembles extant C. florida (Manchester. fers from other modern and loss,] mastixioids b\ 1994b). the occurrence ol paired dorsal infolds in each loe- Nyssa, with a disjunct distribution in eastern llle. giving a \\ -shaped cross section of the looiile \sia. eastern North \iiieii. a. and Central \mciiea. and seed in contrast to the usual - or V-shape. I has an excellent fossil record in the Northern This genus was lust described based on specimens Hemisphere (Eyde, 1997). Nyssa fruits have woody from the Karl) Eocene of England (Reid & Chan- composed stones of fibers and have one to three dler, 1933) and was subsequently recognized in the loeules with apical-dorsal germination \al\es. middle Eocene of Oregon (Manchester, 1994b) and Transverse sections of pcrinineralized specimens Paleocene Wyoming Haggard, of (Tiffnev 1996). cC reveal that the endocarps are composed of tortuous Elaeocarpaceae. Sloanea occurs today in libers, lack an axial bun. and possess apical dot tropical and subtropical America, Asia, and Aus- lie. valves— s\ndromc germination spun sal a diagnostic of Fruits of four valve- correspond- tralia. to five \ t.v.si/. The genus is well c| lies, a led in he o< cue ing lo those o( extant Sloatira occur in the I'alen- i i I I and later I'ertiaiv ol North \ineriea (Manchester, cetic ol North \iiieri. a. where llicv have gone under 1994b) and Europe (Mai, 1995). In Asia, extends the name Carpalithrs 2A-C; New- s/>mo.we> (Kig. it from the Oligocene to Recent (Eyde, 1997). berry, 1898: 138, pi. 68, figs. 2, and in the 3), Mdsti.Mu. which occurs in Asia today, is well Paleooeue of Oreenland (Kig. LM>. K: "Cistanra lin- known lor its excellent representation In fruit- in ger?' in Heer, 1869: 470, pi. 45, fig. 2). These re- the European Tertiary, extending from the Eocene & (Reid Chandler. 1933) to the Miocene (Kirchhei- into the Kocene of the Kockv Mountain region. Slo mer, 1957; Mai. 1993). Mastixia endocarps are an- ancaccarpum a genus based on similar spiny is a alomicalb miiiiI.ii to \\ssa. but the endocarps have valve from lower Olijiocene ol Hungary (Rtiskv. 1« 1 1 more pronounced dorsal infolds, resulting loo- 1962). in Eucommiaceae. Eucommia, native only to central China lodav. has di-tm. live samaroid fruits endocarp. lathei than being confined to the apical (Fig. 3A). Such fruits have an excellent record in end Mastixia also occurs in the Kocene of Oiegon the Kocene of the western and southeastern United & and & California (Manchester. 1994b; Tiffney Hag- States (Kig. 38: Call Dilcher. 1997), and extend- gard, 1996). In addition to Mastixia itself, there is ing into the Migocene 01 Miocene of southern Mex- ( & complex closeb dated genera mas- a ol cxlincl ol ico (Fig. 3C; Magall6n-Puebla Cevallos-Ferriz, i. Europe The tixioids in the Tertiary of (Kirchh. imer. 1994a). generic determination of these fruits is 1936, 1957; Mai, 1993) distinguished by differenc- unquestionable because of the unique morphology and venation, and the presence of latex strands ob- sence of gum/resin servable the fossils u enriespond position « III || III | tending back Cretaceous to the late (Maastrichlian: to the laticifers of extant fruits (Szafer, 1954; Tra- & & Knobloch The Mai. 1986). living mastixioid ge- lau, 1963; Call Dilcher, 1997). In Asia, Eucom- i.u- 'in\ i- a native of \sia today, but has mia is well documented b\ fruits in the Kocene of fossil octillion, es in North America (Slockev et al.. Yubari, Hokkaido, Japan (Huzioka, 1961), the low- & 1998) and Europe (Eyde Xiang, 1990). The tax- er Oligocene of Kiin Kerish. Ka/akhsian \klune- I Biogeographical Relationships 1991), and the Miocene Kraskino flora of lau, 1963; Mai, 1995). The North American fruits r, asan Basin, south Primorye, Russia (Ahlaev et are about half as large as fruits of the extant species 1993). The numerous fruit records in Kurope ami are -liuhtK rn< t a-\ nmielrieal in the place- . & end from the Oligocene to the Pleistocene (Tra- merit of stigma the apex (Call Dilcher, at fruit ;vw — China, A: P. C. Silvestri 7378. B. Eucommia eocenica (Bern) Brown from the Bo\a\ Cla\ Mi^i^ippi. 137.57- — I'il. I I- M & Eucommia 8218. C. I, g n Puebla Cevallos-Ferriz from the Oligocene or Miocene Pie Vara For tie ill = .nation. I'uehla. Mexieo. I(,\1-I'H 237:5 t,V2. Seal.- bars cm. ]

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