A commented synopsis of the pre-pleistocene fossil record of Carex (Cyperaceae).
Carex graceii J. Thomasson [FS], in Am. J. Bot. 70: 437 (1983).Type material: Holotype: J. R. T. Site 50, "seed" clusters locality X, in Scout Canyon, NE 1/4, Sec. 5, T1 5 N, R42W, Garden County, Nebraska, J. R. Thomasson BP483a (exact topographic data in files of FHKSC). Illustrated in Thomasson (1983, SEM), Figs. 1, 2, 3, 4 and 5.
Distribution: USA (Nebraska).
Locality records: Thomasson (1983, SEM).
Time interval: Middle to late Miocene.
Kind of remains: Achenes and utricles, permineralized.
Carex gurnardi E. Reid & M. Chandler [FS], Catalogue of Cainozoic Plants in the Department of Geology, vol. 1, The Bembridge Flora 77 (1926).
Type material: Bembridge, Isle of Wight, United Kingdom, Holotype: VI7556 (NHMUK). Illustrated in Reid and Chandler (1926, photo); Plate IV, Fig. 19.
Distribution: United Kingdom (Isle of Wight).
Locality records: Reid & Chandler (1926, photo).
Time interval: Late Eocene (Priabonian).
Kind of remains: A putative utricle, no achene (Mai, 1999).
Carex major V.P. Nikitin [FS], in Paleocarpology and stratigraphy of the Paleogene and Neogene strata in Asian Russia 163 (2006).
Type material: Holotype: Middle Miocene (Langhian) beds of the borehole No. 34 near the Kruglikovo village, Novosibirsk oblast'--NG, collection 06.56-34-19.5, specimen Cl-4/2. Illustrated in Nikitin (2006, drawing); Plate 20, Fig. 15.
Locality records: Nikitin (2006, drawing); see also Appendix SI.
Distribution: Russian Federation (Western Siberia).
Time interval: Oligocene-Miocene boundary to Middle Miocene; doubtful from the early Pleistocene (Gelasian).
Kind of remains: Achenes and utricles.
Notes: The original illustration in the protologue does not provide enough information to systematically place this species. For this, the revision of the original material is needed.
Subgenus Vignea
The most typical achenes displayed by species from subgenus Vignea have their widest portion below the middle and mostly have a two flexa (abrupt changes in curvature) along their sides (Fig. 3a). However, it is a character that many species and sections do not share. Achenes narrowly oblong are found in the sections Ovales and Physoglochin, a character shared with some species from the Caricoid clade (see the last epigraph at the end of the checklist for additional information). Almost all the species in the subgenus display biconvex achenes. Those species with 3-stigmas (C. gibba Wahlenb. (section Gibbae Kuk.), C. kobomugi Ohwi and C. macrocephala Willd. ex Spreng. (section Macrocephalae Kuk.)) display achenes from biconvex to obtusely-trigonous. What does seem to be a character somewhat restricted to this subgenus is the shortly cylindrical style base, jointed to the not-lignified style (Jimenez-Mejias & Martinetto, 2013; Fig. 3a), a character that seems to be very rare in subgenus Carex (Villaverde et al., in prep.). A few sections within the subgenus display a more elongated cylindrical style base (e.g., sections Glareosae and Ovales; Villaverde et al, in prep.) but such a structure culminates in a clear-cut junction with the not-lignified segment of the style.
Section Incertcie sedis
Carex spp.
Distribution: Belarus, Poland, Russian Federation (European Russia, Western Siberia, Yakutia).
Locality records: Dorofeev (1960c; 1962b; 1963, photo; 1965; 1966a; 1967a; 1972, photo; 1977, photo; 1979), Lancucka-Srodoniowa (1979, photo), Dorofeev (1988, photo); see also Appendix SI.
Time interval: Late early Miocene to late Pliocene (Piacenzian), doubtful from the Oligocene.
Kind of remains: Achenes.
Notes: Shape and also some additional characters of the remains figured by Dorofeev (1963, 1972, 1977, 1988) and Lancucka-Srodoniowa (1979), as well as the specific comments provided by the authors, indicates affinities of these remains to subgenus Vignea.
Carex acutimontana Mai [FS], in Palaeontographica Abt. B, 250 (1-3): 38 (1999).
Type material: Holotype: Bhg. Oberoderwitz 1/69, 128 m (Florenzone VI, Untermiozan), MfN Berlin (Slg. Mai, Nr 1993/7406a). Illustrated in Mai (1999, photo); Plate 21, Fig. 3.
Distribution: Germany.
Locality records: Mai (1999, photo).
Time interval: Early Miocene.
Kind of remains: Achenes and utricles.
Notes: Utricle and achene characters, apart from the style, resemble C. disticha and, according to Mai (1999), also extant species of sections Ovales (C. cristatella Britton, C. tribuloides Wahlenb.) and Vulpinae (Carey) Christ, both from subgenus Vignea. However, the studied material of C. acutimontana lacks the diagnostic style character of having a jointed, shortly cylindrical style base of subgenus Vignea, but it is definitely non-jointed until 3/8 of the achene's length. The many-nerved utricle in one of the types is nearly complete and shows a short beak, ca. 1/15 to 1/20 of the utricle length. None of the modern species observed by us combines a many-nerved utricle, short beak, biconvex narrowly elliptical achene and style non-jointed until to 1/3 or 1/2 of the achene's length. This species may be a stem-taxon of the Vignea clade, with ambiguous style-base. See also notes under C. loliacea for materials with possible affinities with C. acutimontana.
Carex helmensis Mai & H. Walther [FS], in Quartarpalaontologie 7: 84 (1988). [Fig. 5k]
Type material: Holotype: Berga (Pliozan), MMG, Dresden, Coll. Mai, Nr. 5602. Illustrated in Mai and Walther (1988, photo); Plate XII, Fig. 11.
Distribution: Germany.
Locality records: Mai and Walther (1988, photo).
Time interval: Pliocene.
Kind of remains: Achenes.
Notes: One of the two type specimens has the style junction clearly marked at the very apex of the achene. The achene's surface cells are compatible with an affinity to C. canescens and other species of section Glareosae, but possibly also to other species of subgenus Vignea. In our opinion the characters available in the two types are so few that they do not allow yet a more accurate placement than to subgenus Vignea. Only two other achenes from the same original locality (Berga) are represented in the MfN collection. It is a certainly rare species occurring late in the Cenozoic of Europe.
The record by van der Burgh and Zetter (1998) is here tentatively placed under C. klettvicensis.
Carex marchica Mai [FS] in Palaeontographica Abt. B, 250 (1-3): 41 (1999)
Type material: Holotype: Bhg. Staakow 157/62, Spremberger Schichten, Untermiozan (Florenzone III), MfN Berlin (Slg. Mai, Nr. 1993/1766). Illustrated in Mai (1999); Plate 22, Fig. 4.
Distribution: Germany, Poland.
Locality records: Zastawniak (1992, photo) [as C. cf. ungeri; cf. Mai (2000)], Mai (1999, photo), Mai (2000, photo).
Time interval: Early to late Miocene.
Kind of remains: Achenes.
Notes: Mai (1999) noted affinities of this fossil with the extinct C. elongatoides and the extant C. elongata. He also reported affinities with C. paniculata L. However these can be completely ruled out after studying the holotype of C. marchica. Carex paniculata displays somewhat inflated achenes, narrowed at the base and often accompanied by spongy utricle remains. Conversely the achene of C. marchica is quite different, with its biconvex and flat, not inflated achenes. The short style base, ca. 1/ 15 of the achene length, supports assignment of C. marchica to subgenus Vignea. It is sufficiently distinct from the other species described in the region, but the distinction from C. elongatoides seems to be only based on the smaller dimensions and should be reevaluated in more detail.
Carex protogaea Mai [FS] in Palaeontographica Abt. B, 256: 38 (2000)
Type material: Holotype: Tagebau Kausche-Klara II, Hangendtone (Obermiozan, Florenzone XIII), MfN Berlin, Slg. Mai Nr. 1993/4697c. Illustrated in Mai (2000, photo); Plate 15, Fig. 6. [non Mai (2000, SEM); Plate 15, Figs. 7-10 (SEM); see comments below].
Distribution: Gennany.
Locality records: Mai (2000, photo).
Time interval: Late Miocene.
Kind of remains: Achenes.
Notes: There is an error by Mai in the assignment of the achene of Plate 15 Figs. 7-9 (Mai, 2000) to this species. It is obviously a trigonous achene belonging to another species. In the type (Mai, 2000: Fig. 6) the style junction is ambiguous--the style may be broken and in this case it could be conspecific to C. acutimontana whose achenes are not very visible because they are covered by the utricles. From C. helmensis, C. protogaea differs in having a thinner achene and a regular elliptic outline, and from C. marchica in having a larger size. It should be kept as a separate species pending discovery of more complete materials.
Section Ammoglochin Dnmort
Carex brizoides L. [BS] -type
Distribution: Italy.
Locality records: Martinetto (1994b), Martinetto et al. (2007).
Time interval: Late Pliocene
Kind of remains: Achenes.
Notes: Some of the Italian fossil achenes are nearly (if not completely) identical to the type material of C. ungeri Mai and Walther (1988). In the richer Italian assemblage there is a very good agreement in all details and morphological variation between the Pliocene achenes and the extant C. brizoides, so that placement in section Ammoglochin seems to be well assessed. However, it should be verified that this same morphology does not occur in other sections.
Carex colchica J. Gay [BS] -type
Distribution: Germany.
Locality records: van der Burgh (1978, photo; 1983) [as C. ligerica].
Time interval: Middle Pliocene.
Kind of remains: Achenes.
Notes: Taxonomical affinities to be confirmed.
Carex ungeri Mai & H. Walther [FS], in Quartarpalaontologie 7: 87 (1988) [Fig. 3a and b]
Type materials: Holotype: Rippersroda (Pliocene), MfN Berlin (originally published as at MMG, Dresden), Coll. Mai, Nr 1135. Illustrated in Mai and Walther (1988, photo); Plate XII, Figs. 18-19.
Distribution: Germany; doubtful from Poland.
Locality records: Mai et al. (1963, photo) [as C. elongata; Mai and Walther (1988)], Mai and Walther (1988, photo). Doubtful records at Lancucka-Srodoniowa (1979) [as C. cf. elongata-, Lesiak (1994)] and Lesiak (1994, photo).
Time interval: Pliocene; doubtful from middle and early Miocene.
Kind of remains: Achenes and utricles.
Notes: The visible characters of the type material are shared with C. brizoides L. (Fig. 3a-d), a species that is represented only by immature reference material in the MfN modern collection, which could explain why Mai and Walther (1988) did not cite the affinity to it. See also notes in C. brizoides-type. The record in Zastawniak (1992) has been reinterpreted as C. marchica (Mai, 2000).
Section Chordorrhizae Meinsh
Carex chordorrhiza L.f. [BS]--like
Distribution: Canada (Nunavut).
Locality records: Matthews and Ovenden (1990).
Time interval: Pliocene.
Kind of remains: Achenes.
Notes: Taxonomical affinities to be confirmed.
Section Elongatae (Kunth) Kuk
Carex elongata L. [BS] -type
Distribution: Czech Republic, Poland, Russian Federation (European Russia).
Locality records: Szafer (1954), Dorofeev (1962b, photo), Knobloch (1989, photo).
Time interval: Pliocene.
Kind of remains: Achenes.
Notes: The records listed under this name are needed of revision and do not testify the sure occurrence of C. elongata prior to the Pleistocene. They may have affinities to C. elongatoides. Indeed a report from Mai (1965) [as C. cf. elongata] was reinterpreted as C. elongatoides (Mai & Walther, 1988). Lesiak (1994) also suggested that the record of Lancucka-Srodoniowa (1979) [as C. cf. elongata] may be this species, although we consider it in need of revision. An earlier report from Mai et al. (1963) was reinterpreted as C, ungeri (Mai & Walther, 1988).
Carex elongatoides Lanc.-Srod. [FS] in Acta Palaeobotanica 20: 84 (1979)
Type material: Nowy Sacz I (268, 269, 280, 303), Nowy Sacz II (33). Illustrated in Lancucka-Srodoniowa (1979, photo); Plate XIV, Figs. 4, 5 and 6.
Distribution: Italy, Germany, Poland; doubtful in Austria.
Locality records: Mai (1965, photo) [as C. cf. elongata; see Mai & Walther, 1988], Lancucka-Srodoniowa (1979, photo), Mai and Walther (1988, photo), Zastawniak (1992, photo), Martinetto (1994a), van der Burgh and Zetter (1998), Mai (2000, photo), Cavallo and Martinetto (2001). Doubtful records at Czaja (2003, photo) [as C. cf. elongatoides], Mai (2004, photo) [as C. cf. elongatoides], and Meller (2011. photo) [as C. cf. elongatoides].
Time interval: Late early Miocene (late Burdigalian) to late Pliocene (Piacenzian), and possibly early Pleistocene.
Kind of remains: Achenes and utricle remains.
Notes: The assignment to subgenus Vignea is well supported by the style junction close to the achene's apex, and achene outline and epidermal cells size are compatible with an affinity to C. elongata L. (section Elongatae), although further study would be needed for a more confident sectional placement. Also, the possible transition between C. elongatoides and C. elongata warrant further study.
Section Glareosae G. Don
Carex sp.
Distribution: Poland.
Locality record: Lancucka-Srodoniowa (1979, photo) [as C. sp. 1],
Time interval: Late early Miocene (Burdigalian).
Kind of remains : Achenes and an utricle.
Notes: Lancucka-Srodoniowa (1979) suggested an affinity to section Heleonastes (= section Glareosae). Utricle and achenes may well have affinities to C. acutimontana. The sectional affinities are in need of reevaluation.
Carex canescens L. [BS] -type
Distribution: Russian Federation (European Russia).
Locality records: Dorofeev (1962b, photo).
Time interval: Probably Pliocene.
Notes: Taxonomic affinity to C. canescens needs to be reevaluated. The report from Mai (1965) has been reclassified as C. panormitana-type.
Carex canescentoidea Mai [FS], in Mai & Walther, Abh. Mus. Mineral. Geol. Dresden. 38: 132 (1991)
Type material: Flolotype: Oberoderwitz 1/69, 128 m, Tone mit Blattern (Untermiozan, VI), Sig. Mai, Nr. 7401. Illustrated in Mai and Walther (1991, photo); Plate 17, Fig. 25.
Distribution: Germany.
Locality records: Mai and Walther (1991, photo), Mai (1999, photo), Mai (2000, photo), Czaja (2003).
Time interval: Early to late Miocene.
Kind of remains: Achenes.
Notes: In the three type specimens (achenes), the style junction is not clearly marked and the style could also be broken. However, the achene's surface cells are compatible with an affinity to C. canescens and other species of section Glareosae. Despite Mai's report of affinities to sections Glareosae, Ovules or Stellulatae (Mai, 1999), in our opinion the observed characters point to close affinities with only section Glareosae. The single achene type collection of C. remotoides (at MfN) shows strong affinities to the C. canescentoidea and these two names may be synonyms (see below comments under C. remotoides).
Carex loliacea L. [BS] -type
Distribution: Italy, Poland.
Locality records: Lancucka-Srodoniowa (1979, photo); Lesiak (1994, photo); Martinetto (1994a; 1994b, photo); Martinetto and Mai (1996); Bertoldi and Martinetto (1995), Martinetto et al. (2007).
Time interval: Early to late Pliocene.
Kind of remains: Achene and utricles.
Notes: Despite their original citation as related to C. loliacea the sectional placement needs reevaluation. The materials of all the records cited above may have affitinies to C. acutimontana.
Section Heleoglochin Dumort
Carex appropinquata Schum. [BS] -type
Notes: The record by van der Burgh and Zetter (1998) is here tentatively placed under C. klettvicensis.
Carex diandra Schrank [BS] -type
Distribution: Canada (Nunavut), Russian Federation (Western Siberia).
Locality records: Matthews and Ovenden (1990); see also Appendix SI.
Time interval: Middle Mioceneto late Pliocene.
Kind of remains: Achenes.
Notes: Taxonomical affinities to be confirmed.
Section Ovales (Kunth) Christ
Apart from the records cited under this epigraph, there is a set of samples whose taxonomic ascription to either section Ovales or the Caricoid clade is problematic. These additional records are listed at the end of this checklist.
Carex bohemica Schreb. [BS] -type
Distribution: Russian Federation (Western Siberia).
Locality records: see Appendix S1.
Time interval: Late Pliocene.
Notes: Taxonomical affinities to be confirmed.
Carex klarae Mai [FS], in Palaeontographica Abt. B, 256: 35 (2000).
Type material: Holotype: "Grube Klara II bei Kausche, Hangendtone"--MFN Berlin, Slg. Mai Nr. 1993/4720. Illustrated in Mai (2000, photo, SEM); Plate 15, Fig. 1.
Distribution: Belarus, Germany.
Locality records: Mai (2000, photo, SEM), Velichkevich and Zastawniak (2003, SEM), Mai (2004, photo).
Time interval: Late Miocene to Pliocene.
Kind of remains: Achenes and utricle remains
Notes: Affinities with section Ovales (C. muskingumensis Schwein.) have been previously reported (Mai, 2000, 2004). The type specimen is a well-preserved achene with a complete long-cylindrical jointed style-base which represents the most important character for placement within section Ovales. Such style character is also compatible with the European C. bohemica (also section Ovales), which showed a comparable length of the style-base, before the junction. New observations confirm the similarity of the style structure and cell pattern to those of North American species as C. muskingumensis.
Carex maackii Maxim. [BS] -type
Distribution: Japan.
Locality records: Momohara and Saito (2001).
Time interval: Late Miocene (Tortonian).
Kind of remains: Achenes.
Notes: The lenticular achene with ovoid lateral outline is similar to C. maackii, although sectional placement deserves further study.
Section Phaestoglochin Dumort
Carex muricata L. [BS] -type Distribution: United Kingdom.
Locality records: Reid and Reid (1907b, photo).
Time interval: Early Pleistocene.
Kind of remains: Achenes.
Notes: Taxonomical affinities to be confirmed.
Carex spicata L. [BS] -type
Distribution: Germany, Russian Federation (Western Siberia).
Locality records: Madler (1939, photo); see also Appendix SI.
Time interval: Pliocene.
Kind of remains: Utricle containing the achene, achenes.
Notes: The image provided by Madler (1939) shows a perfectly preserved utricle (unfortunately lost during World War II) that was assigned to "Carex sp. sect. Vignea". Actually several utricle characters point to an affinity to C. spicata: dimensions 5 x 2 mm, narrowly ovate outline, with progressive attenuation into a slender beak, which is bifid in its upper half. We consider this a fairly possible record of section Phaestoglochin whose occurrence in the Pliocene could be eventually confirmed by the Russian material in need of study (Appendix S1).
Section Physoglochin Dumort
Carex davalliana Sm. [BS] -type
Distribution: Germany, the Netherlands.
Locality records: von Bulow and Mai (1992), photo), Mai (2004, photo), and this paper (see notes here below).
Time interval: Pliocene to early Pleistocene.
Kind of remains: Achenes.
Notes: Additional Carex davalliana-type achenes and utricle remains are present in the MfN collection (coll. Mai, Nr. 7831) with the label "C. reidii" from the locality Belfeld (Early Pleistocene, the Netherlands; see Westerhoff et al. (1998)). It seems that Mai never described this species, that it is just mentioned in a dichotomous key (Mai, 1999). Despite his statement that these fossils represented C. paucifloroides (notes in the specimens' cards), the shortly jointed style is a better match for C. davalliana. More accurate comparative analyses of all these C. davalliana-type specimens are needed in order to decide if they represent fossil records of the modern species or a yet to be described fossil-species.
Carex dioica L. [BS] -type
Distribution: United Kingdom.
Locality records: Reid and Reid (1907b, photo).
Time interval: Early Pleistocene.
Kind of remains: Utricle.
Notes: Taxonomical affinities to be confirmed.
Section Remotae (Asch.) C. B. Clarke
Carex remota L. [BS] -type
Distribution: Italy.
Locality records: Martinetto and Mai (1996), Basilici et al. (1997), Martinetto et al. (2007).
Time interval: Supposed latest early Pliocene and late Pliocene.
Kind of remains: Achenes.
Notes: Taxonomical affinities to be confirmed.
Carex remotoides Mai [FS], in Palaeontographica Abt. B, 256: 37 (2000).
Type material: Flolotype: Tongrube Ilse bei Grossrachen, Graue Flaschentone der Raunoer Folge (Obermiozan, Florenzone XIII), MfN Berlin, Slg. Mai Nr. 1993/4506b. Illustrated in Mai (2000, photo); Plate 15, Fig. 11.
Distribution: Germany.
Locality records: Mai (2000, photo, SEM).
Time interval: Middle to late Miocene.
Kind of remains: Achenes.
Notes: Mai (2000) reported affinities of this fossil to C. remota L. We have studied the single achene type collection of C. remotoides at MfN and wc think that it does not clearly differ from C. canescentoidea. Further study is needed to demonstrate the taxonomic autonomy of C. remotoides from C. canescentoidea, and to confirm its sectional placement.
Section Stellulatae (Kunth) Christ
Carex echinata Murray [BS] -type
Distribution: The Netherlands.
Locality records: Reid and Reid (1907a, photo).
Time interval: Early Pleistocene.
Kind of remains: Achenes.
Notes: Taxonomical affinities to be confirmed.
Subgenus Carex
Species of subgenus Carex display the greatest variation in achene shape of all the major Carex groups, ranging from biconvex to trigonous, and from ovate to obovate, rhomboid or orbicular, with one to two lateral flexa (abrupt changes in curvature). Some species from certain groups display lateral invaginations (sections Lageniformes (Ohwi) Nelmes, Mitratae Kuk. (Dai et al, 2010), Phacocystis (Jimenez-Mejias & Martinetto, 2013), Indicae Tuck. (Wheeler, 2002), Rhomboidales (Dai et al., 2010), and Vesicariae (Reznicek & Gonzalez-Elizondo, 1995; Ball & Reznicek, 2002)). Some important innovations in the shape of the style base are mostly or totally confined to this subgenus, as cylindrical neck-like appendages (section Lageniformes; Dai et al, 2010), or strongly lignified elongated styles (which within subgenus Carex seems to be exclusive to the clade where sections Carex, Paludosae, Phacocystis, and Vesicariae are nested; Villaverde et al, in prep.).
Section Incertae sedis
Carex spp.
Distribution: Belarus, Germany, the Netherlands, Poland.
Locality records: Reid and Reid (1907a, photo; 1915, photo), Raniecka-Bobrowska (1959, photo), Velichkevich and Zastawniak (2003).
Time interval: Late Miocene to early Pleistocene (Gelasian).
Kind of remains: Achenes and utricles.
Notes: Shape of the remains listed here indicates affinities to subgenus Carex. Most of the records of Carex spp. do not show useful characters for sectional placement.
"Carex atrofusca Schkuhr" [BS] -type
Distribution: Italy.
Locality records: Martinetto and Mai (1996), Cavallo and Martinetto (2001, photo), Ciangherotti et al. (2007), Martinetto et al. (2007).
Time interval: Late Pliocene (Piacenzian) and possibly early Pleistocene.
Kind of remains: Achenes.
Notes: Despite these fossils being described as morphologically similar to the extant C. atrofusca (section Aulocystis Dumort.), they probably are not related to this species or its allies. This material was thought to resemble C. atrofusca on the basis of a comparison, carried out by one of the present authors (E.M.) with the photographs in Berggren (1969). The narrowly elliptic outline with distinct carpophore and straight thin style base are diagnostic characters shared between the fossils and C. atrofusca. These same characters has been observed also in C. strigosa (section Strigosae Christ; see Ercole et al., 2012). Therefore sectional affinities need reevaluation.
Carex eigensis Mai [FS] in Palaeontographica Abt. B, 250 (1-3): 39 (1999).
Type material: Holotype: Berzdorf auf dem Eigen; Hangendschichten, Untermiozan Bhg. Oberoderwitz 1/69, 128 m (Florenzone VI, Untermiozan), MfN Berlin (Slg. Mai, Nr 1993/7406a). Illustrated in Mai (1999, photo); Plate 21, Fig. 13.
Distribution: Germany.
Locality records: Mai (1999, photo), Czaja (2003).
Time interval: Early Miocene.
Kind of remains: Achenes.
Notes: The four type specimens (achenes) differ in preservation, but are compatible with being interpreted as a single species. As all the styles are broken, and the visible characters are shared by several modern species from different sections (e.g., C. distans L., section Spirostachyae, and C. rostrata, section Vesicariae), we cannot propose a sectional assignment. However, we agree with Mai (1999) that these specimens document a fossil-species not covered by any other taxonomic name in the area or time period of pertinence.
Carex gothanii Mai & H. Walther [FS] in Quartarpalaontologie 7: 84 (1988).
Type material: Holoype: Kranichfeld (Pliocene), MfN, Berlin. Illustrated in Mai and Walther (1988, photo); Plate XI, Fig. 14.
Distribution: Germany, Poland.
Locality records: Mai (1965, photo) [as C. cf. flaw, Mai & Walther, 1988], Mai and Walther (1988, photo), Mai and Wahnert (2000).
Time interval: Pliocene.
Kind of remains: Achenes.
Notes: Mai and Walther (1988) compared the single trigonous achene recovered to the Iberian C. reuteriana Boiss. However, this latter species does not have trigonous achenes and these are thin-walled and with less apparent cell pattern than in C. gothanii, suggesting that this taxonomic affinity needs to be reevaluated. The holotype is at present badly damaged (E. M., pers. obs.), although documented by good photographs. In the MfN collection this rare species occurs with a single other achene from the locality of Ruszow (Poland). The shape of these achenes is very peculiar, oblong (with broad faces whose width is rather constant from near the apex to near the base), thus having broadly obtuse apical and basal angles. A vague affinity can be suggested only with the narrower achenes of C. strigosa, but it is not sufficient to confirm assignment to a section.
Carex hartauensis Mai [FS], in Mai & Walther, Abh. Mus. Mineral. Geol. Dresden., 38: 132 (1991). [Figs. 3b and 4d-h]
Type material: Holoype: Tongrube Hartau (Florenzone VI, Untermiozan), Zentralsammlung ehem. ZGI Berlin (Slg. Mai, nr. 1047). Illustrated in Mai and Walther (1991, photo); Plate 17, Fig. 27.
Distribution: Czech Republic, Germany, Poland.
Locality records: Buzek and Holy (1964, drawing, photo) [as C. sp.], Mai (1964, photo) [as C. sp.], Mai and Walther (1991, photo), Mai (1999, photo; 2000, photo, SEM) [see notes under C. klettvicensis for additional C. hartauensis materials misidentified in Mai (2000)], Mai and Wahnert (2000), Czaja (2003).
Time interval: Late Oligocene (Chattian) to Pliocene.
Kind of remains: Achenes and utricles.
Notes: Achenes reported as having affinities to sections Hymenochleaenae Drej. ex Bailey (C. venusta Dewey) and Limosae (Mai & Walther, 1991; Mai, 1999). The holotype is a utricle with many nerves, whose achene is not visible. From the type locality only a few other remains assigned by Mai (in the MfN collection) to the same species are available. Other specimens are available from 18 Miocene sites in the same area. These assemblages bear more than 50 specimens: Klettwitz 2, Berzdorf Hg, and Kausche-Klara II (Fig. 3b). In these three assemblages utricles predominate over isolated achenes. The utricle-achene association is documented without doubt (Fig. 4d-h). The short utricle beak is mostly destroyed, except in one specimen, and this suggests that it was poorly lignified or even scarious. The achene's style is always broken at ca. 1/10 the achene's length, which does not permit a sectional assignment, yet the many-nerved utricle, the bisymmetrical achene, longer than wide, and the thin style all suggest that possible affinities could be sections Paludosae and Vesicariae. The preservation of the style just for 1/10 achene's length could indicate that the distal part was not lignified, as occurs in C. acutiformis. Such living species share all the relevant characters with C. hartauensis, but can be distinguished from the fossils by having a truncate (vs rounded) base on the utricles and a smaller length/width ratio of the achenes.
Carex hisatica Mai [FS] in Palaeontographica Abt. B, 250 (1-3): 40 (1999)
Type material: Holotype: Bohrung Hartau 1/69, Hauptmittle (Florenzone VI, Utermiozan)--MfN Berlin (Slg. Mai, Nr. 1993/7480). Illustrated in Mai (1999); Plate 22, Fig. 5 (photo).
Distribution: Germany.
Locality records: Mai (1999, photo), Czaja (2003).
Time interval: Early to early middle Miocene.
Kind of remains: Achenes.
Notes: Considered similar to the extinct C. graciosa Negru (section Carex) and the extant C. oligosperma Michx. (Mai, 1999). We studied the type material at MfN and several other sections could be considered similar. Another possible match may be the extant C. pendula, to which it agrees in shape, dimensions, style base and cell pattern. However, C. lusatica achenes display a larger shortly substipitated base with a basal callus, a character rarely observed in C. pendula or the allied fossil species C. limosioides and C. plicata. The incomplete style of all the studied materials does not allow for evaluation of affinities to section Vesicariae, to which this taxon could also belong.
Carex strigosoides Lanc.-Srod. [FS] in Acta Palaeobotanica, 20: 88 (1979).
Type material: Holotype: Nowy Sqcz I (272, 294), Nowy S3.cz II (31, 32). Illustrated in Lancucka-Srodoniowa (1979, photo); Plate XIV, Figs. 26-28.
Distribution: Germany, Poland
Locality records: Mai (1965, photo) [as C. acutiformis (Mai & Walther, 1988)], Lancucka-Srodoniowa (1979, photo), Mai and Walther (1988, photo), van der Burgh and Zetter (1998) [mispelled as "C. strichosoides"].
Time interval: Late early Miocene (late Burdigalian) to Pliocene.
Kind of remains: Achenes and utricles.
Notes: Perhaps related to C. strigosa (Lancucka-Srodoniowa, 1979). Taxonomical affinities need reevaluation.
Carex zhilinii Balueva & V.P. Nikitin [FS], in S. G. Zhilin, Paleocarpology and stratigraphy of the Paleogene and Neogene strata in Asian Russia 163 (2006).
Type material: Holotype: Lower Miocene (Aquitanian) beds of the borehole No. 170 at the Gorsky Log village, Omsk oblast'--NG, collection IIK.59-170-56.5 (H3390 Komarov Botanical institute), specimen Cl-6/3. Illustrated in Nikitin (2006, photo); Plate XX, Fig. 17.
Locality records: Nikitin (2006, photo); see also Appendix SI.
Distribution: Russian Federation (Western Siberia); doubtful from Kazakhstan. Time interval: Late Oligocene to Middle Miocene; doubful from the Miocene-Pliocene boundary.
Kind of remains: Achenes.
Notes: Possibly similar to C. graciosa. Taxonomical affinities need evaluation.
Section Acrocystis Dumort
Carex globosiformis (uglobosaeformis") Lanc.-Srod. [FS] in Acta Palaeobotanica 20: 87 (1979).
Type material: Nowy Sqcz I (272, 276). Illustrated in Lancucka-Srodoniowa (1979, photo); Plate XIV, Figs. 12a-c.
Distribution: Poland.
Locality records: Lancucka-Srodoniowa (1979, photo, drawing), Lesiak (1994, photo).
Time interval: Early to middle Miocene.
Kind of remains: Achenes and utricle remains.
Notes: Apparently shows morphological affinities with some North American extant species of the section Acrocystis (Lancucka-Srodoniowa, 1979). Lesiak (1994) recorded additional samples (C. sp. 1) as probably related to this species. Sectional placement in need of reevaluation.
Carex pilulifera L. [BS] -type
Distribution: Germany, Poland.
Locality records: Mai and Walther (1988, photo), Lesiak (1994, photo), van der Burgh and Zetter (1998), Mai and Wahnert (2000, photo).
Time interval: Middle Miocene to Pliocene.
Kind of remains: Achenes.
Notes: Sectional affinities to be confirmed.
Section Chlorostachyae Meinsh
Carex capillaris L. [BS] -type
Distribution: Russian Federation (Kamchatka).
Locality records: see Appendix S1.
Time interval: Miocene-Pliocene boundary.
Notes: Taxonomical affinities to be confirmed.
Section Carex
Carex atherodes Spreng. [BS] -type
Distribution: Russian Federation (Western Siberia).
Locality records: see Appendix SI.
Time interval: Miocene-Pliocene boundary.
Notes: Taxonomical affinities to be confirmed.
Carex carpophora Mai & FI. Walther [FS] in Quartarpalaontologie, 7: 83 (1988).
Type material: Holotype: Berga (Pliocene), MMG, Dresden, Coll. Mai, Nr. 5598a. Illustrated in Mai and Walther (1988, photo); Plate XI, Fig. 7.
Distribution: Belarus, Germany, Italy, Poland, Ukraine; doubtful in European Russia.
Locality records: Szafer (1947, photo) [as Carex sect. Frigidae], Dorofeev (1955b), photo) [as C. sp. 1 (Mai & Walther, 1988)], Mai and Walther (1988, photo), Mai (1995, photo), Cavallo and Martinetto (2001, photo), Velichkevich and Zastawniak (2003, photo); doubtful records in Dorofeev (1963, photo) [as Carex sect. Frigidae (Mai & Walther, 1988)], Negru (1986, photo) [as "C. marii-srodoniowii" p.p.; see comments below], and Mai (2008, photo) [as C. cf. carpophora].
Time interval: Miocene to late Pliocene and possibly early Pleistocene; cited as doubtfi.il from the Late Oligocene (Chattian).
Kind of remains: Achenes.
Notes: The shape of the materials from Mai's collections housed at MfN cannot be detected in any modern European species. However, the style is very characteristic, very similar to C. hirta, quite thick and not attenuated at the apex, sometimes even enlarged, with a small rounded to conical termination, often asymmetrical. Nevertheless, the style in C. carpophora is definitely more robust than in C. hirta and the termination is closer to the achene's apex; also the maximum width of the achene is closer to the base rather than closer to the apex. This supports C. carpophora as a valid fossil-species possibly belonging to section Carex.
In the original publication of the name C. mariae-srodoniowae, Negru (1986) pictured three achenes. The specimens pictured in Plate 35, Figs. 4 and 6 differ from the one in the Fig. 5 because of their more robust style, not attenuated at the apex and terminating in a rounded point. This morphology resembles samples of the extant C. hirta extracted from soil (Martinetto et al, 2014a), which show a kind of junction between the strongly lignified proximal part of the style, and the scarcely lignified distal one. The differences with the studied materials of C. carpophora are also very small. A detailed comparative study should be carried out to confmn the identity of these materials as C. carpophora. See comments under C. graciosa for further discussion about Negru's taxonomic concept of the name C. mariae-srodoniowae.
Carex graciosa Negru [FS], Meot. flora. Izd. Schtiinca: 146 (1986). [Fig. 4a-c]
Type material: Holotype: coll. num. 20I1K, Meotis, Bolshoy Gontan Odessa, Ukraine, late Miocene. Illustrated in Negru (1986, photo); Plate 35, Figs. 1, 2 and 3.
Synonym: Carex mariae-srodoniowae ("marii-srodoniowii") Negru, Maot. Flora Sevemo-Zap. Pricemonomor'ja: 145 (1986), nom. inval., illustrated in Negru (1985: Plate 35, Fig. 5; but see notes below).
Distribution: Germany, Ukraine.
Locality records: Negru (1986 p.p., photo; see comments below), Mai (1989, 1999, photo; 2000, photo, SEM) [all as "C. marii-srodoniowii"], Czaja (2003, SEM) [as "C. marii-srodoniowii"].
Time interval: Early to Late Miocene.
Kind of remains: Achenes.
Notes: Negru (1986) mentioned affinities of C. graciosa to C. flava L., but the elongated lignified style of the achenes from the type collection is incompatible with C. flava. Among the studied samples, materials from Klettwitz 2 (MfN 3804) show the most complete styles, but these are also broken at 1/2 or 3/4 of the achene length and show no junction up to this level. Instead, there is an apparent attenuation (Fig. 4b). An immature specimen shows that style lignifications occurred early in achene development (Fig. 4c). Utricles have been never found, apart from small fragments adhering to the achenes. Despite the morphological resemblance of C. graciosa achenes to those of C. hirta, placement in section Carex must not be taken for sure, and alternative systematic affinities with section Vesicariae may also be possible. In our study of the German material of the MfN we observed in some samples (especially the one from the locality Klettwitz 2 [MfN 3804]) a mixture of achenes matching the morphology of those described and pictured as C. graciosa and "C. marii-srodoniowii" by Negru (1986). Therefore, we suggest that the name C. mariae-srodonowiae (which is indeed invalid) should be considered as a synonym of C. graciosa. In the description of the name C. mariaesrodonowiae ("C. marii-srodonowii") by Negru (1986), the author failed in designating a particular material as a holotype, which therefore remained invalid. He provided instead for the C. mariae-srodonowiae collections cited in the paper two different numbers: 20ITK and 21IIK. Negru's concept of C. mariae-srodoniowae seemed to be based particularly on the achene figured in Plate 35, Fig. 5 of Negru (1986). The other two achenes illustrated in the intended original description (Negru, 1986, Plate 35, Figs. 4 and 6) may belong to a separate species, possibly C. carpophora (see comments under this name).
Carex hirta L. [BS] -type
Distribution: The Netherlands, Russian Federation (European Russia, Western Siberia), United Kingdom.
Locality records: Reid and Reid (1907a, b, photo), Dorofeev (1962b, photo); see also Appendix SI.
Time interval: Pliocene to early Pleistocene.
Kind of remains: Achenes, utricle.
Notes: Taxonomical affinities need reevaluation.
Carex lasiocarpa Ehrh. [BS] -type
Distribution: Czech Republic, Germany, Poland, Russian Federation (Western Siberia). Locality records: Rudolph (1935, photo), van der Burgh (1978, photo; 1983), Dyjor et al. (1998), Mai and Wahnert (2000); see also Appendix SI.
Time interval: Pliocene and possibly early Pleistocene.
Kind of remains: Achenes.
Notes: Taxonomical affinities need reevaluation.
Section Ceratocystis Dumort
Carex flava L. [BS] -type
Notes: Records reported by Mai et al. (1963) and Mai (1965) were reclassified as C. flavicarpa and C. gothanii respectively.
Carex flavicarpa Jim.-Mejias & Roalson [FS], nom. nov.
Nom. subs.: Carex flaviformis ("flavaeformis") Lancucka-Srodoniowa in Acta Palaeobotanica 20: 89 (1979), nom. illeg:, non Carex flaviformis Nelmes, Kew Bull. 10: 84 (1955).
Etymology: 'flavicarpa" derives from the Latin words flavus (yellow), in reference to Carex flava L., and carpus (fruit), as the fossil achenes of C. flavicarpa resemble those of C. flava.
Type material: Nowy Sacz I (268). Illustrated in Lancucka-Srodoniowa (1979); Plate XIV, Figs. 18-19 (photo).
Distribution: Germany, Italy, Poland.
Locality records: Mai et al. (1963) [as C. flava], Lancucka-Srodoniowa (1979, photo), Mai and Walther (1988, photo), Mai (1989, drawing, photo), Zastawniak (1992, photo), van der Burgh and Zetter (1998), Mai (2000, photo), Cavallo and Martinetto (2001). Time interval: Late early Miocene to Pliocene, and possibly early Pleistocene.
Kind of remains: Achenes.
Notes: Taxonomical affinities need reevaluation. General shape of C. flavicarpa remains could also match species from section Sylvaticae Rouy.
Carex hostiana DC. [BS] -type
Distribution: The Netherlands.
Locality records: van der Burgh (1987).
Time interval: Late Miocene.
Kind of remains: Achenes.
Notes: Taxonomical affinities need reevaluation.
Carex hostianoides Mai [FS] in Natur und Landschaft im Bezirk Cottbus 11: 34 (1989).
Type material: Holotype: Wischgrund-Boeschung bei Kostebrau/Niederlausitz. Illustrated in Mai (1989); Plate VIII, Fig. 13 (photo), Fig. 12a (drawing, reconstruction).
Distribution: Germany.
Locality records: Mai (1989, drawing, photo), Mai (2000, photo, SEM).
Time interval: Middle to late Miocene.
Kind of remains: Achenes.
Notes: Despite the fact that Mai (2000) considered this species to be related to C. hostiana (section Ceratocystis), he also reported similarities to C. hartmanii Cajander (section Racemosae G. Don). For this reason, the sectional placement of C. hostianoides needs to be reevaluated.
Carex lepidocarpa Tausch. [BS] -type
Distribution: Italy, the Netherlands.
Locality records: Westerhoff et al. (1998), Cavallo and Martinetto (2001).
Time interval: Supposedly from late Pliocene (Piacenzian) to early Pleistocene.
Kind of remains: Achenes.
Notes: Taxonomical affinities need reevaluation. In our revision, we detected possible affinities of the achenes reported by Cavallo and Martinetto (2001) to C. demissa Homem., a species also from section Ceratocystis.
Section Confertiflorae Franchet ex Ohwi
Carex ischnostachyaSteud. [BS] -type
Distribution: Japan.
Locality records: Yamakawa et al. (in press, photo).
Time interval: Late Pliocene (Piacenzian).
Kind of remains: Achenes.
Notes: Sectional placement uncertain; despite the fact that these remains show close similarities to this species, similar morphotypes may be found in other sections (cf. Yamakawa et al., in press). Further studies are needed to confirm the sectional placement of such materials.
Section Lageniformes (Ohwi) Nelmes
The cylindrical neck-like appendage at the apex allows to unequivocally placing these remains in this section.
Carex sp.
Distribution: Japan.
Locality records: Momohara and Saito (2001; as C. formosensis-Xype).
Time interval: Late Miocene (Tortonian).
Kind of remains: Achenes.
Notes: The fossils consist in distinct elongated-subcylindrical achenes, not found in extant species of section Lageniformes.
Carex breviscapa C. B. Clarke [BS] -type [Fig. 5e]
Distribution: Japan.
Locality records: Momohara and Saito (2001; as C. formosensis-type).
Time interval: Late Miocene (Tortonian).
Kind of remains: Achenes.
Section Molliculae Ohwi
Carex sp.
Distribution: Japan.
Locality records: Momohara and Saito (2001; as Carex sect. Extensae).
Time interval: Late Miocene (Tortonian).
Kind of remains: Achenes.
Notes: The fossil materials indicated as "section Extensae" in Momohara and Saito (2001) comprise several morphotypes of achenes with trigonous cross-section. At least part of these materials seems to belong to a species from section Molliculae; however, this designation also likely includes several morphotypes that may be assignable to other sections.
Carex aphanolepis Franch. & Savat. [BS] -type
Images: Yamakawa et al. (in press).
Distribution: Japan.
Locality records: Yamakawa et al. (in press, photo).
Time interval: Late Pliocene (Piacenzian).
Kind of remains: Achenes.
Notes: Taxonomical affinities to be confirmed.
Section Paludosae G. Don (including section Tumidae Kuk.)
Carex acutiformis Ehrh. [BS] -type
Distribution: Germany, Italy, Poland, United Kingdom.
Locality records: Reid and Reid (1907b, photo), Lancucka-Srodoniowa (1979, photo), van der Burgh (1987), Mai and Walther (1988, photo), Lesiak (1994, photo), Mai (1995), van der Burgh and Zetter (1998).
Time interval: Late early Miocene to early Pleistocene.
Kind of remains: Achenes and utricles.
Notes: We studied the materials record in Mai and Walther (1988) and Mai (1995) and they did not show enough characters for a reliable assignment to the extant C. acutiformis, so their systematic placement is in need of revision. Furthermore, the images provided by Reid and Reid (1907b) and Lancucka-Srodoniowa (1979) suggest that also their actual systematic affinities are doubtful. Remains recorded by Mai (1965) were determined to be C. strigosoides.
Carex riparia Stokes [BS] -type
Distribution: Germany, the Netherlands, Russian Federation (Western Siberia), United Kingdom.
Locality records: Mai (1965, photo), Reid and Reid (1907a, b, photo), Mai and Walther (1988, photo), van der Burgh and Zetter (1998); see also Appendix SI.
Time interval: Pliocene to early Pleistocene.
Kind of remains: Achenes and utricles.
Notes: Remains recorded by Mai et al. (1963) were determined to be C. praehirta.
Section Paniceae (Carey) Christ
Carex panicea L. [BS] -type
Distribution: Germany, Italy, Poland.
Locality records: Szafer (1947, 1954), van der Burgh (1983 p.p., photo), Cavallo and Martinetto (2001); the record of van der Burgh (1983) from Hambach is considered doubtful (see here below).
Time interval: Pliocene.
Kind of remains: Achenes.
Notes: The robust, enlarged and jointed style displayed by these samples point out affinities with C. panicea. The distinction of these records from C. praehirta does not seem to be clear. Among the samples pictured in van der Burgh (1983), all from Fortuna-Garsdorf, one sample (Plate 4, Fig. 5, achene at the left, and Fig. 7) seems to agree with these diagnostic characters, whereas at least another achene (Plate 4, Fig. 5, achene at the right) resembles C. pseudocyperits. Given this confusion, as the sample from Hambach is not pictured, we considered that its actual resemblance to C. panicea is needed of revision. In any case, further studies would be necessary to confirm the placement of these samples under section Paniceae.
Carex praehirta Mai & Walther [FS] in Quartarpalaontologie 7: 85 (1988).
Type material: Holotype: Rippersroda (upper Pliocene), MMG, Dresden, Coll. Mai, Nr 1145. Illustrated in Mai and Walther (1988, photo); Plate XII, Fig. 20.
Distribution: Germany.
Locality records: Mai et al. (1963, photo) [as C. riparia\ Mai and Walther (1988)], Mai and Walther (1988, photo), Mai and Wahnert (2000, photo), Mai (2004, photo).
Time interval: Late Miocene to Pliocene.
Kind of remains: Achenes.
Notes: After the description of C. praehirta by Mai and Walther (1988), Mai (1999) proposed its synonymization with C. graciosa Negru. In our revision we studied the type material of C. praehirta and we detected a style pattern (very robust, enlarged and jointed) that is not particularly similar to the type of C. graciosa and the extant C. hirta. Therefore, we keep C. praehirta and C. graciosa as separate species. Actually, the holotype of C. praehirta shows agreement of all diagnostic characters with C. panicea L. The achenes of this species are rather variable, although they are not known to have such a distinctly apparent substipitate base as found in the types of C. praehirta. Regardless, we consider these German fossils to be much better placed in section Paniceae. Some of the records here reported as C. panicea-type may belong to C. praehirta.
Carex vaginata Tausch [BS] -type
Distribution: Russian Federation (Western Siberia).
Locality records: see Appendix SI.
Time interval: Late Pliocene.
Kind of remains: Achenes.
Notes: Taxonomical affinities need reevaluation.
Section Phacocystis Dumort
Records published as C. acuta, C. aquatilis, C. cespitosa and C. nigra have probably been used by authors to indicate morphological affinity to section Phacocystis species rather than to denote a real taxonomic relationship with any of the particular extant taxa. Despite this, for simplicity of organization and reference to previous works we listed such records under the correspondent extant species epigraph when we could not propose a better fit. All of these records need to be reevaluated following the guidelines provided by Jimenez-Mejias and Martinetto (2013) for carpological characters in section Phacocystis.
Carex sp.
Distribution: Japan.
Locality records: Momohara and Saito (2001) [as Carex sect. Carex]
Time interval: Late Miocene (Tortonian).
Kind of remains: Achenes.
Notes: Several morphotypes are included (A. M., pers. obs.). Affinities to other Asian sections with biconvex achenes (e.g., section Gradies Kuk.) cannot be ruled out.
Carex acuta L. [BS] -type
Notes: The records by van der Burgh (1987, photo), Dyjor et al. (1998) and Mai and Wahnert (2000) [as C. gracilis] are here tentatively placed under C. klettvicensis.
Carex aquatilis Wahlenb. [BS] -type
Distribution: Canada (Nunavut and Yukon) and United States (Alaska).
Locality records: Matthews and Ovenden (1990).
Time interval: Pliocene.
Kind of remains: Achenes.
Notes: Records from Martinetto (1994a, b), Bertoldi and Martinetto (1995), Cavallo and Martinetto (2001), Ciangherotti et al. (2007) and Martinetto et al. (2007, 2014b), have been revised as C. panormitana-type.
Carex blysmoides P. I. Dorof. [FS], The Problems of the Palaeobotany 60 (1986).
Type material: Holotype: Dvorets, BIN n. 493-33; illustrated in Dorofeev (1986, photo); Plate IV, Fig. 11.
Distribution: Belarus, Germany.
Locality records: Dorofeev (1986, photo), von Bulow and Mai (1992), photo) [as Eleogiton fluitans; Mai, 2004], Mai (2004, photo), Velichkievich and Zastawniak (2007, photo).
Time interval: Pliocene.
Kind of remains: Achenes.
Notes: Considered by Mai (2004) to be related to the East Asian C. thunbergii Steud. The two achenes figured in this publication share with the narrowest achenes of the modern C. buekii all of its characters except the narrow stipitate base (see Jimenez-Mejias & Martinetto, 2013). The abundant utricle remains visible at the base of Dorofeev's (1986) achenes definitely point to the assignment to section Phacocystis. The presence of utricle remains with thick nerves (Velichkevich & Zastawniak, 2007) could also indicate similarities with other very closely related species, as C. buekii utricles are totally or almost totally nerveless.
Carex cespitosa L. [BS] -type ("caespitosa")
Notes: The records by Mai (1965), Mai and Walther (1988), von Billow and Mai (1992), Mai (1995, 2004), and Martinetto (1994a, b) were revised as C. panormitana-type (see Jimenez-Mejias & Martinetto, 2013); the records by Zastawniak (1992) and van der Burgh and Zetter (1998) have been tentatively placed here under C. klettvicensis.
Carex elata All. [BS] -type
Distribution: Italy
Locality records: Jimenez-Mejias and Martinetto (2013; photo); the record of Martinetto et al. (2012, photo) [Fig. 5h of this paper] must be considered as doubtful.
Time interval: Early Pleistocene (Gelasian).
Kind of remains: Achenes and utricles.
Notes: Jimenez-Mejias and Martinetto (2013) already confirmed the strong morphological affinities of the studied remains to the extant C. elata on the basis of both, utricle and achene, characters. These fossils from the Gelasian could indeed belong to C. elata or its immediate ancestor. The materials cited here as doubtful C. elata-type remains from the early Pleistocene (Martinetto et al., 2012; Fig. 5h of this paper) are very similar to C. klettvicensis in the observed mesoscopic characters, and may belong to a different taxon.
Carex klettvicensis Mai [FS], in Palaeontographica Abt. B, 250 (1-3): 40 (1999). [Fig. 2b]
Type material: Holotype: Bhg. Lubbenau 2/62, Spremberger Folge; Untermiozan (Florenzone III)--MfN Berlin (Slg. Mai, Nr. 1993/1668). Illustrated in Mai (1999, photo); Plate 22, Fig. 3.
Distribution: Germany, doubtful from Poland and the Netherlands.
Locality records: Mai (2000 p.p., photo [excluding Plate 14 Figs. 18-20 and 23-24, see notes here below]), Mai and Wahnert (2000). Records by Gregor (1982, SEM) [as C.jlagellata, Plate 14, Fig. 24], van der Burgh (1987, photo) [as C. acuta], Zastawniak (1992, photo) [as "C. caespitosa L. foss.", cf. Mai (2000)], Dyjor et al. (1998) [as C. acuta], van der Burgh and Zetter (1998, photo) [as C. appropinquata, C. cespitosa, and C. helmensis], Mai (1999, photo), Mai and Wahnert (2000) [as C. gracilis], and one of the achenes recorded by Mai (2004) may also belong to this species (see notes below). Other records are here re-classified as C. maiana (see notes below).
Time interval: Early Miocene to Pliocene, doubtful from the late middle Miocene.
Kind of remains: Achenes, the utricle cited in Mai (2000) belongs to C. hartauensis (see notes here below).
Notes: Mai (1999) reported this species as unambiguously belonging to section Acutae Fries (= Phacocystis). However, he erroneously recorded the "nordamerikanische [North American] Carex morrowii Boott.", a name belonging to a Japanese species of section Mitratae Kuk., as similar to C. klettvicensis. This was due to the incorrect identification of a sample in the MfN modern reference collection, very similar in outline to C. klettvicensis, but belonging to an unknown species of subgenus Vignea (E. M., pers. obs.). This error points out the necessity of maintenance, revision, and improvement of the modern fruit and seed collections (see Martinetto et al., 2014a).
The information provided by the four fossil achenes preserved in the C. klettvicensis type collection from Ltibbenau 2/62 is poor. We see similarities in the robustness of the achene and cell pattern to species of section Phacocystis. The longest preserved style base is 2/11 of the achene's length, and for this reason the affinity to subgenus Vignea may not be totally ruled out, although the achene outline of C. klettvicensis is uncommon among members of subgenus Vignea.
Carex klettvicensis seems to be a rare and poorly understood species. Only the holotype, plus two other achenes from the same layer seem to agree in relevant characters and thus confidently represent C. klettvicensis. A single achene from the Miocene of Lubtheen (Mai, 2004; Plate 8, Figs. 8 and 9; also Fig. 5g of this paper) may belong to this species. No utricle remains are known. Mai (2000; Plate 14, Fig. 24) later associated achenes from the sites Klettwitz 2, 3 and 10 and an utricle from the site Klettwitz 2 with the name C. klettvicensis, but these materials disagree with the characters seen in the holotype. The utricle was revised by Mai himself as C. hartauensis in the MfN collection, as the author possibly realized the similarity of structure to the abundant utricles of C. hartauensis that occur in the same layer (Klettwitz 2). The other achenes from Klettwitz 2 (Mai, 2000; Plate 14, Figs. 18-20, 23), Klettwitz 3 (Mai, 2000; Plate 14, Figs. 12-17) and Lubtheen 2 (Mai, 2004; Plate 15, Figs. 1, 2, 3 and 4) disagree with the holotype in having a larger size of surface cells (2035 vs 15-20 [micro]m) and a broader base, so that they are here described as C. maiana (see subgenus Carex, section Phacocystis). About the samples from Klettwitz 10 we lack enough information to refuse or confirm its identity as C. klettvicensis, so its identity remains doubtful for us. Furthermore, a lenticular achene figured by Gregor (1982, Plate 14, Fig. 24) could also belong to this species due to the narrow substipitate base. However its complete style is nearly identical to that of C. maiana, and a verification of the cell pattern would be required for a reliable assignment to either of these species.
After the examination of the materials pictured as Pliocene records of C. acuta and C. cespitosa, we found that the cited materials do not share the representative characters of these species (Jimenez-Mejias & Martinetto, 2013). We found instead similarities in the shape with the materials belonging to C. klettvicensis. Therefore, we tentatively include these pre-Pleistocene records as possible C. klettvicensis. Also the materials cited here as doubtful C. elata-type remains from the early Pleistocene (Martinetto et al., 2012) are very similar to C. klettvicensis in the observed mesoscopic characters. We encourage the revision of such materials.
Carex maiana Martinetto [FS] (Appendix 1). [Figs. 2c, 4i-j, 5i]
Type material: Holotype: MfN Berlin (Slg. Mai, Nr. 1993/3865b). Illustrated in Mai (2000, photo); Plate 14, Fig. 12.
Distribution: Germany.
Locality records: Mai (2000, photo; Plate 14, Figs. 18-23) [as C. klettvicensis], Mai (2004, photo) [as C. klettvicensis].
Kind of remains: Achenes, rarely with sticking utricle remains (scanty) showing a few very fine nerves.
Time interval: Middle to late Miocene.
Notes: The original material of this species was labeled as C. praeacuta in Mai's collection at MfN. However, it seems that this name was never formally published, apart from being mentioned in a dichotomous key (Mai, 1999). Here, we formally described this species as C. maiana in Appendix 1. The biconvex achenes with lignified non-jointed style, basal callus (very small) and 20-35 pm wide cells point to section Phacocystis. Yet the flatfish style regularly truncated in a straight breaking point (Fig. 2c) is not known in modern species, and other extant sections may be also considered for future comparison. An achene from the Miocene site Kleinleipisch-Roemerkeller (MFN coll. Mai Nr. 6190), completely covered by remains of the utricle, most probably belongs to this species (Fig. 5i). Other available remains point to a thin utricle with 2-4 thin nerves on each face (Fig. 4j). The distinction of C. maiana from C. klettvicensis is based on the larger size of the cells in the central part of the achene's faces (20-35 vs 15-20 pm), the broader base about 1/3 of the achene's width), and the differences in style base morphology (Fig. 2b and c).
Carex nigra (L.) Reichard [BS] -type
Distribution: Czech Republic, Germany, Italy.
Locality records: van der Burgh (1978, photo; 1983), Buzek et al. (1985, photo), Mai and Walther (1988, photo), Knobloch (1989, photo), von Bulow and Mai (1992), Bertoldi and Martinetto (1995), Basilici et al. (1997), Westerhoff et al. (1998), Mai (2004, photo, SEM), Kvacek and Teodoridis (2007, photo), Martinetto et al. (2007).
Time interval: Pliocene to early Pleistocene (see notes below).
Kind of remains: Achenes.
Notes: These few Pliocene or supposed Pliocene records of achenes are partly compatible with the morphology of C. nigra, but many of them lack important characters such as the shortly lignified cylindrical style-base or the conspicuous basal callus (Jimenez-Mejias & Martinetto, 2013). Also, the available materials are too scarce to perform a confident evaluation of their variation and assess a definite affinity to C. nigra. Thus, all them (except Buzek et al. (1985); see below) must be considered in need of revision. At least some these fossils may be related to C. klettvicensis. Others could represent one of more fossil-species not yet described. The rich achene assemblage described by Buzek et al. (1985) seems to be the one most reliably documenting the occurrence of C. nigra, and is probably of Early Pleistocene age (see Teodoridis et al., in press). This age is in agreement with other reliable old records of this species, which dates back not earlier than the Pleistocene (e.g., Ghiotto, 2010; Jimenez-Mejias & Martinetto, 2013).
Carex panormitana Guss. [BS] -type [Fig. 4k]
Distribution: Germany, Italy, the Netherlands, Poland.
Locality records: Mai et al. (1963, photo) [as C. vulpina, Mai and Walther (1988)], Mai (1965, photo) [as C. canescens, Mai and Walther (1988)], Mai and Walther (1988, photo) [as C. cespitosa], von Bulow and Mai (1992), photo) [as C. cespitosa], Martinetto (1994a, b, photo) [as C. gr. cespitosa], Bertoldi and Martinetto (1995) [as C. aff. aquatilis], Mai (1995, 2004, photo) [as C. cespitosa], Martinetto and Mai (1996) [as C. cf. aquatilis], Cavallo and Martinetto (2001) [as C. aquatilis], Ciangherotti et al. (2007) [as C. aff. aquatilis], Martinetto et al. (2007) [as C. aff. aquatilis], Jimenez-Mejias and Martinetto (2013, photo), Martinetto et al. (2014b) [as C. aff. aquatilis]. Newly detected in the MfN collection also for Gennany (Gorsbach, Nordhausen), the Netherlands (Obel; Fig. 4k), and Poland (Kroscienko, Mizema II).
Time interval: Pliocene to early Pleistocene.
Kind of remains: Achenes, rarely with sticking utricle remains showing several faint nerves.
Notes: Jimenez-Mejias and Martinetto (2013) already showed the strong morphological affinities of these remains to the extant C. panormitana and the allied C. reuteriana. These species occurs nowadays in the Mediterranean (Jimenez-Mejias et al., 2014), suggesting the occurrence in the past of taxa allied to them much more northwards than they are today.
Section Pomcystis Dumort
Carex pallescens L. [BS] -type
Distribution: Germany, Italy, Russian Federation (European Russia).
Locality records: Dorofeev (1962b), van der Burgh (1978, photo), Martinetto and Mai (1996).
Time interval: Pliocene.
Kind of remains: Achenes.
Notes: Although taxonomical affinities need confirmation, these records probably belong to section Porocystis.
Section Rhomboidales Kuk
Carex sp. [Fig. 5m]
Distribution: Japan.
Locality records: Momohara and Saito (2001; as Carex sect. Praecocces).
Time interval: Late Miocene (Tortonian).
Kind of remains: Achenes.
Notes: The combination of the rhomboidal outline together with the very thick strongly lignified style-base allow the unambiguous placement of these remains in section Rhomboidales (see Dai et al., 2010).
Section Rhynchocystis Dumort
Several fossil achenes assigned to the fossil-species C. limosioides and C. plicata show distinct affinities to C. pendula based on shape, dimensions, wall thickness, shortly cylindrical lignified style base, and narrow achene base. The scarce differentiation of C. limosioides and C. plicata, and the large variability of the fossils assigned to both species, deserve further investigation. In addition, C. lusatica, also have affinities to this section (see comments in the correspondent epigraph).
The fossil records listed below, together with more recent fossil records (e.g., Rinaldi et al., 2013), seem to indicate a long-lasting presence of species (perhaps chronospecies) belonging to section Rhynchocystis in Europe from the early Miocene to present.
Carex limosioides Negru [FS], Meot. flora. Izd. Schtiinca, 146-147 (1986).
Type material: Illustrated in Negru (1986; photo); Plate 34, Fig. 13.
Distribution: Germany, Ukraine.
Locality records: Negru (1986, photo), Mai (1999, photo; 2000, photo, SEM), Czaja (2003, SEM).
Time interval: Early to late Miocene.
Kind of remains: Achenes
Notes: Considered to be related either to section Limosae (Negm, 1986) or section Vesicariae (Mai, 1999). However, Mai (pers. comm. 2006) did not see the original specimens of Negru (1986) and assigned several specimens from the Miocene of Germany to this species only based on the original description and rather poor illustrations. Actually, Mai named his material "C. leporimontana n. sp." in the MfN collection, but later decided not to describe the species and assigned his fossil achenes to C. limosioides. The German sample with best preserved material is "Berzdorf Hg" (see also Czaja, 2003), with ca. 50 achenes, and the new comparisons to Negru's images by the authors confirm that it may belong to the same species, even if the achene lengths are slightly different: 1.5-2.0 in Negru's material versus 1.3-1.8 in the Berzdorf Hg sample. Furthermore, the distinction of the German specimens assigned to C. limosioides and C. plicata is not clear, and in some cases could be simply explained by different preservation (e.g., in Czaja, 2003; Plate 17, Figs. 1, 2 and 3: major abrasion and removal of the outer periclinal walls; in Plate 17, Figs. 10-12: poor abrasion and partial to total preservation of the outer periclinal walls).
Carex pendula Huds. [BS] -type
Notes: The record by Mai and Walther (1988) has been reclassified as C. plicata.
Carex plicata Lanc.-Srod. [FS], in Acta Palaeobotanica 20: 90 (1979). [Fig. 5f]
Type material: Nowy Sacz I (57, 180, 193), Nowy Sacz II (18, 23, 30, 31, 39). Illustrated in Lancucka-Srodoniowa (1979, photo); Plate XIV, Figs. 20-25.
Distribution: Germany, Italy, Poland.
Locality records: Lancucka-Srodoniowa (1979, photo), Mai and Walther (1988) [as C. pendula (Mai, 1999)], Martinetto (1994a, SEM; 1994b), Mai (1995, photo), Martinetto and Mai (1996), Mai (1999, photo; 2000, photo, SEM), Czaja (2003, SEM), Ciangherotti et al. (2007); doubtful at Zastawniak (1992, photo) [as C. cf. plicata], Cavallo and Martinetto (1996) [as C. cf. plicata], Martinetto et al. (2007) [inch C. aff. pendula], Mai (2008, photo).
Time interval: Early Miocene to late Pliocene; a doubtful record from the late Oligocene (Chattian).
Kind of remains: Achenes and utricles
Notes: There are few to no characters to separate the shorter achenes of C. plicata from C. pendula, and also from those of the fossil-species C. limosioides. The original publication (Lancucka-Srodoniowa, 1979) describing C. plicata shows a very variable length range, which is not observed in the extant C. pendula. That these samples might represent a mixture of different species should not be discounted, and reevaluation of the original collection at Krakow is needed to verify if these materials really represent a single species. The late Oligocene C. plicata record cited by Mai (2008) is needed of revision.
Section Spirostachyae Drejer ex Bailey
Carex binervis Sm. [BS] -type
Distribution: Germany.
Locality records: Mai and Walther (1988, photo), von Bulow and Mai (1992, photo), Mai (2004, photo).
Time interval: Pliocene.
Kind of remains: Achenes.
Notes: Some of the pictured achenes preserve the strongly constricted substipitate bases typical of section Spirostachyae (E. M. and P. J.-M., pers. obs.). Nevertheless, sectional placement is in need of reevaluation.
Carex laevigata Sm. [BS] -type
Distribution: Germany, Italy, United Kingdom.
Locality records: Reid and Reid (1907b, photo) [as C. cf. helodes], Martinetto and Mai (1996), Mai and Walther (1988, photo).
Time interval: Pliocene to early Pleistocene.
Kind of remains: Achenes, utricle.
Notes: Taxonomical affinities need reevaluation.
Section Squarrosae J. Carey
This section is currently endemic to North America. The species C. szaferi documentes its presence in the Old World through the Miocene to the early Pleistocene.
Carex szaferi P. I. Dorof., Simbuginskaya flora: 54. In: Goretsky G. I. (ed.) Fauna i flora Simbugino. Nauka, Moskva. (1977). [Fig. 51]
Type material: Holotype: BIN N587-13. Illustrated in Dorofeev (1977, photo); Plate IV, Fig. 28.
Distribution: Belarus, Czech Republic, Germany, Italy, the Netherlands, Poland, Russian Federation (European Russia, Western Siberia); doubtful from the Czech Republic.
Locality records: Reid and Reid (1915, photo) [as C. flagellata p.p. (Dorofeev, 1977; Mai & Walther, 1988; see Plate 3, Figs. 23-26)], Szafer (1938, photo) [as C. flagellata], Szafer (1947, photo) [as C. rostrata (Dorofeev, 1977; Velichkevich & Zastawniak, 2003)], Raniecka-Bobrowska (1959, photo) [as C. aff. rostrata (Dorofeev, 1977; Velichkevich & Zastawniak, 2003)], Dorofeev (1977, photo), Yakubovskaya (1982, photo) [as C. cf. szaferi], Martinetto and Mai (1996), Mai and Walther (1988, photo), Knobloch (1989, photo), Basilici et al. (1997), van der Burgh and Zetter (1998, photo), Cavallo and Martinetto (2001, photo), Velichkevich and Zastawniak (2003, photo), Martinetto et al. (2007); doubtful in Buzek et al. (1985, photo) and Kvacek and Teodoridis (2007) [as C. cf. szaferi]; see also Appendix S1.
Time interval: Late Miocene to Pliocene, and possibly also early Pleistocene.
Kind of remains: Achenes and utricles.
Notes: Similarities were reported with section Vesicariae (Mai & Walther, 1988), and later with Squarrosae J. Carey (Velichkevich & Zastawniak, 2003). Our own observations reveal that the achenes of this taxon, and especially the way these are very loosely involved by the strongly inflated utricles, are an almost perfect match for the achenes and utricles of C. squarrosa L. van der Burgh and Zetter (1998) considered C. szaferi to be the smaller fraction of the material included within C. flagellata, probably because achenes of both species frequently occur together (Velichkevich & Zastawniak, 2003). However, both taxa have distinct characters, with the achenes of C. szaferi being narrower, with sharper angles in cross-section, and with the base not distinctly widened (Fig. 51), whereas the fruits of C. flagellata are wider, with more rounded angles in cross-section, and with the base distinctly widened, platform-like (Fig. 5b). It would be highly desirable a critical revision of the materials ascribed to C. szaferi and to species belonging to section Vesicariae (specially C. flagellata) given the number of misidentifications and disagreements between authors.
Section Vesicariae Fries ex Rouy (Includes Sections Lupulinae Tuckerm. ex J. Carey and Pseudocypereae Tuckerm. ex Kiik.)
There is a very rich record for this section, as the aquatic habitats where most species of this section grow are more readily fossilized than other habitats. Given the wide observed variability, as well as the frequent disagreements between authors, a critical revision of the fossil materials ascribed to section Vesicariae would be highly desirable (see also comments under C. szaferi).
Carex sp.
Distribution: Germany
Locality records: Gregor (1982, Plate 14, Fig. 23 and possibly Fig. 25, photo) [as C. flagellata].
Time interval: Late middle Miocene (Serravallian).
Kind of remains: Achenes.
Notes: The two achenes of the Miocene locality Achldorf figured by Gregor (1982) as C. flagellata do not show the characters of this species, and the visible details seem to suggest two different species (see comments under C. klettvicensis). In the trigonous specimen of Plate 14, Fig. 23 in Gregor (1982), the outline points to an affinity to the achene figured by Lancucka-Srodoniowa (1966), without style, which was suggested to be similar to C. rostrata. The achene figured by Gregor (1982) is trigonous and has a fairly long and thick style, which also suggests affinity to section Vesicariae. Considering shape and dimensions of the achene, these fossils may also have affinities to C. graciosa. Another achene figured by Gregor (1982, Plate 14, Fig. 25) from the locality Thalham has a different shape, but it seems to be also trigonous and display a fairly long and thick style, which points to affinity to section Vesicariae.
Carex capricornis Meinsh. ex Maxim. [BS] -type
Images: Yamakawa et al. (in press, photo).
Distribution: Japan.
Locality records: Yamakawa et al. (in press).
Time interval: Late Pliocene (Piacenzian).
Kind of remains: Achenes and utricle remains.
Notes: These samples are different from the extant C. capricornis Meinsh. ex Maximowicz because of their longer fruits (2.5-3.0 mm vs. 1.5-2.0 mm) (cf. Yamakawa et al., in press).
Carex flagellata C. Reid & E. Reid [FS], The Pliocene flora of the Dutch-Prussian border: 69 (1915) [Fig. 5a and b]
Type material: Swalmen, Reuver, Brunssum. Illustrated in Reid and Reid (1915); Plate III, Figs. 22-26 (photo).
Synonym: Carex maii Erw. Knobloch, in Sbor. geol. ved 19: 173 (1983). Holotypus: Kranichfeld (MfN), illustrated in Mai (1965: Plate 4, Fig. 30, photo).
Distribution: Belarus, Bulgaria, Czech Republic, Germany, Italy, the Netherlands, Poland, Russian Federation (European Russia, Western Siberia, Yakutia, Kamchatka), Ukraine; doubtful records from Austria.
Locality records: Reid and Reid (1915, photo) [p.p. Mai & Walther, 1988: Plate 3, Fig. 22], Szafer (1954, photo), Raniecka-Bobrowska (1959, photo), Dorofeev (1962a, photo; 1963, photo), Mai et al. (1963, photo), Dorofeev (1966a, b, 1969), photo; 1971; 1977, photo), van der Burgh (1978, photo), Knobloch (1981a, photo), van der Burgh (1987), Negru (1986, photo), Palamarev and Petkova (1987, photo), Mai and Walther (1988, photo), Martinetto (1994b, photo), Palamarev (1994, photo), Martinetto and Mai (1996), Basilici et al. (1997), Dyjor et al. (1998), van der Burgh and Zetter (1998, photo), Westerhoff et al. (1998, photo), Mai (2000, photo, SEM), Mai and Wahnert (2000, photo), Ciangherotti et al. (2007); doubtful records from Szafer (1961, photo) [as C. cf. flagellata], Knobloch (1981b, photo) [as C. flagellata], Yakubovskaya (1982, photo) [as C. cf. flagellata], Kovar-Eder et al. (2006) [as C. cf. flagellata], Martinetto et al. (2007), and Martinetto et al. (2014b) [as C. cf. flagellata]', see also Appendix SI.
Time interval: Late Oligocene to early Pleistocene.
Kind of remains: Achenes and utricles.
Notes: Variously considered to be related to section Vesicariae itself (Reid & Reid, 1915), or section Lupulinae J. Carey (Mackenzie) (van der Burgh & Zetter, 1998; Mai & Wahnert, 2000). Current phylogenetic estimates demonstrate these two sections are intertwined and are now considered consectional (e.g., Waterway et al., 2009). There is not extant taxon in Europe similar in morphology to C. flagellata. While some authors saw relationships with C. szaferi, with which C. flagellata uses to occur in admixture (Velichkevich & Zastawniak, 2003), others even considered both to be synonyms. However, they seem to be distinct species (see C. szaferi paragraph). A critical revision of the materials belonging to both taxa is necessary to identify possible misidentifications. A record by Dorofeev (1963) is now considered as C. rostratapliocenica. Gregor's (1982) C. flagellata-type records are here considered as a generic Carex sect. Vesicariae record, and a possible C. klettvicensis (see above). Materials pictured in Szafer (1954) were considered to be C. szaferi by Velichkevich and Zastawniak (2003), however we prefer to be conservative and cite them under the name in the original publication.
Knobloch (1989) proposed C. maii as an additional species similar to C. flagellata that occurred in the European fossil record. He reported the very broad basal attachment as the main difference between both species. Carex flagellata fossils assemblages from Italy show a very broad variation in this particular character, so it should not be considered relevant to separate C. maii from C. flagellata (E. M., pers. obs.). Thus, we regard both names as synonyms.
Carex omoloica P. I. Dorof. ex Jim.-Mejias, S. Popova & Martinetto [FS] (Appendix 1)
Carex omoloica P. I. Dorof., Tretichnye flory basseyna reki Omoloya (Tertiary floras of Omoloy river floras) In I. T. Vassilchenko (ed.), Istoriya flory i rastitelnosty: 81 (1972), nom. inval.
Type material: Holotype: Khapchan-Khaya, outcroop 352. Illustrated in Dorofeev (1972, photo); table IX, Fig. 2.
Distribution: Russian Federation (Yakutia).
Locality records: Dorofeev (1972, photo); see also Appendix SI.
Time interval: Late Miocene; doubtful from the early Pliocene
Kind of remains: Achene and utricle remains.
Notes: Achenes resemble C. flagellata but smaller and less elongated (Dorofeev, 1972). The name C. omoloica P. I. Dorof. was originally not validly published because no holotype was designated (ICN, McNeill et al., 2012). We designate a holotype among Dorofeev's (1972) protologue illustrations to let this name be validly published (Appendix 1).
Carex pseudocyperoides Lanc.-Srod. [FS] in Acta Palaeobotanica 20: 91 (1979).
Type material: Nowy Sacz I (57, 271, 291, 300, 311, 346, 348), Nowy Sacz II (18, 23, 29(7), 32). Illustrated in Lancucka-Srodoniowa (1979, photo); Plate XIV, Figs. 14-17.
Distribution: Germany, Poland, Russian Federation (Western Siberia), Ukraine.
Locality records [inch C. pseudocypems-type records from Miocene; the distinctive characters of the two species do not seem clear to us]: Lancucka-Srodoniowa (1979, photo), Negru (1986, photo), Lesiak (1994, photo), Mai (1999, photo; 2000, photo, SEM), Mai and Wahnert (2000, photo) [as C. pseudocypems], Czaja (2003). Doubtful record from Szafer (1961) [as cf. C. pseudocypems]', see also Appendix S1.
Time interval: Early to late Miocene.
Kind of remains: Achenes and utricles.
Notes: Mai labeled some of the MfN collections as "Carex zittawensis", but this name seems to have never been published, since these fossils are cited under C. pseudocyperoides in Mai (2000). In the MfN collection C. pseudocyperoides is documented by records from 10 localities (1-3 achenes each). There are only two specimen-rich samples: "Berzdorf Hg" and "Hartau 1". Incomplete utricles, showing ca. 10 nerves per side, are only present at Hartau 1, and the association to the achenes is ambiguous. Achenes with the best preserved styles are in the Berzdorf Hg material, and show a distinct style attenuation at 1/3 achene's length. This last character, as well as the utricle characters reported by Mai and Wahnert (2000), suggest a clear relationship of this taxon to C. pseudocypems, as Lancucka-Srodoniowa (1979) pointed out in her original description. There seems to be a discontinuous documentation of C. pseudocypems-type remains from the Lower Miocene to the Holocene. It remains necessary to test if Miocene forms can be really separated or if these conform a chronospecies.
Carex pseudocypems L. [BS] -type Fig. 51).
Distribution: Czech Republic, Germany, Italy, the Netherlands, Poland.
Locality records: Szafer (1947, 1954, photo), Mai et al. (1963, photo), Mai and Walther (1988, photo), Knobloch (1989, photo), von Billow and Mai (1992), Martinetto (1994a, SEM; 1994b), Bertoldi and Martinetto (1995), Martinetto and Mai (1996), Westerhoff et al. (1998), Mai and Wahnert (2000, photo), Girotti et al. (2003), Mai (2004), Ciangherotti et al. (2007), Martinetto et al. (2007), Martinetto et al. (2014b, SEM); a doubtful record in van der Burgh (1983, photo) [as C. panicea p.p.: Plate 4, Fig. 5, achene at the right].
Time interval: Early Pliocene to early Pleistocene.
Kind of remains: Achenes and utricles.
Notes: Miocene remains cited as C. pseudocypems have been transferred to C. pseudocyperoides. See above comments about the affinities with this putative species. Several Pliocene records of C. pseudocypems are represented by abundant specimens including perfectly preserved achenes and complete utricles with beaks (Fig. 51). The good degree of conservation of these remains even allow to rule out other closely related species as a match, such as C. capricornis or C. comosa Boott.
Carex rostrata-pliocenica V.P. Nikitin [FS], in Miocene of the Mamontova Gora: 135 (1976).
Basionym: Carex rostrata? f. pliocenica, Nikitin Pliocene and quat. Floras of Voronezh: 120 (1957) Plate III, Fig. 25.
Type material: Brown coal of locality Kriviborie. Illustrated in Nikitin (1957, photo); Plate III, Figs. 25-26.
Locality records: Nikitin (1957, photo) [as C. rostrata f. pliocenica], Dorofeev (1963 [as C. flagellata p.p.]; 1969, photo; 1977, photo), Nikitin (1976, photo), Dorofeev (1979, 1986); see also Appendix SI.
Time interval: Miocene to early Pleistocene (Velichkevich et al., 1998).
Kind of remains: Achenes.
Distribution: Poland, Russian Federation (European Russia, Yakutia).
Notes: Nikitin (1957) originally reported achenes of this species to be similar to C. rostrata and C. flagellata, but larger than the former and smaller than the latter (2.0-2.5 x 0.9-1.1 mm) and with a surface sculpture displaying a more marked relieve, describing also a utricle to be 3.3 x 1.7 mm. He considered C. rostrata-pliocenica to resemble some atypically narrow achenes that occur among modern C. rostrata samples (particularly immature materials). We agree with Nikitin that this fossil-species is certainly similar to the modern C. rostrata and also to C. vesicaria, but the differences with the extant taxa are not clear to us. Identical remains in western Europe (e.g., Mai and Walther (1988), Martinetto (1994b)) have been named C. rostrata or C. cf. rostrata (see below). A comparative analysis of the fossil material from Eastern and Western Europe is needed, as well as a detailed comparison to modern materials. For the sake of simplicity we list under this epigraph materials that have already been transferred to C. rostrata-pliocenica by previous authors, and we keep under the C. rostrata/C. vericaria-type epigraph those that have not. The record by Velichkevich and Zastawniak (2003) is here considered as C. yakubovskayae. The record by Martinetto et al. (2012) is here transferred to C. rostrata-type epigraph.
Carex rostrata Stokes [BS] -type and Car ex vesicaria Huds. [BS] -type
Distribution: Europe (Czech Republic, Germany, Italy, the Netherlands, Poland, European Russia, United Kingdom), Asia (Japan, Russian Federation (Western Siberia)), and North America (Canada, Yukon; USA, Alaska).
Locality records: Reid and Reid (1907b, photo), Szafer (1954, photo), Dorofeev (1962b, photo; 1963, photo; 1965), Lancucka-Srodoniowa (1966, photo), van der Burgh (1978, photo; 1983), Buzek et al. (1985), Mai and Walther (1988, photo), Matthews and Ovenden (1990), von Bulow and Mai (1992), Martinetto (1994b), Westerhoff et al. (1998), Momohara and Saito (2001), Mai (2004, photo and SEM), Kvacek and Teodoridis (2007), Martinetto et al. (2007), Martinetto et al. (2012) [as C. rostrata-pliocenica]', see also Appendix SI.
Time interval: Middle Miocene to early Pleistocene.
Kind of remains: Achenes and utricles.
Notes: Modern C. rostrata achenes are very variable and difficult to distinguish from C. vesicaria (E. M., pers. obs.), therefore any suggestion of affinity to either of the species should be avoided for fossil records. The records by Szafer (1947) and Raniecka-Bobrowska (1959) has been both transferred to C. szaferi (Velichkevich & Zastawniak, 2003 and Dorofeev, 1977 respectively). Also records have been transferred to C. rostrata-pliocenica (see epigraph above), but differences between both species does not seem clear to us.
Carex yakubovskayae Jim.-Mejlas, S. Popova & Martinetto [FS], nom. nov. [Fig. 5c and d]
Nom. subs.: Carex curvata Yakubovskaya, in Paleokarpologiceskije issledovanija Kajnozoja: 59 (1982) nom. illegy non Carex curvata Knaf, Flora 30: 184 (1847).
Etymology: Dedicated to T. V. Yakubovskaya, who originally described this species as C. curvata T. V. Yakubovskaya.
Type material: Holotype: coll. num. 306-51 (306-Ya), Dvorets, Gomel oblast', Russian Federation, borehole n. 4, drilling depth 28.7-29.0 m; late Pliocene (early Dvorets subsuite). Illustrated in Yakubovskaya (1982, photo); Tab. II, Figs. 17-22 [as C. curvata].
Distribution: Belarus, the Netherlands, Russian Federation (European Russia).
Locality records: Reid and Reid (1907a, photo) [as C. sp.], Nikitin (1957, photo) [as C. sp. 15] Dorofeev (1977, photo) [as Carex sp. 1], Yakubovskaya (1982, photo), Velichkevich (1990), Velichkevich and Zastawniak (2003, photo) [as C. rostratapliocenica].
Time interval: Early Pliocene to early Pleistocene.
Kind of remains: Achenes and utricles.
Notes: Style robustness and achene shape point to affinities of this fossil with section Vesicariae. It has been suggested to be a synonym of C. rostrata-pliocenica, as paleobotanists considered the typical lateral invagination on the achene wall not to be "an important feature for species recognition" (Mai, 1999; Velichkevich & Zastawniak, 2003). Indeed, this invagination is a peculiar feature of strong taxonomic value in Carex (P.J.-M., pers. obs.), that in section Vesicariae is only known in the extant North American C. tuckermannii Dewey (Ball & Reznicek, 2002) and the South American C. luridiformis Reznicek & S. Gonzalez and C. setigluma Reznicek & S. Gonzalez (Reznicek & Gonzalez-Elizondo, 1995). This example points out the need of stronger connections between palaeobotanists and plant taxonomists.
Records that may Belong Either to the Caricoid Clade or to Section Ovules
Here we cite biconvex or obtusely trigonous oblong achenes with well-preserved elongated style bases. This combination of characters is a good match for either section Ovales (subgenus Vignea) or to certain members of the Caricoid clade (e.g., section Leucoglochin Dumort. s.L, former genus Kobresia). While the comparison of modern materials readily allows the distinction of these two very different groups of sedges on the basis of other additional characters (e.g., epidermal cell-pattern), in the studied fossil remains these characters are not so straightforward to interpret. Members of the Caricoid clade display a great morphological variation (Global Carex Group, 2015) that is also found in achenes morphology (P. J.-M., pers. obs.). Achenes from species of the Caricoid clade range from trigonous and broadly ovate or obovate in some groups, to flattened biconvex narrowly oblong in other groups. A special effort is needed to better understand the carpological variation within this group, which will permit a more confident approach to interpreting the fossil remains listed under this epigraph.
Carex sp.
Illustration: Velichkevich and Zastawniak (2003, photo).
Distribution: Belarus.
Locality records: Velichkevich and Zastawniak (2003) [as C. sp. 1],
Time interval: Pliocene.
Kind of remains: Achenes.
Notes: The materials reported by Velichkevich and Zastawniak (2003) were considered similar to the Asian C. remotiuscula Wahlenb. (subgenus Vignea, section Remotae). The two figured achenes seem to be almost identical to those of the modern C. bohemica Schreb. The appearance of the style remain, with a truncation at 1/5 of the achene's length, associated to the persistence of central filament, suggests a jointed style with a distal deciduous portion, a pattern that is typical for subgenus Vignea.
Carex paucijlora Lightf. [BS] -type
Distribution: Belarus, Russian Federation (European Russia, Western Siberia, Altay).
Locality records: Dorofeev (1959c; 1960a, c; 1962b, photo; 1963 [p.p.]; 1965; 1966a, photo; 1967a, c; 1968). See also Appendix SI.
Time interval: Miocene toearly Pleistocene.
Kind of remains: Achenes.
Notes: These fossil materials cited as C. pauciflora should probably belong to C. paucifloriformis or C. paucijloroides. Revision is needed.
Carexpaucifloriformis ("paucifloraeformis") V.P. Nikitin [FS], in Baranova et al, The Miocene of the Mamontova Gora 173 (1976).
Type material: Holotype: Miocene deposits "Besheul horizon" Kartashevo, Irtysh river. Illustrated in Nikitin (1976, photo); Plate 45, Fig. 31.
Distribution: Kazakhstan, Russian Federation (Western Siberia, Yakutia).
Locality records: Dorofeev (1963, photo) [as C. cf. pauciflora], Dorofeev (1969, photo) [as C. sp. 3], Nikitin (1976, photo). Doubtful at Dorofeev and Tjulina (1962). See also Appendix S1.
Time interval: Late Oligocene to Pliocene, doutbful for the early Oligocene.
Kind of remains: Achenes and utricle remains.
Notes: These fossil achenes definitely resemble the achenes of C. pauciflora Light., but they are smaller (13-2.0 x 0.6-0.9 in C. paucifloriformis vs. 2.123 x 0.8-1.0 mm in C. pauciflora (Ball & Recnizek, 2002)), and the general appearance is clearly different. They also differ from C. microglochin (Berggren, 1969) in the shorter style. Nikitin (1976) reported the epidermic cell structure as "ambiguous". From the extinct C. klarae Mai (section Ovales, subgenus Vignea) this species differs in the absence of nerves in the utricle remains and the different surface sculpture of the achene. According to Velichkevich and Zastawniak (2003) the achenes of C. paucifloriformis are also similar to C. paucifloroides, but are smaller and possess utricle remains with a smaller number of nerves.
Carex paucifloroides Velichkevich [FS], in N. A. Makhnach (ed.) Stratigrafiya I Paleontografiya Antropogena: 125 (1975). [Fig. 41]
Type material: II-73-2-1, Dvorets, richikhinsky rayon, Gomel oblast', Russian Federation, late Pliocene. Illustrated in Velichkevich (1975); Plate 1, Fig. 24.
Distribution: Italy, Germany, Belarus.
Locality records: Velichkevich (1975, photo), Yakubovskaya (1982), Mai and Walther (1988, photo), von Billow and Mai (1992, photo), Martinetto (1994a) [as C. cf. paucifloroides], Bertoldi and Martinetto (1995), Velichkevich and Zastawniak (2003, photo), Mai (2004, photo), Velichkevich and Zastawniak (2007, photo), Martinetto et al. (2007). In this paper we confirm a report of C. paucifloroides for the locality Momello-Lanzo in NW Italy (Fig. 41), preliminarily reported by Martinetto (1995).
Time interval: Early to late Pliocene. The species seems to pass into the Pleistocene (Velichkevich & E. Zastawniak, 2006).
Kind of remains: Achenes and utricle remains.
Notes: The specimens assigned to C. paucifloroides in the collection MfN show a long style-base, a very fine cell pattern, and a cross-section either biconvex or trigonous, which indicates the Caricoid clade as a good possible match. Affinities with section Ovales (subgenus Vignea) may not be totally ruled out due to the characteristics of the style. All of the studied samples of C. paucifloroides showed a distinct restriction of the style base at 1/3 achene's length, a character that is not common in the modern members of the Caricoid clade. On the other hand, the observed cell pattern is finer than in the studied species of section Ovales, as well as in some members of the Caricoid clade (e.g. C. pulicaris L.).
Conclusions
Paleobiogeography and Paleoecology of Carex
Little can be said about the biogeography of the fossil record of Carex due to the geographical bias in the fossil sampling and taxonomic uncertainty of many taxa, but at least three different patterns can be inferred for certain fossil remains.
1- Temporal Persistence. For certain types of samples, such as fossils belonging to section Rhynchocystis and C. pseudocyperus-Uke remains (including C. pseudocyperoides and other C. pseudocyperus-type fossils), their presence is recorded continuously in Europe since the Miocene. For the sections Lageniformes and Rhomboidales, both mostly endemic to East Asia (Dai et al., 2010), their presence in this area can be traced back to the late Miocene (Japan). Despite the fact that the persistence of Tertiary relict taxa in East Asia and North America is a fairly well-known phenomenon (Milne & Abbott, 2002; Huang et al., 2015; Tang, 2015; and references therein), reports of the species of such nature are more rare in Europe and mainly refer to woody plants (e.g., Palamarev, 1989; Willis & Niklas, 2004). Our data suggests that at least species from section Rhynchocystis and Carex pseudocyperus-likc plants may be herbaceous Tertiary relicts in the Old Continent.
2- Extinction. Two fossil species from Europe are reported to belong to species groups currently absent in this territory. These are C. yakuboskayae (allied to American species of section Vesicariae, as C. tuckermannii), and C. szaferi (belonging to the North American endemic section Squarrosae). To this should be added C. flagellata, of uncertain affinities within section Vesicariae but definitely not related to any European extant taxon. Extinction of elements of American or Asian affinity in Europe has been long reported for other plant groups (Szafer, 1954; Svenning, 2003; Postigo-Mijarra et al., 2009; Martinetto et al., in press). It is not a surprise to find this pattern in Carex also. Similarly C. panormitana-type plants are today known to occur only in the Mediterranean (Jimenez-Mejias et al, 2014), whereas fossil remains have been found in the Pliocene of Central Europe and Northern Italy, where they are currently absent. It suggests the occurrence of these taxa in the past much more northwards than they are currently found, probably promoted by the formerly warmer climate conditions in these areas (see Utescher et al., 2011 and references therein).
3- Absence. There are a large number of extant Carex species known from Siberia (see Egorova, 1999). However, the fossil record of the genus in this area is, despite of its abundance, restricted to a few species during the Miocene and Pliocene (see Appendix SI). There is a shift in the composition of the aquatic flora of Siberia between the warmer and drier Tertiary and the cooler and wetter Neogene (Dumikin, 2010), probably due to the immigration into Siberia of exogenous elements. Carex seems to follow this pattern, becoming a more diverse genus in Siberia in more recent times.
The knowledge about the paleoecology of Carex is partially limited for the same reasons as for the biogeography: sampling bias and taxonomic uncertainty. To this it should be added the lack of an exhaustive paleocological characterization for most fossil sites. Nevertheless, the association of most remains to freshwater environments seems to indicate an ecology linked to these wet habitats. It is particularly remarkable for the remains ascribed to sections Carex, Phacocystis and Vesicariae, suggesting some degree of niche conservationism since at least the Miocene. Conversely, extant groups that are largely confined to dry environments seem to be totally absent from the fossil records (e.g., sections Aulocystis Dumort., Halleranae (Asch. & Graebn.) Rouy, or Mitratae).
Summary
Our research shows that there is an abundant fossil record potentially assignable to species of genus Carex of which most neobotanist were not aware of. This includes up to 83 different names applied to remains that we considered to be reliably assigned to Carex, plus a number of samples not identified to species level. All these come from near 550 different sites. Additionally, another 23 Carex names have been applied to remains of doubtful identity. In any case, further research is needed to confidently assess the affinities of many fossils to particular groups of species, sections or subgenera.
The temporal range of the fossils confidently belonging to Carex is from the late Eocene of England (C. colwellensis and C. gurnardi, an achene and a utricle respectively) to recent times. Remains of this genus seem to become abundant from the middle Miocene onwards. Most fossils are ascribed to extinct species. Ten fossil-species seems to be recorded also for the Pleistocene (C. carpophora, C. dorofeevii, C. elongatoides, C. flagellata, C. flavicarpa, C. frequens, C. yakubovskayae, C. major, and C. rostrata-pliocenica), although probably some of these records need revision (e.g., C. dorofeevii, C. major). The use of extant species names applied to fossil samples is common for Pliocene material, with very few examples dating back to the Miocene, and most of these probably referring to a morphological similarity rather than actual taxonomic identification.
We are aware that most of the fossils reviewed come from a relatively small number of places, with a strong bias to Central Europe. Carex fossils have been found so far from Europe, Asia, and North America. The fossil record of Carex and many other plant genera and families would be greatly increased if more attention were paid to these mesofossils (e.g., Smith et al, 2008). The systematic community should be more aware of the amount of fossil data awaiting evaluation in this and other plant groups. More complete basic research on fossil morphology and taxonomic affinity, as well as morphology and anatomy in modern plant groups for context, is required for phylogeneticists to be able to integrate the "known fossil record" for many plant groups into other studies.
We would like to encourage the neobotanical and paleobotanical communities to work together on broad-scope systematic projects like the one we present here. We hope that this study and others such as this will help to uncover the many neglected fossils that, among other results, provide a more complete understanding of evolutionary diversification of lineages across temporal and spatial scales.
DOI 10.1007/s12229-016-9169-7
Acknowledgments We would like to acknowledge C. Hiller, K. Puffert, and especially S. Schultka from the Museum fur Naturkunde of Berlin for assistance in consulting Mai's collection; A. Hvalij for providing critical literature and access to the databased information from Nikitin's papers and collections; Z. Zhou and P. Grote because of discussion about occurrence of Carex fossils in China and South East Asia respectively; J. Kvacek and V. Turek (National Museum of Prague) for the pictures and information from the type specimen of C. antiqua A. Slavik; D. de Franceschi for updated information about Saporta's collection; T. V. Yakubovskaja for permission reproducing the pictures of C. yakubovskayae, and G. E. Rodriguez Palacios for technical support during bibliography digitalization.
This project was funded by NSF-DEB Award #1256033 to EF1R.
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Appendix 1
Nomenclatural adjustments implying newly formal descriptions provided in this paper.
Carex dorofeevii Egor, ex Jim.-Mejias, S. Popova & Martinetto, spec, nova
Carex antiqua P. I. Dorof., Treticnye floiy Urala: table IV, Figs. 18-22. (1970) nom. inval.
Carex dorofeevii Egor. Sedges Russia: 673 (1999) nom. inval.
Carex dorofeevii Mai in Palaeontographica Abt. B, 250 (1-3): 36 (1999) nom. inval.
Holotype: K464 (Komarov Botanical Institute), Polevskoy, Miocene browncoal deposits (Russian Federation): specimen illustrated in Dorofeev (1970, photo: Plate IV, Fig. 21 [as C. antiqua]).
Type stratum: Polevskoy, brown coal deposits, Miocene.
Description: Achenes 2.5-3.0 x 1.2-1.7 mm, narrowly obovate to elliptical in outline, trigonous, slightly curved. Faces concave in cross section, almost equal in width; edges thick at the base. Upper half of the achene acute, gradually narrowing into the style base. Style base up to 1/7 of the achene length, rounded-triangular, not very thick, often bent. Lower half of the achene gradually tapering into a narrow base. Base of the achene bearing a more or less definite, thin callus. Wall thick, leathery, and elastic; surface brown, shiny, with large cells with shallow cell lumina pits.
Etymology: Dedicated to P. I. Dorofeev, who originally described this species as C. antiqua P. I. Dorof. We follow a previous invalid intent by Egorova (1999) (see comments on the checklist main text).
Carex maiana Martinetto, spec, nova (Figs. 2c and 4i-j)
Holotype: MfN Berlin (Slg. Mai, Nr. 1993/3865b). Illustrated in Mai (2000) [Plate 14, Fig. 12],
Type stratum: Clay layer with conifer cones ca. 10 m above the "Spezialton" layer, Klettwitz 3 in Mai (2000), stratigraphic unit "Meuroer Folge" of lower Lusatia.
Paratypes (cited in the stratigraphic order suggested by Mai (2000,2004)): Liibtheen 2/75 (MFN coll. Mai Nr 10132), Klettwitz 1 (MFN coll. Mai Nr. 865), Klettwitz 2 (MFN coll. Mai Nr. 1183b), Klettwitz 3 (MFN coll. Mai Nr. 3865b, 3885b; Figs. 2c and 4i-j), Kleinleipisch 2 (MFN coll. Mai Nr. 3968), Nochten 1 (MFN coll. Mai Nr. 8129), Broethem (MFN coll. Mai Nr. 4566b). Doubtful record (achene covered by utricle remains): Kleinleipisch-Roemerkeller (MFN coll. Mai Nr. 6190a) [cited as C. klettvicensis from Kleinleipisch, Mai (2000); Fig. 5i of this paper].
Description: Achenes 1.2-1.6 x 1.0, elliptic to narrowly obovate, narrowly biconvex. Faces slightly concave; edges thick at the apex. Upper half of the achene rounded, abruptly contracted into the style base. Style base up to 2/5 of the achene length, flattened, regularly truncated in a straight breaking point. Lower half of the achene tapering into a wide base, about 1/3 achene width. Base of the achene bearing a thin callus. Wall surface with isodiametric, polygonal cells, 20-35 pm wide (mean 28 pm on the central part of the achene face), giving to the achene an apparent meshed pattern. Utricle remains thin, probably with 2-4 thin nerves on each face.
Etymology: Dedicated to D. FI. Mai, who originally identified this species as a new species and named it "C. praeacuta " in the labels of his collection at the MfN, although it was never formally described.
Carex omoloica P. I. Dorof. ex Jim.-Mejias, S. Popova & Martinetto, spec, nova
Carex omoloica P. I. Dorof., Tretichnye flory basseyna reki Omoloya (Tertiary floras of Omoloy river floras) In I. T. Vassilchenko (ed.), Istoriya flory i rastitelnosty: 81 (1972), nom. inval.
Holotype: K274 (Komarov Botanical Institute): specimen illustrated in Dorofeev (1972, photo: table IX, Fig. 2).
Type stratum: Khapchan-Khaya outcrop 352.
Description: Achenes 1.2--1.5 x 1.0-1.2 mm, subrhomboid to obovate, trigonous. Faces slightly concave, almost equal in width; edges thick, rounded, sharper at the base. Style as long as or longer than the achene length, sharply trigonous at the base, turning rounded-triangular at the middle, gradually tappering into a curved end. Walls thick, elastic. Utricles ovate, thin, flatfish, with seven veins on each face.
Etymology: The name omoloica, chosen by Dorofeev (1972), refers to the Omoloy river in Yakutia (Russian Federation) where the remains of this species were first found.
P. Jimenez-Mejias (1,6) * E. Martinetto (2) * A. Momohara (1) * S. Popova (4) * S. Y. Smith (5) * E. H. Roalson (1)
(1) School of Biological Sciences, Washington State University, Pullman, WA 99164-4236, USA
(2) Dipartimento di Scienze della Terra, Universita dcgli Studi di Torino, Via Valperga Caluso 35, 1-10125 Torino, Italy
(3) Graduate School of Horticulture, Chiba University, 648 Masudo, Chiba 271-8510, Japan
(4) Komarov Botanical Institute of Russian Academy of Sciences, St. Petersburg 197376, Russia
(5) Department of Earth & Environmental Sciences and Museum of Paleontology, University of Michigan, Ann Arbor, MI 48109, USA
(6) Author for Correspondence; e-mail: pjimmej@gmail.com
Published online: 17 June 2016
The two first authors contributed equally to the preparation of this paper.
Electronic supplementary material The online version of this article (doi: 10.1007/s 12229-016-9169-7) contains supplementary material, which is available to authorized users.
Table 1 Alphabetical list of Carex names at species rank published to refer fossil materials, corresponding tentative systematic placement provided in this paper, and additional pertinents comments on the reliability of the remains (including synonymy) Systematic Name in references placement Notes C. acuta L. See Carex klettvicensis C. acutiformis Subgenus Carex, Systematic Ehrh. section Paludosae affinities of the fossils needed of revision C. acutimontana Mai Subgenus Vignea, section incertae sedis C. acutior Saporta Doubtful as Carex C. amissa Hcer Doubtful as Carex C. anderssoni Heer Doubtful as Carex C. antiqua P. I. Carex dorofeevii Dorof. C. antiqua Heer cf. Limnocarpus (Najadaccae) C. antiqua A. Doubtful as Carex Slavik C. aplwnolepis Subgenus Carex, Systematic Franchct & Savatier section Molliculae affinities of the fossils not completely reliable C. apiculata Doubtful as Carex Saporta C. appropinquata See Carex Schum. klettvicensis C. aquatilis Subgenus Carex, Systematic Wahlcnb. section Phacocystis affinities of the fossils needed of revision "C. atrofusca Subgenus Carex, Schkuhr." section incertae sedis C. atherodes Subgenus Carex, Systematic Spreng. section Carex affinities of the fossils needed of revision C. berggreni Heer Doubtful as Carex C. binervis Sm. Subgenus Carex, Systematic section affinities of the Spirostachyae fossils needed of revision C. blysmoides Subgenus Carex, Dorof. section Phacocystis C. bohemica Scherb. Subgenus Vignea, Systematic section Ovales affinities of the fossils needed of revision C. breviscapa C. B. Subgenus Carex, Clarke section Lageniformes C. brizoides L. Subgenus Vignea, section Ammoglochin C. burrardiana Doubtful as Carex Dawson C. canescens L. Subgenus Vignea, Systematic section Glareosae affinities of the fossils needed of revision C. capillaris L. Subgenus Carex, Systematic section affinities of the Chlorostachyae fossils needed of revision C. capricomis Subgenus Carex, Meinsh. section Vesicariae C. carpophora Mai & Subgenus Carex, H. Walther section Carex C. canescentoidea Subgenus Vignea, Systematic Mai section Glareosae affinities of the fossils needed of revision C. cespitosa L. See Carex klettvicensis and C. panonnitana C. chordorrhiza Subgenus Vignea, Systematic L.f. section affinities of the Chordorrhizae fossils needed of revision C. clarkii E. W. Applied only to Berry leaf remains; doubtful as Carex C. colchica J. Gay Subgenus Vignea, Systematic section Ammoglochin affinities of the fossils needed of revision C. colwellensis M. Incertac sedis Chandler Carex C. comuta Saporta Cladium biconte Saporta C. curvata Carex yakubovskayae Yakubovskaya C. davalliana Sm. Subgenus Vignea, Systematic section affinities of the Physoglochin fossils needed of revision C. diandra Schrank. Subgenus Vignea, Systematic section affinities of the Heleoglochin fossils needed of revision C. diffusa Saporta Doubtful as Carex C. dioica L. Subgenus Vignea, Systematic section affinities of the Physoglochin fossils needed of revision C. dorofeevii Egor, Incertac sedis ex Jim.-Mejlas, S. Carex Popova & Martinetto C. echinata Murray Subgenus Vignea, Systematic section Stellulatae affinities of the fossils needed of revision C. effossa Hcer Doubtful as Carex C. eigensis Mai Subgenus Carex, section incertac sedis C. elata All. Subgenus Carex, section Phacocystis C. elongata L. Subgenus Vignea, Systematic section Elongatae affinities of the fossils needed of revision C. elongatoides Subgenus Vignea, Systematic Lanc.-Srod. section Elongatae affinities of the fossils needed of revision C. eximia Goppert & Acoropsis eximia Menge (Goppert & Menge) Bogner (Araceae) C. flagellata C. Subgenus Carex, Reid & E. Reid section Vesicariae C. flava L. See Carex flavicarpa and C. gothanii C. flavicarpa Jim.- Subgenus Carex, Systematic Mejlas & Roalson section affinities of the Ceratocystis fossils needed of revision C. flaviformis Carex flavicarpa Lanc.-Srod. C. frequens V.P. Incertae sedis Nikitin Carex C. globosiformis Subgenus Carex, Systematic Lanc.-Srod. section Acrocystis affinities of the fossils needed of revision C. gothanii Mai & Subgenus Carex, Walther section incertae sedis C. graced J. Incertae sedis Thomasson Carex C. graciosa Negru Subgenus Carex, section Carex C. gumardi E. Reid Incertae sedis & M. Chandler Carex C. hartauensis Mai Subgenus Carex, section incertae sedis C. helmensis Mai & Subgenus Vignea, H. Walthcr section incertae sedis C. hirta L. Subgenus Carex, Systematic section Carex affinities of the fossils needed of revision C. hostiana DC. Subgenus Carex, Systematic section affinities of the Ceratocystis fossils needed of revision C. hostianoides Mai Subgenus Carex, Systematic section affinities of the Ceratocystis fossils needed of revision C. hyperborea Heer Doubtful as Carex C. ischnostachya Subgenus Carex, Systematic Steud. section affinities of the Confertiflorae fossils needed of revision C. klarae Mai Subgenus Vignea, section Ch'ales C. laevigata Sm. Subgenus Carex, Systematic section affinities of the Spirostachyae fossils needed of revision C. lasiocarpa Ehrh. Subgenus Carex, Systematic section Carex affinities of the fossils needed of revision C. lepidocarpa Subgenus Carex, Tausch. section Ceratocystis C. lilpopi Kownas Doubtful as Carex C. limosioides Subgenus Carex, Negru section Rhynchocystis C. loliacea L. Subgenus Vignea, Systematic section Glareosae affinities of the fossils needed of revision C. lusatica Mai Subgenus Carex, section incertae sedis C. maackii Maxim. Subgenus Vignea, Systematic section Ovules affinities of the fossils needed of revision C. maiana Subgenus Carex, Martinetto section Phacocystis C. maii Erw. Carex flagellata Knobloch C. major V.P. Incertae sedis Nikitin Carex C. marchica Mai Subgenus Vignea, section incertae sedis C. mariae- Carex graciosa sronodiowae Negru C. misella Heer Doubtful as Carex C. mucronata Heer Doubtful as Carex C. muricata L. Subgenus Vignea, Systematic section affinities of the Phaestoglocliin fossils needed of revision C. nigra (L.) Subgenus Carex, Systematic Rcichard section Phacocystis affinities of the fossils needed of revision C. noursoakensis Doubtful as Carex, Heer also applied to leaf remains C. omoloica P. I. Subgenus Carex, Dorof. cx Jim.- section Vesicariae Mcjias, S. Popova & Martinetto C. palaeocarpa Doubtful as Carex Saporta C. pallescens L. Subgenus Carex, Systematic section Porocystis affinities of the fossils needed of revision C. panicea L. Subgenus Carex, Systematic section Paniceae affinities of the fossils needed of revision C. panormitana Subgenus Carex, Guss. section Phacocystis C. pauciflora Either Caricoid Systematic Lightf. clade or subgenus affinities of the Vignea, section fossils needed of Ovales revision C. paucifloriformis Either Caricoid V.P. Nikitin clade or subgenus Vignea, section Ovales C. paucifloroides Either Caricoid Vclichkevich clade or subgenus Vignea, section Ovales C. pendula Huds. See Carex plicata C. philibertii Doubtful as Carex Saporta C. pilulifera L. Subgenus Carex, Systematic section Acrocystis affinities of the fossils needed of revision C. plicataLanc.- Subgenus Carex, Srod. section Rhynchocystis "C. praeacuta Mai" Carex maiana C. praehirta Mai & Subgenus Carex, H. Walther section Paniceae C. protogaea Mai Subgenus Vignea, section inccrtae sedis C. pseudocyperoides Subgenus Carex, Lanc.-Srod. section Vesicariae C. pseudocyperus L. Subgenus Carex, section Vesicariae C. quinquenervis Name applied only Schmalh. to leaf remains C. recognita Hccr Doubtful as Carex "C. reidii Mai" See Carex davalliana C. rediviva Hecr Doubtful as Carex C. remota L. Subgenus Vignea, Systematic section Remolae affinities of the fossils needed of revision C. remotoides Mai Subgenus Vignea, Systematic section Remotae affinities of the fossils needed of revision C. riparia Stokes Subgenus Carex, section Paludosae C. rochettiana Hecr Fruits belonging to Nymphaeaceae C. rostrata Stokes Subgenus Carex, Sectional placement section Vesicariae fairly probable, but actual taxonomic affinities of the different fossils needed of revision C. rostrata- Subgenus Carex, pliocenica V.P. section Vesicariae Nikitin C. scheuchzeri Hcer Name applied only to leaf remains C. servata Heer Doubtful as Carex C. sodalis Saporta Doubtful as Carex C. spicata L. Subgenus Vignea, section Phaestoglochin C. strigosoides Subgenus Carex, Lanc.-Srod. section incertac sedis C. szaferi P. I. Subgenus Carex, Dorof. section Squarrosae "C. tertiaria Doubtful as Carex; (Unger) Heer" name correctly applied only to leaf remains C. tsagajanica Doubtful as Carex Krassilov C. ungeri Mai & H. Subgenus Vignea, Walther section Ammoglochin C. vaginata Tausch. Subgenus Carex, Systematic section Paniceae affinities of the fossils needed of revision C. vancouverensis Doubtful as Carex Dawson C. vesicaria L. Subgenus Carex, Sectional placement section Vesicariae fairly probable, but taxonomic affinities of the fossils needed of revision C. vulpina L. See Carex panomiitana C. yakubovskayae Subgenus Carex, Jim.-Mejias, S. section Vesicariae Popova & Martinctto C. zhilinii Bauleva Subgenus Carex, & V. P. Nikitin section incertae sedis For a more complete evaluation of the taxonomic accuracy of each name, the reader is referred to the main text of the checklist. Synonym names for fossil species are given. Synonym names of extant species arc omitted and the listed name is the current accepted one according Govaerts et al. (2016) Table 2 Localities where fossil carpological remains of Carex have been reported Reference/ GEOLOGICAL EPOCH Locality Age Taxa / names PALEOCENE Heer Firkanten Paleocene C. anderssoni (1869b) Formation, Cap (Paleobiology *, C. berggreni Staratschin, Database, 2015) *, C. Svalbard, hyperborea *, Norway C. misella * Heer Kongsfjorden Paleocenecf. C. ultima * (1869b) (Kingsbai), Dallmann, 1999) Svalbard, Norway Heer Ifsorisok Middle to C. (1874, Formation, earlylate noursoakensis 1883) Ivssorrigsok, Paleocene *, cf. Carex * Nugssuaq (Selandian- Peninsula, Thanctian, Greenland 61.1-58.7 Ma; Dam et al., 2009) Heer Atanekerdluk Middle C. (1880) Formation, Paleocene noursoakensis Atanekerdluk, (Sclandian; *? Greenland Paleobiology Database, 2015) Krassilov Vladivostok, Cretaceous-Pal C. tsagajanica (1976) Amur, Russian eocene boundary * Federation (Maastrichtian- Danian) Lesquereux South Table Early Paleocene C. berthoudi * (1873, Mountain, (Danian, 66.5- 1878) Golden, 65.9 [+ or -] Colorado, USA 0.2 Ma; Kauffman et al., 1990) EOCENE Chandler Upper Headon Late Eocene C. (1963) Hill (Priabonian; colwellettsis, Formation, see King, 2010) C. sp. Colwell Bay, Isle of Wight, United Kingdom Dawson Burrard Inlet, Eocene C. burrardiana (1895) Vancouver, *, C. British vancouverensis Columbia, * Canada Reid and Thomess Bay, Latest Eocene C. gurnardi, cf Chandler Bembridgc, (Priabonian, Carex * (1926) Isle of Wight, ca. 34 Ma; United Kingdom Collinson & Cleal, 2001; Hayes & Collinson, 2014) OL1GOCENE Alvarez Peguera, Middle cf Carex * Ramis and Mallorca, Oligocene Ramos Spain Guerrero (1986) Dorofeev Novyi Log, Oligocene C. sp. (1952) Sverdlovsk oblast, Urals, Russian Federation Dorofeev Kozyulino, Oligocene C. sp. (1959b, Tomsk oblast, 1963) Western Siberia, Russian Federation Dorofeev Dunayevskiy Oligocene C. spp. (1960b, Yar, Tomsk 1963) oblast, Western Siberia, Russian Federation Dorofeev Belyi Yar, Oligocene C. subg. (1963) Tomsk oblast, Vigneal, C. sp. Western Siberia, Russian Federation Dorofeev Novyi Log, Oligocene C. sp. (1970) Sverdlovsk oblast, Urals, Russian Federation Heer Greith coal Late Oligocene "C. tertiaria" (1855) mine, (Chattian; * Hohenrhone, Paleobiology Switzerland Database, 2015) Heer Rivaz-Monod, Late Oligocene C. mucronata * (1859) Molasse basin, (Chattian; Switzerland Emmanuel et al" 2012) Mai (2008) Oberleichters- Late Oligocene C. carpophora?, bach, (Chattian) C. Bayern, colwellensis, Germany C, plicata? Mai and Thietbarch, Late Oligocene C. hartauensis Walther Saxony, (Chattian) (1991) Germany Saporta Camoins-les- Late Oligocene C. palaeocarpa (1865) Bains, France (Chattian) * Saporta Aix-Basin, Oligocene- C. acutioi *, (1889) Provence, Miocene C. apiculata *, France boundary C. diffusa *, (Chattian- C. philibertii Aquitanian; see *, C. sodalis * Rcichenbacher, 2004) MIOCENE Buzck and Chomutov- Miocene (cf. C. hartauensis Holy Most-Teplice Teodoridis & (1964) Basin (Czech Kvacek, 2006) Republic) Czaja Berzdorf, Supposed early C. (2003) Saxony, and middle canescentoidea, Germany Miocene C. eigensis, C. (Langhian) elongatoidesl, C, graciosa, C. hartauensis, C. limosioides, C. lusatica, C. plicata, C. pseudocyperoi- des Dorofeev Bol'shoy Supposed late C. spp. (1951, Fontan, Miocene 1955a) Odessa, Ukraine Dorofeev Tabaki, Supposed late C. carpophora, (1955b) Odessa, Miocene C. spp. Ukraine Dorofeev Novonikol'- Supposed C. (1955c, skoyc, Miocene pauciflori- 1963) Omsk oblast, fonnis, Western C. subg. Siberia, Vignea, C. spp. Russian Federation Dorofeev Demidovo, Supposed late C. sp. (1955d) Odessa oblast, Miocene Ukraine Dorofeev Strashnyi, Supposed late C. sp. (1955d) Mykolayiv Miocene oblast, Ukraine Dorofeev Kochetovskaya, Supposed C. sp. (1959a) Rostov oblast, Miocene Russian Federation Dorofeev Bol'shaya Supposed late C. spp. (1959a) Orlovka, Miocene Rostov oblast, Russian Federation Dorofeev Lezhanka, Omsk Supposed middle C. pauciflora- (1959c, oblast, Miocene type, C. sp. 1963) Western Siberia, Russian Federation Dorofeev Mozyr', Gomel, Supposed C. sp. (1960a) Belarus Miocene Dorofeev Osovtsy, Supposed C. pauciflora- (1960a) Gomel, Belarus Miocene type Dorofeev Zhitkovichi, Supposed C. spp. (1960a) Gomel, Belarus Miocene Dorofeev Krasnaya Supposed late C. pauciflora- (1960a) Sloboda, Miocene type, C. subg. Minsk, Belarus Vignea Dorofeev Osovtsy, Supposed late C. pauciflora- (1960a) Gomel, Belarus Miocene type Dorofeev Kireyevsk, Supposed C. flagellata, (1960c, Tomsk oblast, Miocene C. pauciflora- 1963) Western type, C. subg. Siberia, Vignea, C. spp. Russian Federation Dorofeev Kireyevsk, Supposed early C. spp. (1960c) Tomsk oblast, Miocene Western Siberia, Russian Federation Dorofeev Gorniy, River Supposed C. flagellata (1962a) Ob', Kahnty- Miocene Mansi Okmg, Tyumen oblast, Western Siberia, Russian Federation Dorofeev Chernoluch'ye, Supposed C. pauciflora- (1963) Omsk oblast, Miocene type, C. spp. Western Siberia, Russian Federation Dorofeev Ebargul, Omsk Supposed C. pauciflora- (1963) oblast, Miocene type Western Siberia, Russian Federation Dorofeev Isaakovka, Supposed early C. carpophoral, (1963) Kireyevsk, Miocene C. flagellata, Omsk oblast, C. Western pauciflori- Siberia, formis, Russian C. rostrata- Federation pliocenica, C. subg. Vignea, C. sp. Dorofeev Kartashovo, Supposed C. pauciflora- (1963) Omsk oblast, Miocene type, C. Western pauciflori- Siberia, formis, Russian C. spp. Federation Dorofeev Korolenka, Supposed C. pauciflora- (1963) Omsk oblast, Miocene type Western Siberia, Russian Federation Dorofeev Krasnoyarka, Supposed early C. sp. (1963) Omsk oblast, Miocene Western Siberia, Russian Federation Dorofeev Yekaterinins- Supposed early C. sp. (1963) koye, Miocene Omsk oblast, (Aquitanian) Western Siberia, Russian Federation Dorofeev Apsheronsk, Supposed late C. sp. (1964) Krasnodar Miocene Krai, Russian Federation Dorofeev Yurovsk, Supposed C. flagellata, (1966b) Tyumen' Miocene C. sp. oblast, Urals, Russian Federation Dorofeev Sivkovo, Supposed C. sp. (1967a) Grodno, Miocene Belarus Dorofeev Detkovichi, Supposed C. sp. (1967b) Brest, Belarus Miocene Dorofeev Zhitkovichi, Supposed late C. pauciflora- (1967c) Gomel, Belarus Miocene type, C. sp. Dorofeev Shiichi, Supposed C. pauciflora- (1968) Gomel, Belarus Miocene- type, C. sp. Pliocene boundary (Messinian- Zanclean) Dorofeev Mamontova Supposed C. flagellata, (1969), Gora, Yakutia, Miocene C. Nikitin Russian pauciflori- (1976) Federation formis, C. rostrata- type, C. sp. Dorofeev Polevskoy, Supposed early C. dorofeevii, (1970) Urals, Russian Miocene C. spp. Federation Dorofeev Khapchan- Supposed late C. omoloica, C. (1972) Khaya, Omoloy Miocene subg. Vignea, river, C. sp. Yakutia, Russian Federation Dorofeev Timirdeckh- Supposed C. spp. (1972) Khaya, Omoloy Miocene river, Yakutia, Russian Federation Dorofeev Novaya Zhizn', Supposed C. spp. (1988) Petrovka, Miocene Ploskoye and Sosnovka, Tambov oblast, Russian Federation Dorofeev Dimitriyevka, Supposed middle C. dorofeevii?, (1988) Kuznetsovka, Miocene C. sp. Lavrovo, Pushkari, and Vol'naya Vershina, Tambov oblast, Russian Federation Dorofeev Dmitriyevka, Supposed late C. spp. (1988) bakumovka, Miocene Beryozovka, (Tortonian) Malinovka, Selezni, and Vol'naya Vershina Erdei and Bukkalja Late Miocene C. sp. Magyari formation, (Messinian, (2011) Bukkabrany, NE 7.5-6.8 Ma) Hungary Friis Odderup Early to middle C. spp., C. (1985) formation, Miocene dorofeevii? Fasterholt, (Burdigalian to Denmark early Langhian, 20.44-15 Ma aprox; cf. Rasmussen et al., 2010) Gregor Achldorf, Late middle C. (1982) Molasse, Miocene klettvicensis?, Germany (Serravallian; C. sect. cf. Bohme et Vesicariae- ah, 2012) type Heer Oehningen Middle Miocene C. amissa *, C. (1859) beds, (Serravallian, effossa *, C. Oeningen, 12.7-11.6 Ma; recognita * Badcn- Paleobiology Wurttcmbcrg, Database, 2015) Germany Heer Hreoavatn- Late Miocene C. rediviva * (1868) Stafholt (Messinian,7-6 Formation, Ma; Denk et al., Hreoavatn, 2011) Iceland. Heer Kenai Group, Late early to C. servata * (1869c) Port Graham, early middle Alaska, USA Miocene (Burdigalian- Langhian; cf. Wolfe & Tanai, 1980) Knobloch Dubnany and Late Miocene C. flagellata, (1981a) Prusanky, (Messinian) C. sp. Moravia, Czech Republic Kovar- Koflach/ Early Miocene C. sp. Eder et Voistberg (Burdigalian; al. (2001) lignite area, 19-17.6 Ma) Oberforf, Styria, Austria Kovar- Mataschen, Late Miocene C. flagellata Eder et Steiermark, ?, C. sp. al. Austria (2006); see also Kovar- Eder and Hably (2006) Kownas Dobrzyn nad Middle Miocene C. lilpopi * (1955) Wista, Poland (Worobiec et al., 2012) Krausel Mainz Castell Early Miocene cf. Carex * (1938) Kvacek et Most Early Miocene C. sp. al. Formation, (2004); Most Basin, Kvaiek and Holesice and Teodoridis Tuchorice, (2007) North Bohemia, Czech Republic Lancucka- "Gdow Bay" Late Miocene C. rostrata- Srodoniowa formation, (Tortonian) type, C. spp. (1966) between Wieliczka, Bochnia, and Gdow, Poland Lancucka- Nowy Sacz Supposed late C. acutiformis- Srodoniowa Basin, Poland early Miocene type, C. (1979) (late elongatoides, Burdigalian; C. flavicarpa, Kovar-Eder et C. al., 2008) globosiformis, C. loliacea- type, C. plicata, C. pseudocyperoi- des, C. strigosoides, C. ungeril, Carex subg. Vignea, Carex sect. Glareosae-type Lahcucka- Wielicza, Early middle C. spp. Srodoniowa Poland Miocene and (Langhian, Zastawniak 15.032-13.82 (1997) Ma; Hohenegger et al., 2014) Lancucka- Poznan clay Late Miocene C. spp. Srodoniowa Formation, (Tortonian- etal. Sosnica, Messinian, 9-5 (1981) Wroclaw, Ma; cf Dyjor et Poland al., 1998) Lesiak Orawa-Nowy Middle Miocene C. acutiformis- (1994) Targ Basin, type, C. Lipnica Mala, globosiformis, Poland C. globosiformis?, C. loliacea- type, C. pilulifera- type, C. pseudocyperoi- des, C. ungeril, C. spp. Mai Hartau, Supposed early C. (1964), Saxony, Miocene acutimontana, Mai (1999) Germany C. hartauensis, C. lusatica, C. pseudocyperoi- des Mai Wischgrund, Supposed late C. flavicarpa, (1989), Brandenburg, Miocene C. graciosa, C. Mai (2000) Germany hostianoides, C. plicata Mai Klettwitz, Supposed middle C. (1989), Brandenburg, and late elongatoides, Mai (2000) Germany (more Miocene (more C. jlagellata, than 10 than 10 C. graciosa, C. sampbng sites, different hartauensis, C. different sampling sites, hostianoides, layers in a 50 different C. maiana, C. m thick layers, in a 50 marchiaca, C. succession) m thick plicata, C. succession) pseudocyperoi- des Mai (1999) Berzdorf, Supposed middle C. Hangcndes, Miocene canescentoidea, Saxony, C. eigensis, C. Germany graciosa, C. hartauensis, C. limosioides, C. lusatica, C. pseudocyperoi- des Mai (1999) Kaltwasser, Supposed early C. graciosa Saxony, Miocene Germany Mai (1999) Lubbenau, Supposed early C. Brandenburg, Miocene klettvicensis Germany Mai (1999) Mittelherwigs- Supposed early C. dorf, Miocene pseudocyperoi- Saxony, des Germany Mai (1999) Oberoderwitz, Supposed early C. acutimontana Saxony Germany Miocene Mai (1999) Reichwalde, Supposed early C. lusatica Saxony, Miocene Germany Mai (1999) Sandforstgen, Supposed early C. Saxony, Miocene pseudocyperoi- Germany des Mai (1999) Schlabendorf, Supposed early C. graciosa, C. Brandenburg, Miocene hartauensis, C. Germany limosioides, C. plicata, C. pseudocyperoi- des Mai (1999) Staakow, Supposed early C. marchica Brandenburg, Miocene Germany Mai (1999) Zittau, Supposed early C. pseudocype- Saxony, Miocene roides Germany Mai (2000) Brothen, Supposed middle C. graciosa, C. Saxony, Miocene hostianoides Germany Mai (2000) Brothen, Supposed late C. elongatoides Saxony, Miocene Germany Mai (2000) Crinitz, Supposed late C. Brandenburg, Miocene pseudocyperoi- Germany des Mai (2000) Grossrarchen, Supposed late C. Brandenburg, Miocene canescentoidea, Germany C. elongatoides, C. Jlagellata, C. remotoides Mai (2000) Kausche-Klara Supposed late C. graciosa, C. II, Miocene hartauensis, Brandenburg, Germany Mai (2000) Kleinleipisch Supposed middle C. klarae, C. (three Miocene pseudocyperoi- outcrops), des, Brandenburg, C. protogaea C. Germany graciosa. Mai (2000) Kleinleipisch- Supposed late C. limosioides, Roemerkeller, Miocene C. maiana, C. Brandenburg, plicata C. Germany hartauensis, Mai (2000) Kostebrau, Supposed late C. limosioides, Brandenburg, Miocene C. maiana, C. Germany maianal, C. graciosa, C. plicata C. graciosa Mai (2000) Meuro, Supposed middle C. graciosa Brandenburg, Miocene Germany Mai (2000) Nochten, Supposed middle C. graciosa, Saxony, Miocene Germany Mai (2000) Plessa, Supposed middle C. hartauensis, Brandenburg, Miocene C. Germany hostianoides, C. plicata, C. pseudocyperoi- des, C. remotoides C. hartauensis Mai (2000) Rauno, Supposed late C. graciosa Brandenburg, Miocene Germany Mai (2000) Rietschen, Supposed middle C. graciosa Saxony, Miocene Germany Mai (2000) Tetta, Saxony, Supposed middle C. graciosa Germany Miocene Mai (2000) Welzow, Supposed middle C. graciosa, C. Brandenburg, and late limosioides Germany (two layers) Mai (2000) Wischgrund- Miocene C. hostianoides Boschung, Supposed middle Brandenburg, Miocene Germany Mai and Brandis, Supposed early C. hartauensis Walther Saxony, Miocene (1991) Germany Mai and Golpa-Nord, Supposed early C. hartauensis Walther Saxony, Miocene (1991) Germany Mai and Leipnitz, Supposed early C. Walther Saxony, Miocene canescentoidea, (1991) Germany Mai and Skoplau, Supposed early C. hartauensis Walther Saxony, Miocene C. hartauensis (1991) Germany Mai and Hartau, Supposed early C. hartauensis Walther Saxony, Miocene (1991) Germany Mai and Nochten-Ost, Late Miocene C. pilulifera- Wahnert Sachsen, (Messinian; type, C. (2000) Germany Paleobiology praehirta, C. Database, pseudocyperoi- des, C. szafeti Matthews Lava Camp, 2015) Late C. rostrata- and Seward Miocene type Ovenden Peninsula, (Messinian, (1990) Alaska, USA >5.7 [+ or -] 0.2 Ma) Matthews Upper Early middle C. sp. and Ramparts, Miocene Ovenden Porcupine (Langhian, 15.7 (1990) River, Alaska, 14.4 Ma; Kunk USA et al., 1994) Meller Hintersch- Late Miocene C. (2011) lagen, (Tortonian) elongatoides?, Austria C. sp. Meller and Mataschen clay Late middle C. sp. Hoffman pit, Fehring, Miocene (2004) Styria, (Serravallian, Austria ~11.6 Ma; see Kovar-Eder & Hably, 2006) Melleretal. Oberdorf, Late early C. spp. (1999) Styria, Miocene (late Austria Burdigalian) Momohara Tokiguchi Late Miocene C. maackii- and Saito Porcelain Clay (Tortonian, 9 type, C. (2001) Fonnation, Ma) vesicaria- Tajimi City, type, Carex Japan sect. Molliculae- type, Care.x sect. Phacocystis- type, Carex sect. Lageniformes- type, Carex sect. Rhomboidales- type Negru Bolshoy Late Miocene C. carpophora?, (1986) Fontan, (Tortonian- C. flagellata, Odessa, Messinian, 8.0- C. graciosa, C. Ukraine 6.0 Ma; cf. limosioides, C. Vasiliev et pseudocyperoi- al., 2011) des Nikitin Kireevskoe, Supposed early C. frequens (2006) Western Miocene Siberia, (Burdigalian) Russian Federation Nikitin Gorskiy Log, Supposed early C. zhilinii (2006) European Miocene Russia, (Aquitanian) Russian Federation Nikitin Kruglikovo, Supposed middle C. major (2006) Novosibirsk Miocene oblast', Western Siberia, Russian Federation Ortuno et Pruedo, Val Late Miocene C. sp. al. (2013) d'Aran, (Tortonian, Llcida, Spain 11.1-8.7 Ma). Palamarev Katina and Supposed C. flagellata and Balscha, Miocene- Petkova Bulgaria Pliocene (1987), boundary Palamarev (1994) Raniccka- Konin brown Supposed late C. flagellata, Bobrowska coal, Miocene C. szaferi, C. (1959) Morzyslaw, subg. Carex, C. Poland sp. Slavik Tuchorie, Early Miocene C. antiqua * (1869) Czech Republic Szafcr Gliwice, Supposed late C. flagellata?, (1961) Poland Miocene C. (Tortonian) pseudocyperoi- des? Teodoridis Brestany Middle or late cf. Carex * and Kvacek clays, Miocene (cf. (2006) Holesice Grygar & Mach, Formation, 2013) Most Basin, Czech Republic Thomasson Ash Hollow Middle or late C. graeeii (1983) Formation, Miocene (13.6- Ogalalla 4.9 Ma) Group, Garden County, Nebraska, USA van der North Rhine- Supposed late C. acutifomns- Burgh Westphalia, Miocene type, C. (1987) Germany flagellata, C. hostiana-type, C. klettvicensis?, C. sp. Velenowski Vrsovice, Supposed early cf. Carex * (1881) Czech Republic Miocene. Zastawniak Gozdnica, Supposed late C. (1992), Poland Miocene. klettvicensisl, Mai (2000) C. elongatoides, C. flavicaipa, C. marchica, C. plicatal, C. spp. PLIOCENE Basilici Sento II, Supposed latest C. flagellata, et al. Piedmont, early Pliocene C. nigra-type, (1997) Italy (Zanclean- C. remota- Piacenzian type, C. boundary; ca. szaferi, C. sp. 3.8-3.5 Ma) Bertoldi Ca' Viettone, Supposed latest C. panormitana- and Turin, early Pliocene type, C. nigra- Martinetto Piedmont, (Zanclean- type, C. (1995) Italy Piacenzian loliacea-type, boundary; ca. C. 3.8-3.5 Ma) paucifloroides, C. pseudocyperus- type, C. sp. von Bulow Trebs Supposed late C. binervis- and Mai borehole, Pliocene type, C. (1992), Hagenow (Piacenzian) blysmoides, C. Mai (2004) District, davalliana- Southwest- type, C. nigra- Mecklenburg, type, C. Germany panormitana- type, C. paucifloroides, C. pseudocyperus- type., C. vesicaria- type, C. spp. Buzek et Vildstejn Supposed C. nigra-type, al. Formation, Pliocene or C. rostrata- (1985), Cheb Basin, probably early type, C. Kvacek and Bohemia, Czech Pleistocene szafei? Teodoridis Republic (cf. Teodoridis (2007) et al., in press) Cavallo Crava di Early Pliocene C. plicatal, C. and Morozzo, (Zanclean; ca. sp. Martinetto Pocapaglia, 4.5-3.8 Ma) (1996) Piedmont, Italy Cavallo Castelletto Supposed late C. atrojusca- and Cervo I, Pliocene type, C. Martinetto Biella, (Piacenzian) elongatoides, (2001) Piedmont, C. lepidocarpa- Italy type, C. panicea-type, C. panormitana- type Cavallo FR S-11, Front Supposedly C. brizoides, and Canavese, Pliocene C. Martinetto Piedmont, pseudocyperus- (2001) Italy type Ciangherotti Ceresole Late Pliocene C. atrofusca- et al. d'Alba, (Piacenzian) type, C. (2007) Piedmont, flagellata, C. Italy panormitana- type, C. plicata, C. pseudocyperus- typc, C. sp. Denk et Tjornes, Early Pliocene C. sp. al. (2011) Skeifa, (Zanclean; 3.9- Iceland 3.8 Ma) Dorofeev Bet'ki, Supposed early C. sp. (1956, Tatarstan, to middle 1957b) Russian Pliocene Federation Dorofeev Kamskiye Supposed C. spp. (1956) Polyany, Pliocene Tatarstan, Russian Federation Dorofeev Naberczhnyie Supposed C. spp. (1956) Chelny, Pliocene Tatarstan, Russian Federation Dorofeev Teren'-Sazy, Supposed C. sp. (1956) Tatarstan, Pliocene Russian Federation Dorofeev Matanov sad, Supposed middle C. pauciflora- (1957a, Rostov oblast, Pliocene typc, C. spp. 1966a) Russian Federation Dorofeev Biklyan', Supposed early C. spp. (1957b) Tatarstan, to middle Russian Pliocene Federation Dorofeev Iglino, Supposed C. sp. (1960d) Bashkortostan, Pliocene Urals, Russian Federation Dorofeev Kumurly, Supposed C. spp. (1960d) Bashkortostan, Pliocene Urals, Russian Federation Dorofeev Verkhne- Supposed C. sp. (1960d) tashevo, Pliocene Bashkortostan, Urals, Russian Federation Dorofeev Verkhnyaya Supposed middle C. sp. (1960d) Khabarovka, Pliocene Bashkortostan, Urals, Russian Federation Dorofeev Bash-Shidy, Supposed C. sp. (1962b) Bashkortostan, Pliocene Urals, Russian Federation Dorofeev Izyak, Supposed C. pallescens- (1962b) Bashkortostan, Pliocene type, C. spp. Urals, Russian Federation Dorofeev Karaganka, Supposed C. rostrata- (1962b) Bashkortostan, Pliocene type, C. subg. Urals, Russian Vignea, C. spp. Federation Dorofeev Novokulevo, Supposed C. canescens- (1962b) Bashkortostan, Pliocene type, C. Urals, Russian elongata-type, Federation C. hirta-type, C. pauciflora- typc, C. subg. Vignea, C. spp. Dorofeev Staroisayevo, Supposed C. sp. (1962b) Bashkortostan, Pliocene Urals, Russian Federation Dorofeev Uspenka, Supposed C. subg. (1962b) Bashkortostan, Pliocene Vignea, C. spp. Urals, Russian Federation Dorofeev Chermoluch'ye, Late Pliocene C. rostrata- (1963) Omsk oblast, type Western Siberia, Russian Federation Dorofeev Nur, Pliocene C. sp. (1965) Bashkortostan, Urals, Russian Federation Dorofeev Pyatiletka, Pliocene C. rostrata- (1965) Bashkortostan, type, C. subg. Urals, Russian Vignea, C. spp. Federation Dorofeev Shelkanovo, Pliocene C. sp. (1965) Bashkortostan, Urals, Russian Federation Dorofeev Staroye Pliocene C. pauciflora- (1965) Baryatino, type, C. Bashkortostan, rostrata-type, Urals, Russian C. subg. Federation Vignea, C. sp. Dorofeev Sychovo, Pliocene C. sp. (1965) Bashkortostan, Urals, Russian Federation Dorofeev Urman, Pliocene C. subg. Vignea (1965) Bashkortostan, Urals, Russian Federation Dorofeev Matanov Sad, Middle Pliocene C. pauciflora- (1966a) Rostov oblast, type, C. subg. Russian Vignea, C sp. Federation Dorofeev Sivkovo, Pliocene C. subg. (1967a) Grodno, Vignea, C. sp. Belarus Dorofeev Tabola, Pliocene C. pauciflora- (1967a) Grodno, type, C. subg. Belarus Vignea, C. sp. Dorofeev Smychok, Pliocene C. pauciflora- (1968a) Gomel, Belarus type, C. sp. Dorofeev Kholmech, Supposed C. earpophora, (1971), Gomel, Belarus Pliocene C. flagellata, Velichkevich C. klarae, C. and paucifloroides, Zastawniak C. szaferi, C. (2003) yakubovskayae, C. subg. Carex, C. sp. [Caricoid clade or section Ovales], C. sp. Dorofeev Simbugino, Supposed late C. flagellata, (1977) Bashkortostan, Pliocene C. rostrata- Urals, Russian (Piacenzian) pliocenica, C. Federation szaferi, C. yakubovskayae, C. sp. (subgenus Vignea), C. spp. Dorofeev Dan'shino, Supposed C. rostrata- (1979) Lipetsk Pliocene type, C. spp. oblast, Russian Federation Dorofeev Dvorets, Gomel Supposed late C. blysmoides, (1986) oblast', Pliocene C. Belarus; see (Piacenzian) paucifloroides, also C. rostrata- Velichkevich type, C. sp. (1975, 1990), Velichkevich and Zastawniak (2007), and Yakubovskaya (1982) Dorofeev Kureyka, Supposed C. spp. and Krasnoyarsk Pliocene Mezhvilk kray, Russian (1956) Federation Dorofeev Mamontova Supposed C. and Gora, Yakutia, Pliocene paucifloroi- Tjulina Russian des?, (1962) Federation C. spp. Dyjoretal. Ruszow, Zary, Supposed C. flagellata, (1998), Poland Pliocene C. gothanii, C. Mai and klettvicensis, Wahncrt C. (2000) klettvicensis?, C. lasiocarpa- type, C. hartauensis, C. spp. Girotti et Temi, Umbria, Pliocene- C. al. (2003) Italy Pleistocene pseudocyperus- boundary type (Piacenzian- Gelasian) Jimenez- W Piedmont Early to late C. panormitana- Mejias and (Lcvone, Nole Pliocene (4- type martinetto Canavese), 2.6 Ma) (2013) Italy; see also Mai (1995) and martinetto (1994a, b). Knobloch Neusield, Pliocene C. flagellata? (1981b) Burgenland, Austria Knobloch Bystrovany, Late Pliocene C. elongata- (1989) Moravia, Czech (Piacenzian) type, C. Republic nigi'a-type, C. pseudocyperus- type, C. szaferi Knobloch Holice, Late Pliocene C. nigra-type (1989) Moravia, Czech (Piacenzian) Republich Madler Frankfurt- Supposed C. spicata-type (1939) Klarbcckcn, Pliocene Hesse, Germany (probably Piacenzian) Mai et al. Rippersroda, Pliocene C. flagellata, (1963), Thuringia, C. flavicarpa, Mai and Germany C. panormitana- Walther type, C. (1988) praehirta, C. pseudocyperns- type, C. ungeri Mai Kranichfeld, Supposed C. (1965), Thuringia, Pliocene elongatoides, Mai and Germany C. flagellata, Walther C. gothanii, C. (1988) panormitana- type, C. pilulifera- type, C. riparia-type, C. strigosoides, C. ungeri Mai RDB Quarry- Late Pliocene C. acutiformis- (1995), Villafranca (Piacenzian, type, "C. Jimenez- d'Asti, ca. 3.3-3.0 Ma) atrofusca"- Mejias and Piedmont, type, C. martinetto Italy carpophora, C. (2013) flagellata, C. laevigata- type, C. loliacea-type, C. panormitana- type, C. plicata, C. pseudocyperus- type, C. remota-type, C. szaferi, C. sp. Mai (2004) Lubtheen 2-75 Supposed C. klarae, C. and 3-75, Pliocene cf. Mecklenburg- klettvicensis, Vorpommem, C. praehirta Germany Mai (2004) Ruterberg, Supposed late C. Mecklenburg- Pliocene elongatoides!, Vorpommem, (Piacenzian) C. nigra-type, Germany C. pseudocyperus- type Mai and Berga, Supposed C. binervis- Walther Thuringia, Pliocene type, C. (1988) Germany carpophora, C. flagellata, C. helmensis, C. laevigata- type, C. paucifloroides, C. pilulifera- type, C. plicata, C. rostrata-type, C. szaferi Mai and Gerstungen, Supposed C. binervis- Walther Thuringia, Pliocene type, C. (1988) Germany elongatoides, C. pseudocyperus- type, C. rostrata-typa, C. szaferi, Mai and Kalten- Pliocene C. acutiformis- Walther sundheim, type, C. (1988) Thuringia, flavicatpa, C. Germany nigra-type, C. panormitana- type, C. riparia-type Mai and Nordhausen, Supposed C. nigra-type, Walther Thuringia, Pliocene C. panormitana- (1988), Germany type, C. and this paucifloroides, paper C. pseudocyperus- type, C. riparia-type, Martinetto Barbania and Supposed C. (1994a), La Cassa, Pliocene elongatoides, Jimenez- Piedmont, C. loliacea- Mcjias and Italy type, C. Martinetto panormitana- (2013) type, C. paucifloroi- des?, C. plicata, C. pseudocyperus, C. spp. Martinetto Dunarobba, Supposed C. flagellatal, (1994a), Umbria, Italy Pliocene, or C. loliacea- Jimenez- early type, C. Mejias and Pleistocene panormitana- Martinetto (Piacenzian- type, C. (2013), Gelasian, 3- pseudocyperus- Martinetto 1.5 Ma) type et al. (2014b) Martinetto Stura di Lanzo Late Pliocene "C. atrofusca"- (1994b), River Fossil (Piacenzian, type, C. Martinetto Forest, 3.0-3.1 Ma) brizoides- et al. Piedmont, type, C. (2007), Italy flagellata, C. Jimenez- loliacea-type, Mejias and C. nigra-type, Martinetto C. plicata, C. (2013) panormitana- type, C. pseudocyperus, C. remota- type, C. rostrata-type, C. spp. Martinetto Roatti, Late Pliocene "C. and Mai Piedmont, (Piacenzian) atrofiisca"- (1996) Italy type, C. panormitana- type, C. pseiidocypems- type Martinetto Front Late Pliocene C. brizoides, et al. Canavese, (Piacenzian, 3 C. pauc if loro (2007) Piedmont, Ma) ides, C. Italy szaferi Matthews Beaver Peat, Early to late C. aquatilis- and Ellesmere Pliocene (4-3 type, C. Ovenden Island, Ma, Matthers & diandra-type, (1990) Nunavut, Tclka, 1997) Carex Canada chordorrhiza- type, C. spp. Matthews Bluefish Supposed C. spp. and Exposure, Pliocene Ovenden Bluefish (1990) Basin, Yukon, Canada Matthews Ch'ijee's Supposed C. aquatilis- and Bluff, Pliocene type, C. Ovenden Bluefish rostrata-type, (1990) Basin, Yukon, C. spp. Canada Matthews Cone Bluff, Pliocene C. spp. and Porcupine Ovenden River, Alaska, (1990) USA Matthews Gubik Late Pliocene C. aquatilis- and formation, (Piacenzian) to type, C. spp. Ovenden Fish Creek, early (1990) Alaska, USA Pleistocene (Gelasian) (3- 2,4 Ma) Matthews Kugruk River Pliocene C. aquatilis- and sites, Seward type, C. Ovenden Peninsula, rostrata-type (1990) Alaska, USA Matthews Lost Chicken Early late C. spp. and Mine, Alaska, Pliocene Ovenden USA (Piacenzian, (1990) 2.9 [+ or -] 0.4 Ma; Matthews et al., 2003) Matthews Niguanak, Pliocene C. spp. and Alaska, USA Ovenden (1990) Matthews Upper Pliocene C. spp. and Ramparts, Ovenden Porcupine (1990) Canyon, Alaska, USA Nikitin Krivoborie, Supposed C. rostrata- (1957) Voronezh, Pliocene pliocenica, C. Russian yakiibovskayae Federation Reid and Reuver, Pliocene C. flagellata, Reid Swalmen and C. szaferi, C. (1915) Brunssum, subg. Carex, C. Limburg, the spp. Netherlands Rudolph Nova Ves, Supposed C. lasiocarpa- (1935) Bohemia, Czech Pliocene type, C. spp. Republic Szafer Kroscienko, Supposed C. carpophora, (1938, Poland Pliocene C. flagellata, 1947), and C. panicea- this paper type, C. panormitana- type, C. pseudocyperus- type, C. szaferi, C. spp. Szafer Mizema, Supposed C. flagellata, (1954), Czorsztyn, Pliocene, C. elongata- and this Poland several layers, type, C. paper most probably panicea-type, early C. panormitana- Pleistocene at type, C. the top rostrata-type, C. pseudocyperus- typc van der Fortuna- Supposed C. colchica- Burgh Garsdorf, Pliocene (early type, C. (1978, Bergheim, Zanclean based Jlagellata, C. 1983) Germany on the flora) lasiocarpa- typc, C. m'gra- type, C. panicea-type, C. pallescens- type, C. pseudocyperus- type?. C. rostrata-type van der Frechen, North Supposed C. rostrata- Burgh Rhine- Pliocene (early type, C. sp. (1983) Westphalia, Zanclean based Germany on the flora) van der Hambach mine, Pliocene C. acutiformis- Burgh Duren, Germany (supposed type, C. (1983), Bmnssummian elongatoides, van der based on the C. flagellata, Burgh and flora) C. flavicarpa, Zettcr C. (1998) klettvicensis?, C. panicea- type?, C. pilulifera- type, C. riparia-type, C. strigosoides, C. szaferi, C. sp. Velichkevich Dvorets, Gomel Supposed late C. blysmoides, (1975, oblast'. Pliocene C. 1990), Belarus; see (Piaccnzian) paucifloroides, Velichkevich also Dorofeev C. and (1986) and yakiibovskayae Zastawniak Yakubovskaya (2007) (1982). Yakubovskaya Dvorets, Gomel Supposed late C. flagellatat, (1982) oblast', Pliocene C. Belarus; see (Piaccnzian) yakubovskayae, also Dorofeev C. spp. (1986), Velichkevich (1975, 1990), and Velichkevich and Zastawniak (2007). Yakubovskaya Kholmech, Supposed middle C. (1982) Gomel oblast', part of early paucifloroides, Belarus Pliocene C. cf. szaferi, (Zanclean) C. yakubovskayae, C. spp. Yamakawa Asanuki, Late Pliocene C. aphanolepis- et al. (in (Conan, Shiga (Piaccnzian, typc, C. press) Prefecture, 2.6 Ma) capricomis- Japan type, C. dispalata- type, C. ischnostachya- typc This paper Gorsbach, Pliocene C. panormitana- Thuringia, type Germany This paper Momello- Supposed C. Lanzo, Italy Pliocene paucifloroides EARLY PLEISTOCENE Cavallo Castelletto Supposed early C. atrofusca- and Cervo II, Pleistocene typc, C. martinetto Biella, (Gelasian) carpophora, C. (2001) Piedmont, elongatoides, Italy C. flavicarpa, C. lepidocarpa- typc, C. szaferi, C. spp. Jimenez- Casnigo, Early C. elata-type Mejias and Lombardy, Pleistocene Martinetto Italy (Gelasian) (2013) Jimenez- Buronzo, Supposed early C. elata-type Mejias and Piedmont, Pleistocene Martinetto Italy (Gelasian) (2013) Martinetto San Pietro di Supposed early C. elata- et al. Ragogna, Pleistocene type?, C. (2012) Friuli- (Gelasian) rostrata-type, Vcnctia C. sp. Giulia, Italy Reid and Tegelen clays, Early C. echinata- Reid Tegelen-sur- Pleistocene type, C. hirta- (1907a) Meuse, (Gelasian; van type, C. Limburg, the den Hoek riparia-type, Netherlands Ostende, 2003; C. Cohen et al., yakubovskayae?, 2013) C. subg. Carex, C. spp. Reid and Castle Eden, Supposed early C. acutiformis- Reid United Kingdom Pleistocene type, C. (1907b) (Gelasian; see dioica-type, C. Jones and Keen hirta-type, C. (1993) laevigata- type, C. muricata-type, C. riparia- type, C. rostrata-type, C. vesicaria- type, C. spp. Westerhoff Belfeld, the Early C. davalliana- et al. Netherlands Pleistocene type, C. (1998), (Gelasian) lepidocarpa- also this type, C. nigra- paper type, C. pseudocyperus- type, C. rostrata-type, C. sp. Westerhoff Maalbeck, Early C. flagellata, et al. Tegelen, the Pleistocene C. sp. (1998) Netherlands (Gelasian) This paper Oebel, Early C. panormitana- Tegelen, the Pleistocene type Netherlands (Gelasian) Records are given by alphabetical and chronological order of publication within geological epochs. Names provided correspond to our final identifications; when the name in the original publication was a different one, it must be traced back in the checklist. Remains doubtfully treated under Carex are accompanied by an asterisk (*). Those confident Carex records of doubtful taxonomic relationships are accompanied by a question mark (?). Records cited from boundaries between epochs are cited within the older epoch
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Title Annotation: | p. 301-345 |
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Author: | Jimenez-Mejias, P.; Martinetto, E.; Momohara, A.; Popova, S.; Smith, S.Y.; Roalson, E.H. |
Publication: | The Botanical Review |
Date: | Sep 1, 2016 |
Words: | 26957 |
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