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

Article Details

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Title Annotation:	p. 301-345
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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Topics:	Cyperaceae Distribution Identification and classification Physiological aspects Fossil plants Distribution Identification and classification Physiological aspects Plants, Fossil Distribution Identification and classification Physiological aspects