The taxonomy and morphological diversity of leaves of Paraprotophyllum (Platanaceae) from the Late Cretaceous of Sakhalin Island

ISSN 0031�0301, Paleontological Journal, 2010, Vol. 44, No. 10, pp. 1270–1280. © Pleiades Publishing, Ltd., 2010

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INTRODUCTION

Elongated entire leaves with a cordate or roundedbase, dentate�emarginate margin, pinnate craspedo�dromous venation, and veins repeatedly dichotomiz�ing near the leaf margin are very characteristic ele�ments of the Late Cretaceous flora of the SakhalinIsland. During the long history of the study of thisCretaceous flora the taxonomic position of such leaveswas defined in different ways: most often, they wereconsidered as members of the genera ProtophyllumLesquereux (Kryshtofovich and Baikovskaya, 1960;Kutuzkina, 1974; Budantsev, 1975; Krassilov, 1979) orPseudoprotophyllum Hollick (Vachrameev, 1976).Herman (1984) described a new genus, Paraprotophyl�lum Herman, based on this material.

Describing Paraprotophyllum, Herman (1984)included into the genus two species: P. ignatianum(Kryshtofovich et Baikovskaya) Herman, which wasoriginally reported from Sakhalin Island as Protophyl�lum ignatianum Kryshtofovich et Baikovskaya (Krysh�tofovich and Baikovskaya, 1960), and the new speciesP. pseudopeltatum Herman, from the northwesternKamchatka Peninsula. However, the characteristics ofboth species were nearly completely based on theKamchatka material. Numerous specimens fromSakhalin Island, treated in various ways by differentscientists, were not revised.

The present paper provides a detailed characteriza�tion of the morphology and variability of leaves ofParaprotophyllum from the classic collections amassedby Heer and Kryshtofovich. Such work seems neces�sary for the further study of numerous new finds of thegenus from several Late Cretaceous localities of Sibe�ria and the Russian Far East (Golovneva, 2005, 2006).Currently, the comparison between the new materials

and type material is difficult, since the Sakhalin mate�rial was not revised, and even photographs of mostspecimens were not published.

Leaves of Paraprotophyllum were first collected bySchmidt and Glen at the coast of the Strait of Tartary,near the village of Mgachi in 1873, and handed over toHeer, who used them for the description of two newspecies of the genus Viburnum L.: V. schmidtianumHeer and V. spinulosum Heer (Heer, 1878).

Later, leaves of Paraprotophyllum were described byKryshtofovich and Baikovskaya (1960) in CretaceousFlora of Sakhalin Island mostly under the genericname Protophyllum: P. ignatianum Kryshtofovich etBaikovskaya, P. longifolium Kryshtofovich et Baik�ovskaya, P. praestans Lesquereux, and P. sternbergiiLesquereux. Other morphologically similar leafimprints from these beds were assigned to earlierdescribed species of Viburnum (V. whymperi Heer andV. lesquereuxii Ward), Cissus L. (Cissus sp.), Dryophyl�lum Debey ex Saporta (D. cf. bruneri Ward), and VitisL. (V. xantholithensis).

In his paper on the Late Cretaceous flora of Sakha�lin Island, Krassilov (1979) united Viburnum schmid�tianum and Protophyllum ignatianum under the nameProtophyllum schmidtianum (Heer) Krassilov. Basedon unclear drawings of Heer (1878), Herman (1984)concluded that these species were not identical andchose Protophyllum ignatianum as the type species ofParaprotophyllum. I have restudied the collection ofHeer, which is kept at the Komarov Botanical Institute(BIN). The collection contains several leaf fragmentsof Viburnum schmidtianum and a leaf with a damagedapex. The last specimen (BIN, no. 3/14) was chosen asthe lectotype of V. schmidtianum (Figs. 1a, 1c). Theimprint clearly shows a rounded base, pinnate�pal�

The Taxonomy and Morphological Diversity of Leaves of Paraprotophyllum (Platanaceae) from the Late Cretaceous

of Sakhalin IslandL. B. Golovneva

Komarov Botanical Institute, Russian Academy of Sciences, ul. Professora Popova 2, St. Petersburg, 197376 Russiae�mail: [email protected]

Received February 5, 2009

Abstract—Collections of leaves of the genus Paraprotophyllum Herman amassed by Heer and Kryshtofovichfrom Sakhalin Island are revised. The morphology and variability of P. schmidtianum (Heer) Golovneva,comb. nov. are analyzed.

Key words: Cretaceous, Sakhalin Island, Platanaceae.

DOI: 10.1134/S0031030110100059

PALEONTOLOGICAL JOURNAL Vol. 44 No. 10 2010

THE TAXONOMY AND MORPHOLOGICAL DIVERSITY OF LEAVES 1271

mate venation with five pairs of secondary veins, onepair of subbasal veins, branchlets of secondary veinsdichotomizing near the leaf margin, and veins craspe�dodromously ending in small teeth near the leaf base.The leaf outline and total number of veins remainunknown, since the apex is lacking. All characters ofthis specimen correspond to characteres of otherleaves of Paraprotophyllum from the Late Cretaceousfloras of Sakhalin Island. Herman (1984) believed thatsuch characters as a rounded base and narrow seg�ments intercepted by lowermost lateral veins areuntypical of the genus Paraprotophyllum; however, thecombination of these characters occur relatively often

in small specimens of the Sakhalin Paraprotophyllum(Pl. 6, figs. 2, 3) and was also observed in medium�sized and large leaves (Pl. 6, fig. 7 and Figs. 2c, 2i,respectively). Therefore, I share the opinion expressedby Krassilov that the leaves described as Viburnumschmidtianum and Protophyllum ignatianum belong toone species, with the name Viburnum schmidtianumhaving priority. A new combination, Paraprotophyllumschmidtianum (Heer) Golovneva, comb. nov., is created.Remains of Viburnum spinulosum are a few leaf fragmentswith a well�preserved leaf margin (Figs. 1b, 1d).The mode of dentation and veins branching near the

(a)

1 cm

(b)

(c) (d)

Fig. 1. Leaves of Paraprotophyllum schmidtianum (Heer) Golovneva, comb. nov. collected by Schmidt and Glen, Late Cretaceous,Sakhalin Island: (a, c) lectotype, BIN, no. 3/14 (Heer, 1878, as Viburnum schmidtianum Heer); (b, d) fragment of leaf margin,BIN, no. 4/93 (Heer, 1878, as Viburnum spinulosum Heer). Scale bar 1 cm.

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Plate Plate 6

1 2

3 4

6

5 7



leaf margin testify to the affinity of these specimens toParaprotophyllum.

The reexamination of all specimens of Paraproto�phyllum from the Late Cretaceous of Sakhalin Islandhas shown that they belong to one variable species.The name Paraprotophyllum schmidtianum (Heer)Golovneva has priority for this species. The extendeddescription of this species given below is based on therevision of the collections of Heer and Kryshtofovichfrom Sakhalin Island.

MATERIAL

This paper deals with investigation of the collec�tions on which the classic monographs by Heer (1878)and Kryshtofovich and Baikovskaya (1960) about Cre�taceous floras of Sakhalin Island were based. The col�lection studied by Heer was amassed by Schmidt andGlen from river outcrops around the village of Mgachi.It is currently kept at the laboratory of paleobotany ofBIN in St. Petersburg, as collection BIN, no. 3. Theother collection was amassed by Kryshtofovich in1917–1929 during seven field seasons. Apart fromspecimens collected by Kryshtofovich, it also containsspecimens collected by the geologist P.I. Polevoi andcollector I.P. Polevoi. This collection is kept at theCentral Research Geological Exploration Museumnamed after the academician F.N. Chernyshev(CNIGR Museum) in St. Petersburg as CNIGR,no. 6256. Besides several specimens the drawings ofwhich were previously published, it includes manyspecimens that have never been imaged. In total, Ihave examined about 80 specimens, of which about35 leaves were complete. The material comes from thefollowing localities of the western coast of SakhalinIsland:

(1) locality 24, Polovinka mine (former Eriksonmine), Polovinka River, 6 km from the sea;

(2) locality 27, Polovinka mine, Polovinka River,6 km from the sea;

(3) locality 146, Lyudvigova pad’ (valley), a tribu�tary of the Aleksandrovka River;

(4) locality 195, Lyudvigova pad’, a tributary of theAleksandrovka River;

(5) locality 424, coast of the Strait of Tartary, directlyto the north of the wharf of the village of Mgachi;

(6) locality 461, coast of the Strait of Tartary, to thenorth of the pad’ Varnak (Vakhtmngy);

(7) locality 463, coast of the Strait of Tartary, themouth of the first creek valley to the south of thepad’ Vakhtmngy;

(8) locality 491, Kommunal’nogo khozyaistvamine (former Petrovskii mine);

(9) locality 726, Vladimirovka River, near the edgeof the ridge, in the upper streams of the brook enteringthe Vladimirovka River from the right opposite to thepad’ Smol’naya;

(10) and locality 906, Nainai River, Nazarovskiibrook.

As far as I can suppose from by the descriptions ofthe localities and plant assemblages (Kryshtofovichand Baikovskaya, 1960), they all belong to the ArkovoFormation, mostly to its lower portion, with theexception of locality 424, which belongs to the upperpart of the Arkovo Formation. Krassilov (1979) con�sidered the plant assemblage from the lower part of theArkovo Formation as Ainusskii floristic horizon of theConiacian and the plant assemblage from the upperpart of the Arkovo Formation as Gilyatskii floristichorizon of the Santonian.

SYSTEMATIC PALEOBOTANY

Herman (1984) included into the genus Paraproto�phyllum two species: P. ignatianum and the new speciesP. pseudopeltatum. However, creating the combinationP. ignatianum, he referred to an ineffectively publishedbasionym (Kryshtofovich and Baikovskaya, 1960).Describing Protophyllum ignatianum, Kryshtofovichand Baikovskaya (1960) did not indicate the type, whichhas been a requirement condition of a valid publicationsince 1958 (International Code …, 2006, art. 37.1).Later, this name was validated by Kutuzkina (1974),who revised the Platanaceae of Russia in the mono�graph Fossil Angiosperms of the USSR. However, com�posing the combination, Herman reffered to the paperby Kryshtofovich and Baikovskaya but not to the paperby Kutuzkina. According to Article 33.5 of the ViennaCode (2006), the citation by Herman of a paper dated1960 can be considered as a bibliographic error in thecitation of a paper dated 1974 and should be corrected.

Family Platanaceae T. Lestiboudois, 1826

Genus Paraprotophyllum Herman, 1984

Ty p e s p e c i e s. Paraprotophyllum ignatianum(Kryshtofovich et Baikovskaya ex Kutuzkina) Herman(Protophyllum ignatianum Kryshtofovich et Baikovskayaex Kutuzkina: Kutuzkina, 1974, Fossil Angiosperms of theUSSR, p. 149, text�fig. 94). (=Paraprotophyllum schmid�tianum (Heer) Golovneva, comb. nov.).

E x p l a n a t i o n o f P l a t e 6

Figs. 1–5. Leaves of Paraprotophyllum schmidtianum (Heer) Golovneva, comb. nov. from the Late Cretaceous of Sakhalin Island:(1) CNIGR, no. 6256/24; (2) CNIGR, no. 6256/195; (3) CNIGR, no. 6256/313; (4) CNIGR, no. 6256/311a; (5) CNIGR,no. 6256/155a; (6) CNIGR, no. 6256/302; (7) CNIGR, no. 6256/320. Full size.

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GOLOVNEVA

(a)

(b)

(d)(f)

(c)

(g)

(e)

(h) (i)

Fig. 2. Morphological diversity of leaves of Paraprotophyllum schmidtianum (Heer) Golovneva, comb. nov. from collectionamassed by Kryshtofovich, Late Cretaceous, Sakhalin Island: (a) CNIGR, no. 6256/318; (b) CNIGR, no. 6256/300;(c) CNIGR, no. 6256/164; (d) CNIGR, no. 6256/313; (e) CNIGR, no. 6256/170; (f) CNIGR, no. 6256/302; (g) CNIGR,no. 6256/273; (h) CNIGR, no. 6256/155a; (i) CNIGR, no. 6256/168. Scale bar 1 cm.



(a)

(b)

(c)

Fig. 3. Leaves of Paraprotophyllum schmidtianum (Heer) Golovneva, comb. nov. from collection of Kryshtofovich, Late Creta�ceous, Sakhalin Island: (a) CNIGR, no. 6256/281; (b) CNIGR, no. 6256/315; (c) CNIGR, no. 6256/273. Full size.

Paraprotophyllum schmidtianum (Heer) Golovneva, comb. nov.

Plate 6, figs. 1–7; Plate 7, figs. 1–4.

Viburnum schmidtianum Heer: Heer, 1878, Mém. Acad. Impér.Scienc. St.�Pétersbourg, ser. VII, vol. 25, no. 7, p. 43, pl. 11,figs. 4–8.

V. spinulosum Heer: Heer, 1878, p. 44, pl. 11, figs. 9, 10.Protophyllum schmidtianum (Heer) Krassilov: Krassilov, 1979,

p. 112, pl. 35, figs. 1–6, pl. 36, figs. 1, 2.P. ignatianum Kryshtofovich et Baikovskaya: Kryshtofovich

and Baikovskaya, 1960, p. 89, text�figs. 31–34, nomen invalid.,designat. typi omissa; Krassilov, 1973, p. 112, pl. 24, figs. 62–65.

Protophyllum ignatianum Kryshtofovich et Baikovskaya exKutuzkina: Kutuzkina, 1974, p. 149, text�fig. 94.

Paraprotophyllum ignatianum (Kryshtofovich et Baikovskaya)Herman: Herman, 1984, p. 76.

Protophyllum longifolium Kryshtofovich et Baikovskaya: Kry�shtofovich and Baikovskaya, 1960, p. 92, pl. 20, figs. 1, 2, nomeninvalid., designat. typi omissa.

P. longifolium Kryshtofovich et Baikovskaya ex Kutuzkina:Kutuzkina, 1974, p. 148, pl. 68, fig. 3.

P. praestans auct. non Lesquereux: Kryshtofovich and Baik�ovskaya, 1960, p. 91, pl. 19, fig. 2, text�fig. 36.

P. sternbergii auct. non Lesquereux: Kryshtofovich and Baik�ovskaya, 1960, p. 95, pro parte, pl. 19, fig. 3, text�figs. 38, 39.

Viburnum whymperi auct. non Heer: Kryshtofovich and Baik�ovskaya, 1960, p. 112, pl. 21, fig. 4.

V. lesquereuxii auct. non Ward: Kryshtofovich and Baik�ovskaya, 1960, p. 112, pl. 21, fig. 5.

Dryophyllum sp. cf. D. bruneri auct. non Ward: Kryshtofovichand Baikovskaya, 1960, p. 73, pl. 17, fig. 1.

Vitis xantholithensis auct. non Ward: Kryshtofovich and Baik�ovskaya, 1960, p. 110, pl. 20, fig. 3.

Cissus sp.: Kryshtofovich and Baikovskaya, 1960, p. 110, pl. 21,fig. 3.

L e c t o t y p e (designated here): BIN, no. 3/14,Sakhalin Island, coast of the Strait of Tartary, near thevillage of Mgachi, Late Cretaceous (Heer, 1878,pl. 11, fig. 4); Figs. 1a, 1c.

D e s c r i p t i o n (Figs. 1a–1d, 2a–2i, 3a–3c,4c–4f). Leaves vary from narrowly ovate to broadlyovate or, more rarely, elliptical. The leaf base is cordateor, more rarely, rounded, or truncated. Often, the leafbase is slightly asymmetrical. The upper part of the leafgradually tapers into a shortly acuminated apex. Theleaves are small, middle�sized, and large, 4–20 cmlong and 3.5–16 cm wide. Most often, the leaves areabout 10–14 cm long. The leaf margin is emarginate�dentate. The teeth are broadly triangular, with anacute apex and wide low sinuses. Usually, the teeth aresmall, 1–3 mm high; occasionally, they are larger, upto 5 mm high, with or without glands.

The venation is pinnate, occasionally palmate�pin�nate and craspedodromous. There are 6–17 pairs oflateral veins, usually, 9–12 pairs. The lateral veinsdeviate from the midrib at an angle of 40°–50° theyare alternating or connivent in pairs, usually straightor, more rarely, slightly arching. Many leaves have lowerlateral veins which deviate at an angle up to 60°–90°.The longest lateral veins are the second–forth pairs of

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Plate Plate 7

1

2

3 4



veins from the base. Numerous deviations, which theybear in their middle and distal portions, can dichoto�mize once or twice. The lowest (basal) veins can runmore or less in parallel to other secondary veins. In thiscase, they are approximately as long as other second�ary veins and bear five or six basiscopic deviations,which in their turn can branch or dichotomize near themargin. However, the basal veins never exceed a half ofthe leaf lamina length. Those basal veins that deviate at

a greater angle than others are much shorter than thesecondary veins situated above them and usually are inparallel to the deviations of these secondary veins. Someleaves with narrowed bases can have shortened second�ary veins in two or three lower pairs. The width of seg�ments intercepted by lowermost lateral veins dependson the angle at which basal veins deviate and on howmuch cordate is the leaf base. These segments vary inshape from wide with numerous long branching devia�

(a)(b)

(c)

(e)

(d)

(f)

Fig. 4. Leaves of the Platanaceae from the Late Cretaceous of Sakhalin Island, collection of Kryshtofovich: (a, b) rounded ovateleaves described as Protophyllum leconteanum Lesquereux (Kryshtofovich and Baikovskaya, 1960); (a) CNIGR, no. 6256/159;(b) CNIGR, no. 6256/160; (c–f) leaves of Paraprotophyllum schmidtianum (Heer) Golovneva, comb. nov.; (c) CNIGR,no. 6256/301; (d) CNIGR, no. 6256/294; (e) CNIGR, no. 6256/190; (f) CNIGR, no. 6256/298. Scale bar 1 cm.

E x p l a n a t i o n o f P l a t e 7

Figs. 1–4. Leaves of Paraprotophyllum schmidtianum (Heer) Golovneva, comb. nov. from the Late Cretaceous of Sakhalin Island:(1) CNIGR, no. 6256/194; (2) CNIGR, no. 6256/158; (3) CNIGR, no. 6256/155b; (4) CNIGR, no. 6256/164. Full size.

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GOLOVNEVA

tions to narrow with simple short deviations. Secondaryveins can be densely situated or more or less widelyspaced. Usually, narrow elongated morphotypes havemore numerous secondary veins. The secondary veinsappear slightly above (in 2–9 mm) the leaf base. Belowthe secondary veins, one or two pairs of subbasal veinsdeviate. The subbasal veins are usually thin, S�curved,and downwards and laterally directed. Some leavesbear well�developed subbasal veins, which are compa�rable to low deviations of the basal veins by the length,width, and number of deviations. The secondary veinsand their deviations are usually ending craspedodro�mously. Occasionally, semicraspedodromous venationwas observed in the leaf base or near the apex. Tertiaryveins are branchy�scalariform and usually curved intheir middle part. The venation of the forth order ispolygonal� or orthogonal�reticulate.

C o m p a r i s o n. Paraprotophyllum is also knownfrom the Late Cretaceous of the Kamchatka Peninsula(Herman, 1984; Herman and Lebedev, 1991), whereP. ignatianum and P. pseudopeltatum were described.Herman believed that the two species occur in both theKamchatka Peninsula and Sakhalin Island. In myopinion, Kamchatka specimens differ from Sakhalinspecimens, P. pseudopeltatum is absent in SakhalinIsland and P. ignatianum (junior synonym ofP. schmidtianum (Heer) Golovneva, comb. nov.) ismissing in the Kamchatka Peninsula. Leaves whichwere reported from the Kamchatka Peninsula asP. ignatianum differ from the Sakhalin leaves in thelarger sizes, wider segments intercepted by lowermostlateral veins, more intensive bifurcation of secondaryveins, larger teeth with distinct glands, and slightly dif�ferent mode of variability. Small specimens of theKamchatka leaves are rounded or broadly oval; largeleaves, more elongated. Small specimens of theSakhalin leaves are usually narrower than large speci�mens and with less developed segments intercepted bylowermost lateral veins and less branchy secondaryveins. In addition, the Kamchatka leaves have moreregular outlines: I have not detected any “ugly” orsharply asymmetric forms, like those commonlyoccurring in specimens of Paraprotophyllum fromSakhalin Island. The variability of the Kamchatkaleaves is mostly expressed in different outlines of thebase and mode of venation in the lower part of the leaf.I incline that only one species of Paraprotophyllum,P. pseudopeltatum, is present in the Kamchatka Penin�sula, but this point of view needs a more detailed justifi�cation by the reexamination of the Kamchatka material.

The leaves of P. pseudopeltatum from the Kam�chatka Peninsula differ from leaves of P. schmidtianum(Heer) Golovneva, comb. nov. in the prevailingpseudopeltate (cordate with connivent lobes) base,higher deviation of basal veins, and a greater number ofsubbasal veins. The leaves described by Herman (1984)from Sakhalin Island as P. pseudopeltatum differ fromthe Kamchatka specimens of P. pseudopeltatum in theabsence of a pseudopeltate base. The leaf shown in Kry�

shtofovich and Baikovskaya (1960, text�fig. 39) has adamaged base, which most probably was rounded orweakly cordate (Fig. 2i). The specimens shown byKryshtofovich and Baikovskaya (1960) in text�fig. 38and pl. 19, fig. 3 are fragments of large leaves with cor�date bases. I believe that both specimens only differ bysizes from other leaves of P. schmidtianum (Heer)Golovneva, comb. nov. Among drawings in Kryshto�fovich and Baikovskaya (1960), only one shows adeeply cordate base with overlapping inner lobes (Kry�shtofovich and Baikovskaya, 1960, text�fig. 37).Unfortunately, this specimen was lost and the correct�ness of the drawing cannot be checked. Therefore,currently there are no grounds to believe thatP. pseudopeltatum is present in Sakhalin Island. Ingeneral, leaves of Paraprotophyllum from both theKamchatka Peninsula and Sakhalin Island are charac�terized by high and mostly overlapping variability.

Apart from Sakhalin Island and the KamchatkaPeninsula, Paraprotophyllum was reported from theTyl’pegyrgynaiskaya flora of the Koryak Upland (Fil�ippova and Abramova, 1993), Armanskaya flora of theOkhotsk�Chukotka volcanogenic belt (Herman,2004), Antibes flora of western Siberia (Shczepetovet al., 2009), and Vilyui floristic assemblage of theLena�Vilyui Depression (Golovneva, 2005). However,the systematic description of this material has not beenyet published.

R e m a r k s. The morphological study of leaves ofP. schmidtianum has shown that they are highly vari�able. Along with typical elliptic and ovate leaves withcordate bases, the collection contains untypical, oftenirregular forms, which were described by Kryshtofov�ich and Baikovskaya (1960) within different generaand species. The study of numerous specimens fromthe Kryshtofovich collection has revealed transitionalforms between these unusual morphotypes and typicalleaves, and these morphotypes were this way attributedto P. schmidtianum (Heer) Golovneva, comb. nov.

Narrow leaves with pinnate venation weredescribed as Protophyllum longifolium (Kryshtofovichand Baikovskaya, 1960). Transitional morphotypesalso have relatively narrow leaf laminas with densersituated secondary veins and a narrow elongated apex(Figs. 3a–3c). This allows me to consider P. longifo�lium as a synonym of Paraprotophyllum schmidtianum(Heer) Golovneva, comb. nov.

Slightly wider leaf laminas were described as Proto�phyllum praestans (Pl. 7, fig. 4; Fig. 2c) and in part asProtophyllum sternbergii (Fig. 2i). The impression thatthese leaves are shortened is increased by the fact thatthe apices in both specimens are missing, which couldhave been attenuated and long enough, as, for exam�ple, in specimens shown in Figs. 2e, 2h. In other char�acters, these leaves do not differ from other leaves ofParaprotophyllum schmidtianum (Heer) Golovneva,comb. nov.

Leaves with rounded bases and narrow segmentsintercepted by lowermost lateral veins were described



as Viburnum whymperi (Kryshtofovich and Baik�ovskaya, 1960, pl. 21, fig. 4). Such outlines are quitecommon among small and medium�sized Sakhalinspecimens of Paraprotophyllum schmidtianum (Heer)Golovneva, comb. nov., as well as in transitional formsfrom more typical weakly cordate leaves (Pl. 6, figs. 2,3, 7; Fig. 2).

Kryshtofovich and Baikovskaya (1960, pl. 21,fig. 5) described as Viburnum lesquereuxii leaves withirregular outlines and a narrow base (Pl. 7, fig. 1;Fig. 2a). Taking into account the rare occurrence andasymmetry of such forms, I consider them as rareirregular morphotypes of Paraprotophyllum schmid�tianum (Heer) Golovneva, comb. nov. Similarly, twomorphotypes with wider spaced and irregularlybranching secondary veins are considered as rareunusual forms of Paraprotophyllum schmidtianum(Heer) Golovneva, comb. nov. (Figs. 2e, 2f). Earlier,such morphotypes were described as Vitis xan�tholithensis and Cissus sp. (Kryshtofovich and Baik�ovskaya, 1960).

In addition, several Platanus�like non�lobate leavesare kept in Kryshtofovich collection, which are notincluded into Paraprotophyllum schmidtianum (Heer)Golovneva, comb. nov. These specimens weredescribed as Protophyllum leconteanum Lesquereux(Figs. 4a, 4b) and Credneria grewiopsoides Hollick(Kryshtofovich and Baikovskaya, 1960, text�fig. 30).These leaves differ from leaves of Paraprotophyllum inthe shorter and wider laminas with a smaller numberof secondary veins (about six as one can judge by theincomplete specimens). The leaves described as Proto�phyllum leconteanum also differ by a wide roundedapex.

Describing Paraprotophyllum from the KamchatkaPeninsula, Herman also reported one more species ofthis genus: P. cordatum (Samylina) Samylina (Hermanand Lebedev, 1991). The study of the type material ofP. cordatum from the Late Cretaceous flora of theOkhotsk�Chukotka volcanogenic belt showed that itsleaves differ from leaves of other species of Paraproto�phyllum in having a smaller number of secondary veins(five to seven) and by the presence of a pelta. Cur�rently, this species is excluded from Paraprotophyllumand considered within Pseudoprotophyllum (Shczepe�tov and Golovneva, 2009). The name Paraprotophyl�lum cordatum is invalid, since no reference to thebasionym was made by Samylina (1988, p. 105), whoformed this combination. The only specimendescribed as Paraprotophyllum cordatum from theKamchatka Peninsula (Herman and Lebedev, 1991)has nine pairs of secondary veins and a cordate baseand unequivocally belongs to the genus Paraprotophyl�lum. However, its specific affiliation should be revised.This specimen has a wide base, wide segments inter�cepted by lowermost lateral veins, and distinct sub�basal veins. No such morphotypes were found inSakhalin Island.

Herman (1984) considered Paraprotophyllumwithin the order Hamamelidales. The majority of ear�lier authors by morphological characters assigned theSakhalin leaves to the Platanaceae (Kutuzkina, 1974;Budantsev, 1975), or artificial groups of Platanus�likeplants (Vachrameev, 1976), or Platanofolia (Krassilov,1979). Krassilov (1973, 1979) studied the epidermis ofthe upper surface of these leaves and concluded that itwas comparable with that of Liquidambar L. I studiedthe epidermis of Paraprotophyllum from Late Creta�ceous deposits of the Timerdyakhskaya Formation ofeastern Siberia (Golovneva, 2006). The combinationof such characters as anomocytic stomata, compoundbases of trichomes, and capitate glands confirms theplatanaceous affinity of this genus. Rounded glandswith thick cuticular folds radiating from the base,which were described by Krassilov, are very similar tobases of compound trichomes of the Platanaceae,whereas the adaxial epidermis of Liquidambar issmooth and lacks either glands or hairs.

M a t e r i a l. BIN, nos. 3/14, 4/93, CNIGR,nos. 6256/155–158, 161, 164, 168–170, 190, 193,194, 272, 273a, 273b, 276a, 276b, 277a, 277b, 281,294, 297–308, 310–316a, 316b, 318–322, SakhalinIsland, Arkovo Formation, Late Cretaceous.

ACKNOWLEDGMENTS

The study was supported by the Russian Founda�tion for Basic Research, project no. 07�04�01227 andthe Program of the Presidium of the Russian Academyof Sciences “Biodiversity”.

REFERENCES

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The taxonomy and morphological diversity of leaves of Paraprotophyllum (Platanaceae) from the Late Cretaceous of Sakhalin Island

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