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Clubmosses with branched microphylls
Clubmosses with branched microphylls
Plants in the Protolepidodendrales are from the Middle Paleozoic and may be the ancestors of the enormous scale trees during the Carboniferous. One defining feature is that these herbaceous plants had forked or branched microphylls, which is unique among clubmosses. Microphylls, by definition, are supposed to be leaves with unforked veins, but this group exhibits an evolution of this feature within the lycophytes.
Ecology & Form
Ecology & Form
Stems
Stems
Herbaceous or sub-woody with isotomous and anisotomous branching;
Prostrate rhizome was probably indeterminate
Leaves
Leaves
Branched microphylls for leaves
No other lycophytes display forked leaves
Some leaves have a ligule
Geologic Age
Geologic Age
Classification
Classification
└Protolepidodendrales †
Diversity
Diversity
Archaeosigillariaceae †
Archaeosigillaria †
Archaeosigillaria †
Vanuxem, 1842; Kidston, 1901
A. conferta
Menéndez, 1965
Devonian of Puerto Rico
Originally named Cyclostigma confertum
A. kidstonii
Kräusel & Weyland, 1949
Mississippian of the UK
A. minuta
Lejal-Nicol, 1970
Late Devonian of Niger
A. nathorstii
Kräusel & Weyland, 1949
Mississippian of Norway
A. picosensis
Kräusel & Dolianti, 1957
Early Devonian of Brazil
A. primaeva
White, 1907
Middle Devonian of New York State
Originally called Lepidodendron primaevum
A. serotina
Kräusel & Weyland, 1949
Originally called Archaeosigillariopsis serotina
A. sigillarioides
Lejal-Nicol, 1970
Late Devonian of Niger
A. subcostata
Danzé-Corsin, 1965
Mississippian of Niger
A. vanuxemii
Grierson & Banks, 1963
Leaves 5-6 mm long, simple, possibly petiolate, with a deltoid lamina that bears three or four opposite pairs of small serrations and tapers to an acute apex tipped by a tiny hair (Senkewich, 1956; Fairon-Demaret & Banks, 1978).
Stems are regarded as characterized by hexagonal patterns produced by subepidermal thick-walled cells that surround the areas of attachment of leaves
Gilboaphyton †
Gilboaphyton †
Arnold, 1937
G. argentinum †
Carrizo & Azcuy, 2006
Mississippian of Argentina
G. changyangense †
Feng, 1984
Late Devonian of China
G. goldringiae †
Arnold, 1937; emended Berry and Edwards, 1996
Middle Devonian of New York
Dichotomous branching.
Leaves spirally-arranged, their enlarged bases forming broadly fusiform to hexagonal patterns delimited by narrow grooves.
Persistent leaves are laminar, divided into a large medial segment and two substantially smaller and shorter distal-pointing opposite lateral segments about midway along the total leaf length.
Protostelic xylem column lobate in outline, exarch.
G. griersonii †
"no younger than Frasnian, and no older than late Eifelian"
Vegetative stems, 8-14.5 mm diameter, and up to at least 11 cm in length, bifurcating probably isotomously.
Stem surfaces divided into spirally arranged, adjoining, longitudinally elongate hexagonal or fusiform swollen leaf bases; 7-8 bases per horizontal turn, separated by narrow grooves, each bearing a persistent leaf just above its center.
Leaves slender, upright, up to 9 mm long and 1.20-1.49mm wide with broad base, divided approximately halfway along their length to give a narrow central segment and two much shorter lateral upwards-pointing segments, all tips acute.
Cortex differentiated into layers, Outer zone of stem tissues most resistant to decay, containing elongate regions of softer tissues seated within the leaf bases connected to a cylinder of similar tissue associated with the leaf trace.
Atasudendraceae
Atasudendron mirum †
Atasudendron mirum †
Senkevitsch et al., 1993
Givetian of Kazakhstan
Blasariaceae †
Barrandeina †
Barrandeina †
Stur, 1882
B. agadirica
Senkevitsch, 1961
Givetian of Kazakhstan
B. aroostookensis
Schopf, 1965
Middle Devonian of Maine
B. dusliana
Štúr, 1881
Givetian of Czech Republic
B. karatauensis
Senkevitsch, 1992
Eifelian of Kazakhstan
B. kolderupii
Kräusel & Weyland, 1934
Devonian
Originally called Psygmophyllum kolderupii
B. orlovii
Jurina, 1982
Givetian of Kazakhstan
B. pectinata
Høeg, 1931
Middle Devonian of Norway
B. perryana
White, 1905
Middle Devonian (?) of Maine
Originally called Anarthrocanna perryana
Barrandeinopsis †
Barrandeinopsis †
Kryshtofovich, 1955
B. antiqua
Radczenko, 1962
Devonian
Originally called Minusia antiqua
B. beliakovii
Kryshtofovich, 1955
Devonian of Russia
B. kirkinskayae
Ananiev, 1962
Middle Devonian of Russia
Blasaria sibirica †
Blasaria sibirica †
Zalessky, 1934
Middle Devonian of Russia
Broeggeria †
Broeggeria †
Nathorst, 1915
Emsian-Eifelian
B. laxa
Ananiev, 1960
Emsian of Russia
B. norvegica
Nathorst, 1915
Eifelian of Norway
B. strobiliformis
Dorf, 1933
Emsian of Wyoming
Cyclostigmataceae †
Cyclostigma †
Cyclostigma †
Haughton, 1859
C. aegypticum
Jongmans & Koopmans, 1940
Carboniferous of Egypt
C. asiaticum
Radczenko, 1957
Carboniferous of Russia
C. australe
Feistmantel, 1879
Carboniferous of Australia
C. brasiliense
Dolianiti, 1962
Visean of Brazil
C. cambricum
Crookall, 1964
Kasimovian of UK
C. dasyphyllum
Mägdefrau, 1936
Carboniferous of Germany
C. distans
Chachlov, 1940
Carboniferous of Russia
C. goeppertii
Doweld, 2017
Famennian of Poland
C. majus
Kidston, 1964
Bashkirian of UK
C. primorskiense
Zimina, 1989
Famennian of Russia
Knorria †
Knorria †
Sternberg, 1825
K. aciculari-acutifolia
Weiss, 1886
Late Devonian of Germany
K. acicularis
Göppert, 1846
Mississippian of Poland
K. acutifolia
Göppert, 1847
Mississippian of Poland
K. anceps
von Eichwald, 1854
Permian of Russia
K. apicalis
von Eichwald, 1854
Pennsylvanian of Ukraine
K. bailyana
Schimper, 1870
Famennian of Ireland
K. cancellata
von Eichwald, 1854
Pennsylvanian of Ukraine
K. cervicornis
Roemer, 1860
Carboniferous of Germany
K. chemungensis
Arnold, 1939
Devonian of Eastern North America
Originally called Sigillaria chemungensis
K. confluens
Göppert, 1850
Mississippian of Germany
K. cylindrica
Roemer, 1860
Carboniferous of Germany
K. dichotoma
Jukes & Salter, 1856
Originally called Sigillaria dichotoma
K. distans
Newberry, 1853
Pennsylvanian of Ohio
K. fusiformis
Roemer, 1850
Mississippian of Germany
K. goeppertii
Roemer, 1843
Mississippian of Germany
K. hildrethii
Newberry, 1853
Pennsylvanian of Ohio
K. imbricata
Sternberg, 1825
Originally called Lepidolepis imbricata
K. jugleri
Roemer, 1843
Mississippian of Germany
K. longifolia
Göppert, 1847
Mississippian of Poland
K. mammillaris
von Eichwald, 1854
Carboniferous of Russia
K. mariana
Michael, 1895
Late Triassic of Poland
K. megastigma
Roemer, 1843
Mississippian of Germany
K. microcostata
Zalessky, 1931
Famennian of Russia
K. polyphylla
Roemer, 1843
Mississippian of Germany
K. princeps
Göppert, 1852
Late Triassic of Poland
K. richteri
Geinitz, 1855
Pennsylvanian of Germany
K. schrammiana
Göppert, 1847
Mississippian of Poland
K. sellonii
Sternberg, 1825
Carboniferous of Germany
K. taxina
Lindley & Hutton, 1833
Moscovian of the UK
K. uralica
Neuburg, 1961
Originally called Knorriopsis uralica
K. verrucosa
Zalessky, 1905
Originally called Selaginites verrucosus
Jurinodendron †
Jurinodendron †
Doweld, 2001
This taxon may be a lepidodendrid
J. aegypticum
J. australe
J. brasiliense
J. brevifolium
J. cambricum
J. dasyphyllum
J. hercynium
J. macconochiei
J. majus
J. pacificum
J. primorskiense
J. sigillarioides
J. sinaicum
J. ungeri
J. ursinum
?
Eleutherophyllaceae
Eleutherophyllum †
Eleutherophyllum †
Stur, 1877
Zimmermannia †
Zimmermannia †
Gothan & Zimmermann, 1932
Lepidosigillariaceae
Lepidosigillaria †
Lepidosigillaria †
Kräusel & Weyland, 1949
Libyaria †
Libyaria †
Lejal-Nicol, 1975
Leptophloeaceae †
Omphalophloiaceae †
Protolepidodendraceae †
Leclercqia †
Leclercqia †
L. andrewsii †
L. complexa †
Banks, Bonamo and Grierson, 1972; Bonamo, Banks and Grierson, 1988
L. uncinata †
Axes branching isotomously or anisotomously at angles of 15–45
1.6–2.6 mm wide, at least 70 mm long
Leaf bases elongate-fusiform, tightly arranged in helices or pseudowhorls
8-10 leaves inserted on the axis per gyre
Leaf consists of a basal segment, dividing just below the midpoint into three major segments, a long narrow central segment, and two further divided opposite lateral segments arranged in three dimensions
Deflected abaxially, tapering distally and terminating in an abaxially tightly recurved hook
Pairs of lateral major segments are divided near the base into two or very often three more or less equal minor segments
Sporophyll similar in morphology to sterile leaf; no strobilus formed
Sporangium ovate to fusiform in lateral view, ovate in adaxial view, attached by an oval pad proximal to the division of the leaf,
Plant homosporous
Above: Reconstruction of Leclercqia complexa
Protolepidodendron †
Protolepidodendron †
There is debate whether Protolepidodendron is a valid genus. The Protolepidodendrales are plants that exhibit forked leaves. The holotype specimen of P. scharyanum (in Prague) is poorly preserved and possibly shows a forked leaf, but the apices of other leaves are missing.
P. scharyanum (Krejci 1880; Krausel & Weyland 1929; Jurina 2009)
Middle Devonian
The axis width is 3.1–3.3 mm and does not change over the entire specimen. Leaves with conical bases and narrowing
The length of fragmentary leaves is 2–3 mm, with a bifurcation approximately in its middle area (2 mm from the base) into two thin segments of apices about 2 mm long.
The apices are at an angle slightly exceeding 90° to each other.
Leaf scars are fusiform, acuminate at both extremities, about 2 mm long and 0.4–0.5 mm wide, are arranged in a very low spiral. There are three or, rarely, four leaf scars in each row.
Sublepidodendraceae
Artschaliphyton unicum †
Artschaliphyton unicum †
Senkevich 1971 (originally called Gilboaphyton goldringae Arnold)
Herbaceous plants with isotomous or pseudomonopodial branching
Leaves are petiolate, permanent, hastate, possibly toothed margin
Leaf scars are circular to oval
Sporangia are unknown
Cervicornus wenshanensis †
Cervicornus wenshanensis †
Pragian compression fossil from Yunnan China
Herbaceous axes with leaves arranged in openly spaced helices
The distal portion of each of the leaves is divided three times, resulting in eight segments
Anatomy is not preserved.
Above: Holotype specimen (Plate 1) of Cervicornus
Clwydia decussata †
Clwydia decussata †
Leaves (originally called Archaeosigillaria kidstoni) are 12 mm in length with a rhomboid lamina whose upturned distal portion tapers into a long hair (Lacey, 1962).
On younger axes the leaves are opposite decussate.
Artschaliphyton unicum †
Artschaliphyton unicum †
Senkevich, 1971 (originally called Gilboaphyton goldringae Arnold)
Herbaceous plants with isotomous or pseudomonopodial branching
Leaves are petiolate, permanent, hastate, possibly toothed margin
Leaf scars are circular to oval
Sporangia are unknown
Colpodexylon †
Colpodexylon †
Axes up to 2.5cm wide with isotomous branching
Lobed protostele with exarch maturation; no secondary xylem
Leaf cushion lacking scars or markings
Leaves are persistent and up to 3 cm long, and 3-forked
Sporangia adaxial and below the forking
C. deatsii †
Frasnian (Late Devonian)
C. gracilentum †
Dou et al., 1983; Xu & Wang, 2011
Middle Devonian of NW China
C. pullumpedes †
Famennian of Waterloo Farm, South Africa
‘Short-tipped’ species
High latitude taxon, probably living in refugia into the Late Devonian
C. mergae †
Famennian of Coombs Hill, South Africa
‘Long-tipped’ species similar to C. deatsii, but with different proportions
High latitude taxon, probably living in refugia into the Late Devonian
C. trifurcatum †
Eifelian (Middle Devonian)
Above: Suggested reconstruction of Colpodexylon deatsii
Below: Suggested reconstructions of Colpodexylon pullumpedes (left) and Colpodexylon mergae (right). Scale bar represents 10 mm (Fig. 3, Harris et al. 2021)
Estinnophyton (?) †
Estinnophyton (?) †
E. gracile †
E. wahnbachense †
Kräusel & Weyland, 1932; Fairon-Demaret, 1979
Originally described as Protolepidodendron wahnbachense from the Siegenian of Germany
Narrow axes with persistent leaves, which are bifurcated and spirally arranged.
The vegetative and fertile leaves bifurcate twice and the four resulting segments are spread in various planes
The fertile leaves support two pairs of stalked elongate sporangia which are directed towards the axis
E. yunnanense †
Lower Devonian (upper Pragian) Posongchong Formation at Zhichang village, Gumu Town, Wenshan District, Yunnan Province, China.
This plant possesses aerial stems with isotomous branching
The leaves are slender, once- or twice-bifurcated, and arranged in low spirals or pseudo-whorls
Fertile leaves are morphologically identical to the vegetative ones, clustered on the axis, forming a loose strobilus-like structure
There are three to six fertile leaves in each gyre
Two ovoid-elongate sporangia are singly attached to the adaxial surface each subtended by a recurved stalk, either at the base of each segment in a once-bifurcate fertile leaf or below the points of successive divisions in a twice-bifurcate fertile leaf
E. yunnanense differs from E. gracile and E. wahnbachense, mainly in possessing two single stalked sporangia on each fertile leaf rather than two pairs of sporangia, as well as in the dimensions of its leaves.
Estinnophyton is considered to have had a closer affinity with primitive sphenopsids than the protolepidodendralean lycopsids
Above: Line drawing of the fertile leaves of E. yunnanense
Haskinsia †
Haskinsia †
Middle Devonian
Grierson & Banks, 1963
Haskinsia was distributed in tropical, warm temperate and high southern latitude, cool (?) temperate zones (Cingolani et al., 2002)
Stems are narrow and isotomous
Leaves are petiolate and helically arranged; leaf lamina is roughly deltoid in shape
H. colophylla †
Grierson & Banks, 1983
Originally called Drepanophycus colophylla Grierson & Banks (1963)
H. hastata †
Berry & Edwards, 1996
H. hastata has hastate leaves and H. sagittata has sagittate leaves
H. sagittata †
Edwards & Benedetto, 1985
Above, left: Reconstruction of Haskinsia colophylla growing on a Lepidosigillaria stump (from Retallack 2018); Above, right: leaves of H. sagittata and H. hastata (Berry & Edwards 1996)
Zhenglia radiata †
Zhenglia radiata †
A small and slender herbaceous I ycopsid, possessing simple, entire, possibly ligulate microphylls, with decurrent, swollen bases
Rhomboid leaf bases, showing lepidodendroid-like phyllotaxic arrangement.
Leaf having a single vein.
Fertile structure consisting of closely arranged sporophylls.
Sporophylls basically identical with microphylls.
Each sporophyll bearing a single, ovoid-elongate sporangium attached adaxially.
Sporangia partly enclosed by lateral extensions of the sporophyll lamina.
Above: Zhenglia radiata † (Fig 2, Hao et al. 2006)
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