New species of Gleichenites CGleicheniaceae, Filicales) from the Upper Triassic of Argentina and Chile
Rafael Herbst
Herbst R 1996. New species of Cleichenites (GJeicheni3ce3e, Filic3les) from the Upper Tri3ssic of Argentim 3nd Chile. Palaeobotanist43(3) : 67-72.
Two new 3nd one undetermined species of Cleichenites 3re described from the Upper Tri3ssic C3n3don Largo Forrn31ion (El Tf3nquilo Group) from S3n13 Cruz Province, Argentin3, the UpperTri3ssic La Terner3 Form31ion, COpi3PO Province, northern Chile, and the Upper Tri3ssic Gomero-T3lc3mavid3 Member of the Concepcion 3re3 in southern Chile. These fmdings together with former ones of Argentin3, Austr31i3 3nd eventu3lly An13rctic3, reinforce the ide3 of 3 possible 3ustral origin of the f3mily Gleicheni3ceae, or at le3st p3n of it. All the Tri3ssic 3nd pan of the Jurassic hitherto known species from the Gondw3n3 re31m are multisorate 3nd thus belong to the primitive members of the f3mily included in the subgenus Diplopteryglum in Holnum's sense.
Key-words--P3beobo13ny, Filic3les, Cleichenites, Upper Tri3ssic, Argentim, Chile.
Rafael Herhst, Pringepa- Conlcet, Cas ilia de Correo 128, 3400 corriente,~Argentina.
RESUME
Se describen tres nueV3S form3s de Cleichenites: un3 del Tri3sico superior de b Form3cion C3n3don Largo (Group El Tranquilo) de la provinci3 S3nta Cruz, Argentina, otra del Triasico superior de 13Form3cion La Terner3, provinci3 COpi3PO, norte de Chile y un3 indeterminada del Tri3sico superior del Miembro Gomero-T3lc3m3vid3 del3re3 de Concepcion en el sur de Chile.
Ademas se brinda un3 sintesis de los hallazgos de este genera y ouos afines del Mesozoico bajo del Gondw3n3 (Tri3sico y .!urJsico de Argentin3 y Australi3) y en b3se 3 que c3si tod3s 13sespecies penencen 31primitivo genero Diplopterygium, se 3n3hz3 brevemente 13posibilid3d de un origen austral p3ra 13F3milia Gleichelli3ceae, 0 al menos de un3 p3rte de est3. ~
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~ ~ ~ ~ ~ ~ Cf~~, :i}jT~f<.1~ ., ciI ~ ~ ~~ ~ 3"~~Tf{f<.1~1~ ~ :i}jr2IchfC:ChI"B~ "B~ "B>rc::ffuf I ctl'hlf-lll>f11 'fiT it ¥IT ~ u ~~ ~~ 31~~ M 'lWT11\ m§3lT~I ~.~"B~"5f'ifTfCf'h1wlt Gl§"ll'"1t J'€lHl'lCfCf ~ ~ ~ ~ ~ ~3lT~nlc'-f IN an earlier paper, Herbst (972) described a new ancestor-Oligocarpia Goeppert which was known species of Gleichenites (G. potrerillensis) emphasiz from impressions of several species from the Car ing the importance of its finding in the Gondwana boniferous of North America and Asia, as well as realm. This was so, because at that time-and even spores from the same age, attributed to them (Abbott, now-a-days most authors supported a holartic origin 1954). for this (and many other) family of ferns. The main After that paper at least three species of argument here was the existence of a presumed Gleichenitesbased on fertile impressions were found 68 THE PALAEOBOTANIST in the Triassic of South America (herein described) plate 4, where the pinnules are clearly imbricated. It and one, also of Triassic age, from Australia (Herbst, has also not been stated in his specific diagnosis. 1974). Other two species were already known from Thus, the usefulness and the validity of this proposal the MiddleJurassic ofArgentina (Herbst, 1962; Herbst are somewhat doubtful. & Anzotegui, 1968); and one more, unfortunately This list of cited forms from the southern hemi sterile from the MiddleJurassic of northern Chile was sphere is relevant to a discussion about the origin of lately found (Herbst & Troncoso, in press). Further the Family Gleicheniaceae. It is noteworthy that more more, based on petrified specimens Millay and Taylor and more Triassic and Jurassic species are found in (990) described Antarctipteris from the Triassic of this region, while the worldwide expansion of the Antarctica, which was doubtfully placed with the genus is mostly from the Cretaceous Period. Triassic Gleicheniaceae. On the other hand, the genus Chan and Jurassic species are rare in the "north". sitheca, originally described from the Lower Permian of Shansi (China), which is considered by some The taxonomy of the Gleicheniaceae is still a authors (Bolkhovitina, 1968) as another possible an subject to debate; there are authors who accept only cestor -or in the line of-Gleichenites, was also found one genus Gleichenia (1-:Iolttum, 1957; Tryon & in South America, although with only one species, Tryon, 1982) with a subdivision in subgenera, to and as late as Upper Triassic (Herbst, 1963). those for who raise these subgenera to generic level In a recent paper Crabtree (988) proposed new (Naiko, 1950; Pichi Sermolli, 1972), the latter author generic name - Gleicheniaceaphyllum for sterile recognizes up to 8 genera. But all workers agree in fronds with a general "gleicheniaceous" morphology, considering Diplopterygium(no matter its taxonomic i.e., small, more or less pecopterid to cladophleboid rank) as a primitive genus in the family and somewhat pinnules, with typical open, fern venation. In his isolated from the rest. discussion he maintains that the genus Gleichenites All the Triassic and most of the Jurassic species Goeppert is invalid being based on a Palaeozoic from the Gondwana land-masses are multisorate, and foliage (a species of Sphenopteris) and also as thus belong to this DiplopterygiumPrimitive "Group". Goeppert's species are not the members of the family. But it is true that up to date no reliable possible This being true was recognized by Seward (926) ancestor from the Carboniferous or Permian has been who revalidated the name as clearly stated by Harris found in the "south" while in the northern hemi (931). Therefore most authors have used Gleichenia sphere some new forms are now and then proposed or Gleichenites ( the distinction of these two names as ancestors; a recent example might be the genus being part of another discussion) for fertile fronds Szea(Yao & Taylor, 1988). with a general resemblance with the fertile fossil and Many modern authors when discussing recent members of the family Gleicheniaceae; it is phylogenetic relations and lor the origin of groups of with this characterization that usage has imposed the plants (and animals as well) still somehow neglect the generic name without further problems. geographical data of the findings relaying more on It has to be stated that Crabtree (988) gave a evolutive trends or facts, and associate or relate or generic diagnosis which includes most of the mor ganisms from very different geographical regions. Of phological characters by which Gleichenites is course, it is always possible to invoque physical con known, but he also states that "pinnules, small alter nections or migration routes to overcome this incon nate, elliptic to deltoid to falcate, connected to one veniency but it is a fact, for this particular case, that another along rachis by thin strip of laminar tissue". from the Permian onwards and through most of the His type-species, G. falcatum does not show this Triassic andJurassic the Gondwana was almost com character.both in his text-figure 4 and figures 1-5 of pletely disconnected from the other land masses. PlATE 1 1,6 G. cachivarltensts; I, Sgo 131O( Ho!olype), x I; 6. CfES-PIl10129, 2,3. G. gallegoi: 2, CfES-PB 10120,1,3; 3, Sgo 1307, x 1,3. x 2. 4, S. Gleichenitessp; 4, CfES-pn 10132, x 3; S, CfES-PI3 10131, X 3. :::"'-l :c u Q Z -< z-< zf= "'-l ~ 0 V) "'-l t3 "'-l V)"- ~ "'-l Z ~ V) a:l C<: :c"'-l 70 THE PALAEOBOTANIST 700 GOo• 5'00 6I?:llrPi;zy\U Map 1 -Loc:llion m:lp of outcrops with Gteichenile,,~ (I) are;! of COw Terner;!. provo Copi:lp6. Chile; (2) :Ire;! ofGomero. pmv. Concepci6n. Chile, :lnd (3) ;!re:l of EI Tranquilo, provo S:lnl:l Cruz. Argentin:1 " o .,/ G Thus, the three species described in this paper are Text-figure 1- A, B :md C, Gteichenilespal/egoi, D, E, Gleicheniles significant in favour of the above exposed ideas, sp.; F, G, Gteichenile cachiuan'lensis ( B:lfS :Ire - I mm, except C which will have to be taken in account when the where =0.5 mm) origin of the Gleicheniaceae or eventually a part of this family represented by the Diplopterygiumgroup lower margin more curved than upper one, apex is established (Map n Naturally much more eviden rounded, Size typically 5-6 mm long x 3-4 mm wide. ces will be necessary for this purpose. Midvein strong, lateral ones simple and straight, 3-4 on each side, basal ones of each pinnule mostly once SYSTEMATIC DESCRIPTION forked. Fertile pinnules with six to seven sporangiate Family ---Gleicheniaceae sori, the pentasporangiate being the most common, Number of sori in basal half of pinnule is generally Genus -Gleichenites Seward 1926 (non Goeppert) higher than in upper half. Sori 0,7 to 1mm in diameter with 4 to 7 sporangia (generally 5-6). Sporangia more Gleichenites gallegoi n. sp, or less pyriform, with a short peduncle (7); size about 0.4 mm high x 0.25-0,28 mm in diameter; in some Text-figure 1 A, B, C; PI.1, figs 2-3 impressions thick-walled cells can be seen apically 1988 Gleichenitessp, in Herbst: 376, pLI, figs 7-10; but the disposition of the annulus is uncertain. pI. 4, figs 30-32, Holotype-CTES-PB no, 10120; Cotype no, 10121 Diagnosis-Fronds of unknown shape and size, (PLl, figs 2 and 3 respectively). at least bipinnate; main rachis 0,8-1 mm thick Largest Locality-Ea, Canadon Largo, Prov, Santa Cruz, segment tJp to 8-9 cm long x 10 cm wide, Pinnae Argentina. linear, up to 6 cm long x 7 mm wide, attached to the Honzon---Canadon Largo Formation, El Tran main rachis at 50°-60°, separated 6-8 mm. Pinnules quilo Group. cladophleboid, slighly falcate, united at the base; Age-Upper Triassic HERBST-NEW SPECIES OF GLEICHENlTES FROM ARGENTINA AND CHILE 71 Discussion-In 1988, I wrongly described the assic of Australia, but it distinctly differs in its soral sterile fragments of Gleichenites sp. as they were characters. • associated in the same slab with the fertile fragments Gleichenites sp. (Herbst, 1988). The abundant and well preserved specimens now available show that these belong to Text-figure 1 D-E; PI. 1, figs 4-5 Cladophlebissp. while those of the new species have quite different shape and venation. DeSCription-Fragments of a bipinnate frond of unknown size and shape. Segment rachis 1 mm wide. On the other hand I had already stated, based on the fertile specimens, that the species was not com Pinnae linear, narrowing towards the apex, 1-5 cm parable to any known form from the Triassic or long x 5-6 mm wide, attached to rachis at 40-45°, Jurassic of Australia, India and Argentina; this state separated 7-8 mm. Pinnules cladophleboid, united at the base, sterile ones 4 x 3 long/wide, fertile 3.5 ment stands. ones x 2.8-3 mm long/wide; lower margin slightly more Gleichenitesgallegoi is similar to Gleichenitessp. curved than upper one, apex rounded, more acute in from Gomero (Chile) described in this paper but sterile pinnules. Midvein marked, lateral ones simple differs in the form of pinnae, pinnules and venation and straight, 2-3 on each side. Occasionally one of (Text-figure 1 A-C). However, more material from the basal ones may fork once. Gomero would show whether they are conspecific. Fertile pinnules are more rounded with three to Gleichenites cachivaritensis n. sp. four sporangiate sori, 0.7-0.8 mm in diameter. Other details are not visible. Text-figure 1 F-G; Pl.l, figs 1, 6 Materiat---eTES-PB NO 10131, 10132. Diagnosis--Fronds bipinnate of unknown size Locality--400 m east of the railroad Gomero sta and shape; largest specimen 10 cm long x 12 cm wide. tion, area of Concepcion, Chile. Main rachis 1-1.2 mm thick. Largest pinnae 6 cm long Horizon-Talcamavida-Gomero Member (sensu x 4- 4.5 mm wide, linear, attached at 80-90° a'nd Tavera Jerez, 1960). separated 4-5 mm. Pinnules pecopterid to cladoph Age-Upper Triassic. leboid, united at the base, lower margin more curved Discussion--Only two fragmentary specimens than upper one, apex rounded to somewhat acute at (one fertile and other sterile) are in the collection, apical pinnae and pinnules. They are typically 2 x 2 therefore it is not possible to establish close com mm long/wide. Midvein strong, with 2-3 simple parisons but it has been mentioned above that this lateral veins at each side; occasionally one or two form is rather similar to G. gallegoi(this paper). Thus veins are once forked near the margin. it is being described as Gleichenites sp. Fertile pinnules 4-5 sporangiate (mostly 4 ACKNO~DGEMENTS sporangiate), rounded sori, 0,7 mm in diameter. Basal sori of pinnules usually bigger than upper ones. Thanks are due to the National Geographic Sporangia appear pyriform, sessile, with some of the Society for a Grant which permitted collecting most cells from the annulus visible at the lateral apical of the materials here described. Also to Mr Oscar region. Gallego (Pringepa-Conicet) and Dr Alejandro Tron Holotype-SGO-PB no 1310 (PI. 1, fig. 1). coso (University of Talca, Chile) for field assistance. Locality-Quebrada de La Cachivarita, area of Co La Ternera, provo Copiapo, lIlo, Region, Chile. REFERENCES Horizon--La Ternera Formation. Triassic. 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