Abstract

Faunipollenites Bharadwaj is considered a junior synonym of Protohaploxypinus Samoilovich emend. Morbey. However, Indian workers claim it is a valid genus due to a poorly defined corpus and absence of folds in distal attachment. In India, a standard method is applied including the oxidization with HNO₃ more than 48 hours (+10’ of KOH). We analyze the effects of that treatment on the morphology of pollen grains of both genera in samples from the Permian of India and Brazil. The same samples are also processed with HCl, HF, two hours of HNO₃ and 2’ of KOH and slides are mounted after each step. Our analysis reveals that distinct or indistinct central body and presence/absence of folds in distal attachment do not change in contrast to the indistinct central body and mostly absence of folds from samples that underwent a longer period of oxidization (24-48 hours and KOH 10’). The synonymization of Faunipollenites to Protohaploxypinus is confirmed. Species of Faunipollenites are reassigned to the revised species of Protohaploxypinus. The usage of the latter genus and its species in Permian biostratigraphic studies of India will improve Gondwanan correlations and paleobiogeographic reconstructions in future studies.

Key words
Faunipollenites; India; morphology; Permian; Protohaploxypinus; taxonomy

INTRODUCTION

Faunipollenites was erected by Bharadwaj (1962)BHARADWAJ DC. 1962. The miospore genera in the coals of Raniganj Stage (Upper Permian), India. The Palaeobotanist 9: 68-106. to include haploxylonoid striate bisaccate pollen grains bearing a central body with an ill defined outline and infrareticulate cappa, recovered from the sediments of the Permian Raniganj Formation of the East Raniganj Coalfield, Damodar Basin India (Fig. 1a). Its usage was questioned or rejected by many researchers (e.g. Hart 1964HART GF. 1964. A review of the classification and distribution of the Permian miospores: Disaccate Striatiti. Procceedings Compté Rendu 5° Congrès International de Stratigraphie et de Géologie du Carbonifère, Paris 1963 1: 1117-1129., Balme 1970BALME BE. 1970. Palynology of Permian and Triassic Strata in the Salt Range and Surghar Range, West Pakistan. In: Kummel B and Teichert C (Eds), Stratigraphic boundary problems: Permian and Triassic of West Pakistan. University of Kansas Special Publication 4: 305-453., Foster 1979FOSTER CB. 1979. Permian plant microfossils of the Blair Athol Coal Measures, Baralaba Coal Measures, and Basal Rewan Formation of Queensland. GSQ 372: 1-154.), except for Indian workers that are still using it. On the other hand, Protohaploxypinus defined by Samoilovich 1953SAMOILOVICH SR. 1953. Pollen and spores from the Permian deposits of the Cherdyn’ and Aktyubinsk areas, Cis-Urals. Trudy Vsesoiuznyi Nauchno-issledovatel’skii Geologo-razvedochnyi Institut, new series, Leningrad 75: 5-57. [in Russian]. was widely discussed by several authors especially Hart (1964)HART GF. 1964. A review of the classification and distribution of the Permian miospores: Disaccate Striatiti. Procceedings Compté Rendu 5° Congrès International de Stratigraphie et de Géologie du Carbonifère, Paris 1963 1: 1117-1129., who formally emended it to exclude non taeniate pollen grains. Balme (1970)BALME BE. 1970. Palynology of Permian and Triassic Strata in the Salt Range and Surghar Range, West Pakistan. In: Kummel B and Teichert C (Eds), Stratigraphic boundary problems: Permian and Triassic of West Pakistan. University of Kansas Special Publication 4: 305-453. proposed the combination of the type species Faunipollenites varius Bharadwaj 1962BHARADWAJ DC. 1962. The miospore genera in the coals of Raniganj Stage (Upper Permian), India. The Palaeobotanist 9: 68-106. to Protohaploxypinus and discarded Hart´s (1964) taxonomic proposal because of the morphological overlapping mainly created with Lunatisporites Leschik emend. Mädler that also displayed a haploxylonoid to slightly diploxylonoid with three-five taeniae. Hence, Morbey (1975)MORBEY SJ. 1975. The palynostratigraphy of the Rhaetian Stage, Upper Triassic in the Delbachgraben, Austria. Palaeöntographica Abt B 152: 1-75. proposed a new emendation of Protohaploxypinus to exclude beyond the non taeniate, the taeniate pollen grains with less than three taeniae in the cappa (Lunatisporites, Lueckisporites). He provided synonymy lists for the genus and several of its species. This amendment was followed by many researchers (e.g. Foster 1975FOSTER CB. 1975. Permian plant microfossils from the Blair Athol Coal Measures, Central Queensland, Australia. Palaeöntographica Abt B 154: 121-171., 1979, MacRae 1988MACRAE CS. 1988. Palynostratigraphical correlation between the Lower Karoo sequence of the Waterburg and Pafuri coal basins and the Hammanskraal plant macrofossil locality, RSA. Memoirs Geological Survey South Africa 75: 1-217., Utting 1994UTTING J. 1994. Palynostratigraphy of Permian and Lower Triassic rocks, Sverdrup Basin, Canadian Arctic Archipielago. GSC 478: 1-107., Stephenson 2015STEPHENSON MH. 2015. Bisaccate pollen from the early Permian OSPZ3a Sub-Biozone of the Lower Gharif Member, Oman. Rev Palaeobot Palynol 212: 214-225.) and among them, Foster (1979)FOSTER CB. 1979. Permian plant microfossils of the Blair Athol Coal Measures, Baralaba Coal Measures, and Basal Rewan Formation of Queensland. GSQ 372: 1-154. discussed the status of many species and provided long lists of synonyms. He discussed the morphological status of the type species Protohaploxypinus latissimus (Luber in Luber & WaltzLUBER AA & WALTZ IE. 1941. Atlas of microspores and pollen of the Palaeozoic of the USSR. Trudy Tsent. Nauchno-issled. Geologo-razv. Institute 139: 1-107. [in Russian, with English summary].) Samoilovich 1953SAMOILOVICH SR. 1953. Pollen and spores from the Permian deposits of the Cherdyn’ and Aktyubinsk areas, Cis-Urals. Trudy Vsesoiuznyi Nauchno-issledovatel’skii Geologo-razvedochnyi Institut, new series, Leningrad 75: 5-57. [in Russian]. although he did not find it in Australia. Instead, this species was described and illustrated by MacRae (1988, Pl. 25, figs. 13, 14) and Utting (1994, Pl. 6, figs. 27-29), based on its original description (i.e. haploxylonoid to weakly diploxylonoid amb, general oval shape in polar view, central body circular to slightly oval in transverse section, large saccus overlap resulting in a narrow cappula of less than 1/3 than the body width, ten taeniae entire and wedge shape). Foster (1979)FOSTER CB. 1979. Permian plant microfossils of the Blair Athol Coal Measures, Baralaba Coal Measures, and Basal Rewan Formation of Queensland. GSQ 372: 1-154. also discussed the taxonomic status of Faunipollenites and tentatively assigned it as a junior synonym of Protohaploxypinus Samoilovich 1953SAMOILOVICH SR. 1953. Pollen and spores from the Permian deposits of the Cherdyn’ and Aktyubinsk areas, Cis-Urals. Trudy Vsesoiuznyi Nauchno-issledovatel’skii Geologo-razvedochnyi Institut, new series, Leningrad 75: 5-57. [in Russian]. emend. Morbey 1975MORBEY SJ. 1975. The palynostratigraphy of the Rhaetian Stage, Upper Triassic in the Delbachgraben, Austria. Palaeöntographica Abt B 152: 1-75.. Indian workers (Venkatachala & Kar 1968VENKATACHALA BS & KAR RK. 1968. Palynology of the Karanpura Sedimentary Basin, Bihar, India-1. Barakar Stage at Badam. The Palaeobotanist 16(1): 56-90., Sinha 1972SINHA V. 1972. Sporae dispersae from Jhingurdah Seam, Singrauli Coalfield (MP) India. The Palaeobotanist 19(2): 175-201., Tiwari 1974TIWARI RS. 1974. Palynological succession in the Barakar type area. Geophytology 3: 166-183.), and especially Tiwari et al. (1989)TIWARI RS, SRIVASTAVA SC, TRIPATHI A & VIJAYA. 1989. Morphographic study of Permian palynomorphs: Callumispora, Parasaccites, Crucisaccites and Faunipollenites. The Palaeobotanist 37(2): 215-266., who treated Faunipollenites in detail, strongly rejected the taxonomic changes proposed for this genus and its type species, on the grounds of an ill defined central body outline. As this taxonomic controversy still requires a clarification, specimens attributed to Faunipollenites in palynological material from the Permian of India (Fig. 1a-d) and Protohaploxypinus in Brazil (Fig. 1e-g) are examined and compared using optical light microscope (Figs. 2-6), confocal laser scanning (Fig. 7) and Scanning electron (Fig. 8). This analysis aims to assess the taxonomic status of Faunipollenites to confirm its synonymization with Protohaploxypinus. Their presence in Permian assemblages of Gondwana and elsewhere will enhance future biostratigraphic correlations and paleobiogeographic reconstructions.

MATERIALS AND METHODS

Procedence of samples

The studied specimens were recorded from the Damodar and Godavari basins in India (Fig. 1a-d) and compared to the material from the Permian of Paraná Basin in Brazil (Fig. 1e-g). The former palynological materials come from the Godavari Basin, obtained from the subsurface levels of the Barakar Formation (early Permian) of Borehole 1007 in Manuguru area (Fig. 1c, further details in Kavali & Jha 2014KAVALI PS & JHA N. 2014. Upper Cisuralian palynology and palaeoclimate of Manuguru area Godavari basin, India and their global correlation. J Earth System Sci 123(7): 1681-1692.), and from the Damodar Basin, obtained from the Lopingian shale (Raniganj Formation) above Seam VI in Sonepur Bazari area of Raniganj Coalfield, near Asansol (Fig. 1d, further information in Jasper et al. 2012JASPER A, GUERRA-SOMMER M, UHL D, BERNARDES-DE-OLIVEIRA MEC, GHOSH AK, TEWARI R & SECCHI MI. 2012. Palaeobotanical evidence of wildfire in the Upper Permian of India: Macroscopic charcoal remains from the Raniganj Formation, Damodar Basin. The Palaeobotanist 61(1): 75-82.). Palynological materials from Brazil were obtained from surface samples of the Paraná Basin at two locations (Fig. 1e-g), one from early Permian of Rio Bonito Formation in southern Paraná State (unpublished) and the other from the Serra Alta and lower Rio de Rastro formations (Guadalupian) in Santa Catarina State (di Pasquo et al. 2018DI PASQUO MM, SOUZA PA, KAVALI PS & FÉLIX C. 2018. Seasonally warmer and humid climates in a lower paleolatitude position of southern Brazil (Paraná Basin): new findings of the Lueckisporites virkkiae zone (late Cisuralian-Guadalupian) in the Serra do Rio do Rastro and neighboring localities. J South Am Earth Sci 82: 143-164.).

Laboratory procedures and repositories

Samples were processed applying HCL (10%, 4 hours) and HF (40%, 18 hours), washed with distilled water, sieved through a 10 µm mesh and slides prepared with +10 residues using Entellan and cellosize products. The residues were divided in two sets and each one oxidized with HNO₃ (80%), one for two hours and the other 24-48 hours, KOH (10%) applied for 2 minutes and after sieving, last set of slides were mounted. These procedures were carried out at three laboratories where the samples (residues, slides, rocks) are housed: 1- India (Birbal Sahni Institute of Palaeosciences, acronym BSIP), 2- Brazil (Institute of Geosciences of the Universidade de São Paulo, acronym IGc, Universidade do Estado do Rio de Janeiro, acronym UERJ DEPA, Universidade Federal do Rio Grande do Sul, acronym MP-P), 3- Argentina (CICYTTP-CONICET-ER-UADER, acronym CICYTTP-Pl). The slides were analyzed using light microscopes Olympus BX61 microscope with DP-25 digital camera using Cell A software in India, Zeiss Scope.A1 camera and software Zen 2.3 lite in Brazil and Nikon Eclipse E200 with a videocamera Labomed (5.0 Mp) in Argentina. England Finder coordinates are used for illustrated specimens. Confocal Laser Scanning Microscope (CLSM) images were obtained at BSIP in India and the CENPES Institute of PETROBRAS in Brazil. Scanning electron microscopy (SEM) images were obtained at BSIP and the Electronic Microscopy Centre of the Institute of Geosciences, Universidade de São Paulo, Brazil.

RESULTS

Morphological comparison of Faunipollenites and Protohaploxypinus

The effect of oxidation on main diagnostic features of well- preserved (not altered and complete) specimens attributed to Faunipollenites from India (Figs. 2-5) and Protohaploxypinus from Brazil (Fig. 6) are compared using light microscope, CLSM (Fig. 7) and SEM (Fig. 8). These features are: (i) saccus condition, (ii) central body, (iii) folds, (iv) striae/ taeniae, and (v) size of specimens. This analysis aims to solve taxonomic misconceptions and discrepancies between those two genera that resulted after the oxidation of samples using HNO₃ not less than 24-48 hours as a standard method in India (P. Kavali, pers. comm.).

Our results detailed below, revealed that most of the morphologic features in the specimens analyzed after each treatment with HCL and HF and after two hours of HNO₃ and 2’ of KOH (Figs. 2-4) are maintained whereas morphological differences are noticed after the application of HNO₃ for 24-48 hours and 10-15’ of KOH (Fig. 5).

(i). Saccus construction: Based on the original and emended diagnoses and descriptions, both Protohaploxypinus (Samoilovich 1953SAMOILOVICH SR. 1953. Pollen and spores from the Permian deposits of the Cherdyn’ and Aktyubinsk areas, Cis-Urals. Trudy Vsesoiuznyi Nauchno-issledovatel’skii Geologo-razvedochnyi Institut, new series, Leningrad 75: 5-57. [in Russian]. and Morbey 1975MORBEY SJ. 1975. The palynostratigraphy of the Rhaetian Stage, Upper Triassic in the Delbachgraben, Austria. Palaeöntographica Abt B 152: 1-75.) and Faunipollenites (Bharadwaj 1962BHARADWAJ DC. 1962. The miospore genera in the coals of Raniganj Stage (Upper Permian), India. The Palaeobotanist 9: 68-106. and Tiwari et al. 1989TIWARI RS, SRIVASTAVA SC, TRIPATHI A & VIJAYA. 1989. Morphographic study of Permian palynomorphs: Callumispora, Parasaccites, Crucisaccites and Faunipollenites. The Palaeobotanist 37(2): 215-266.) are characterized by haploxylonoid to slightly diploxylonoid saccus construction. This feature did not suffer distortions after different processes applied and confirms it is of generic importance.

(ii). Central body: In the original diagnosis of Protohaploxypinus Samoilovich 1953SAMOILOVICH SR. 1953. Pollen and spores from the Permian deposits of the Cherdyn’ and Aktyubinsk areas, Cis-Urals. Trudy Vsesoiuznyi Nauchno-issledovatel’skii Geologo-razvedochnyi Institut, new series, Leningrad 75: 5-57. [in Russian]., the different features of central body were not mentioned, whereas in the emended diagnosis stated by Morbey (1975, a distinct or not well defined circular to oval central body is described having transversal (= vertical, t-a) or longitudinal (l-a) elongation. These variations depended on the species recognized, and among them some are characterized by indistinct to diffused central body (P. latissimus, P. varius, P. diagonalis), others possess a distinct central body (P. amplus, P. limpidus, P. rugosus, P. samoilovichi, P. jacobii, P. harti, P. haigii, P. microcorpus) and some others present transitional forms, from diffuse to well defined (P. goraiensis, P. perexiguus, P. rugatus). Thus, the nature of central body is of specific importance. In the emended diagnosis of Faunipollenites, Tiwari et al. (1989)TIWARI RS, SRIVASTAVA SC, TRIPATHI A & VIJAYA. 1989. Morphographic study of Permian palynomorphs: Callumispora, Parasaccites, Crucisaccites and Faunipollenites. The Palaeobotanist 37(2): 215-266. distinguished Faunipollenites from Protohaploxypinus because the central body is indistinct and the striations define the edge of the central body of the species of the Permian and Triassic of India. Contrarily, several species of Faunipollenites and specimens illustrated displayed distinct central bodies (e.g. Bharadwaj 1962BHARADWAJ DC. 1962. The miospore genera in the coals of Raniganj Stage (Upper Permian), India. The Palaeobotanist 9: 68-106., Pl. 17, Figs. 223, 224, 225 and Pl. 18, Figs. 230, 231, 233; Bharadwaj & Srivastava 1969BHARADWAJ DC & SRIVASTAVA SC. 1969. A Triassic microflora from India. Palaeöntographica Abt 125(4-6): 119-149., Pl. 26, Figs. 34, 35, 38; Tiwari et al. 1989TIWARI RS, SRIVASTAVA SC, TRIPATHI A & VIJAYA. 1989. Morphographic study of Permian palynomorphs: Callumispora, Parasaccites, Crucisaccites and Faunipollenites. The Palaeobotanist 37(2): 215-266., Pl. 26, Figs. 1, 2, 4, 5), including the type species of Faunipollenites (Bharadwaj 1962BHARADWAJ DC. 1962. The miospore genera in the coals of Raniganj Stage (Upper Permian), India. The Palaeobotanist 9: 68-106., Pl. 18, Fig. 230).

In our study almost all specimens analyzed from Brazil (Protohaploxypinus) and India (Faunipollenites) maintained their features of central body (amb, striation, cappula, folds) and sacci (Figs. 2-4, 6-8) after each treatment with HCl, HF, and 2 hours of HNO₃ and 2’ of KOH. Instead, attenuation or distortion of the features mainly concerning central body and folds occurred after a longer period of oxidation (more than 24-48 hours, Figs. 5 and 7). Therefore, we confirm that erroneous morphographic diagnoses and descriptions of species especially in India are likely due to their standard methodology. Therefore, the argument of indistinct central body and absence of folds claimed mainly by Tiwari et al. (1989)TIWARI RS, SRIVASTAVA SC, TRIPATHI A & VIJAYA. 1989. Morphographic study of Permian palynomorphs: Callumispora, Parasaccites, Crucisaccites and Faunipollenites. The Palaeobotanist 37(2): 215-266. to maintain Faunipollenites separated from Protohaploxypinus is rejected herein. It is notice that specimens with transitional features between species of both genera are observed in the studied samples irrespective the treatment applied.

(iii). Nature of cappula and folds associated to sacci attachment on distal face: The shape and width of the cappula are also important specific features of both genera equally developed depending on the nature and extent of saccus attachment to the central body (broad, narrow, convex, concave, straight). Folds are related to the attachment zones of sacci and central body in the cappula. This feature is common in many striate/non- striate bisaccate and monosaccate pollen grains and it is mainly used to differentiate species except for Cannanoropollis and Plicatipollenites (see discussion in Azcuy & di Pasquo 2000AZCUY CL & DI PASQUO MM. 2000. Palynology of the Late Carboniferous from the Tarija Basin, Argentina: a systematic review of monosaccate pollen genera. Palaeöntographica Abt B 253: 103-137. and references). The presence or absence of intexinal folds in Protohaploxypinus is not considered a generic feature (see Foster 1979FOSTER CB. 1979. Permian plant microfossils of the Blair Athol Coal Measures, Baralaba Coal Measures, and Basal Rewan Formation of Queensland. GSQ 372: 1-154.). Some species of Protohaploxypinus display very distinct zones of attachment (P. limpidus, P. amplus, P. rugosus, P. hartii, P. hagii), others are devoid of distal folds (P. varius, P. goraiensis, P. rugatus, P. diagonalis), whereas some are characterized by occasional folds (P. samoilovichi, P. microcorpus, P. jacobii, P. perexiguus). In the diagnosis of Faunipollenites, Bharadwaj (1962)BHARADWAJ DC. 1962. The miospore genera in the coals of Raniganj Stage (Upper Permian), India. The Palaeobotanist 9: 68-106. did not mention folds in the cb-sacci attachment used to maintain both genera separated (Tiwari et al. 1989TIWARI RS, SRIVASTAVA SC, TRIPATHI A & VIJAYA. 1989. Morphographic study of Permian palynomorphs: Callumispora, Parasaccites, Crucisaccites and Faunipollenites. The Palaeobotanist 37(2): 215-266.).

As discussed above, specimens of both taxa bearing folds before the stronger oxidation process reveal poorly defined folds or their absence. Therefore, the variability of this feature at the generic level is prevented and must be maintained as part of the specific characters.

(iv). Striations: Striations in the original definition of Protohaploxypinus and Faunipollenites show overlapping in number and disposition (simple, forked, cuneiform). Morbey (1975)MORBEY SJ. 1975. The palynostratigraphy of the Rhaetian Stage, Upper Triassic in the Delbachgraben, Austria. Palaeöntographica Abt B 152: 1-75. in his emended diagnosis of Protohaploxypinus stated a minimum number of five longitudinal taeniae or six striae to avoid the overlapping with the haploxylonoid Lunatisporites (3-4 taeniae separated by slightly wide striae) and the haplo-diploxylonoid Lueckisporites (two taeniae separated in the middle by a wider striae called platea by Vijaya 1992). The original diagnosis of Faunipollenites states 8-12 horizontal striations (Bharadwaj 1962BHARADWAJ DC. 1962. The miospore genera in the coals of Raniganj Stage (Upper Permian), India. The Palaeobotanist 9: 68-106.) and the emended diagnosis states 6-20 (Tiwari et al. 1989TIWARI RS, SRIVASTAVA SC, TRIPATHI A & VIJAYA. 1989. Morphographic study of Permian palynomorphs: Callumispora, Parasaccites, Crucisaccites and Faunipollenites. The Palaeobotanist 37(2): 215-266.). We observed that striations are slightly distorted after a longer period of oxidation, so this feature is not reliable to differentiate both genera. Hence, it is necessary to be careful to distinguished species when long oxidation processes are applied.

(v). Size: The size is an irrelevant feature to differentiate both genera (e.g. Lindström et al. 1997LINDSTRÖM S, MCLOUGHLIN S & DRINNAN AN. 1997. Intraspecific variation of taeniate bisaccate pollen within Permian glossopterid sporangia, from the Prince Charles Mountains, Antarctica. Int. J Plant Sci 158: 673-684.) because their type species Protohaploxypinus latissimus (major diameter of 83 µm) and Faunipollenites varius (MD 106 µm) are overlapped in their size ranges (i.e. 64 µm – 180 µm and 78-92 µm respectively). The size of specimens in general is increased in oxidized samples (see Smith & Butterworth 1967SMITH AHV & BUTHERWORTH MA. 1967. Miospores in the coal sequence of the Carboniferous of Great Britain. Palaeontological Association, Spec Pap Palaeontol 1: 1-324.).

Systematic palaeontology

A critical morphological analysis of specimens from Permian assemblages of India and Brazil was carried out. Their comparison with species of Protohaploxypinus and Faunipollenites described and illustrated in selected literature of India and elsewhere allowed us to support that Protohaploxypinus is the senior genus of Faunipollenites. The species of Faunipollenites as well as some of Protohaploxypinus are synonymized with the convalidated species of Protohaploxypinus revised herein (Fig. 9, Table I). Species addressed in this section are listed in alphabetical order.

Faunipollenites

F. bharadwajii Maheshwari, 1967

F. circumstriatus Maheshwari, 1969

F. congoensis (Bose & Kar) Tiwari, Srivastava, Tripathi & Vijaya, 1989

F. copiosus Bharadwaj & Salujha, 1965

F. enigmatus Maheshwari, 1969

F. gopadensis Bharadwaj & Srivastava, 1969

F. magnus (Bose & Kar) Tiwari, Srivastava, Tripathi & Vijaya, 1989

F. minor Salujha, 1965SALUJHA SK. 1965. Miospore assemblage of Seam IX of east Raniganj Coalfield. The Palaeobotanist 13(3): 227-238.

F. multistriatus Srivastava, 1979SRIVASTAVA AK. 1979. Studies in the Glossopteris flora of India 44. Raniganj plant megafossils and miospores from Auranga Coalfield, Bihar. The Palaeobotanist 26(1): 72-94.

F. parvus Tiwari, 1965TIWARI RS. 1965. Miospore assemblage in some coals of Barakar Stage (Lower Gondwana) of India. The Palaeobotanist 13(2): 168-214.

F. singrauliensis Sinha, 1972

Protohaploxypinus

P. amplus (Balme & Hennelly) Hart, 1964

P. bhardwajii Foster, 1979

P. chalonerii Clarke, 1965CLARKE RFA. 1965. British Permian saccate and monosulcate miospores. Palaeontology 8: 322-354.

P. goraiensis (Potonié & Lele) Hart, 1964

P. hartii Foster, 1979

P. jacobii (Jansonius) Hart, 1964

P. kumaonensis (Lankhapal, Sah & Dube) Hart, 1964

P. limpidus (Balme & Hennelly) Balme & Playford, 1967

P. microcorpus (Schaarschmidt) Clarke, 1965

P. micros Hart, 1964

P. panaki Utting, 1994

P. perfectus (Naumova) Samoilovich, 1953

P. rugosus (Jansonius) Foster, 1979

P. samoilovichii (Jansonius) Hart, 1964

P. sewardii (Virkki) Hart, 1964

P. varius (Bharadwaj) Balme, 1970

Infraturma STRIATITI Pant, 1954

Genus PROTOHAPLOXYPINUS Samoilovich emend. Morbey, 1975

TYPE SPECIES. Protohaploxypinus latissimus (Luber in Luber & Waltz) Samoilovich, 1953 [SD].

REMARKS. We agree with Foster (1979, see also Foster & Gomankov 1994FOSTER CB & GOMANKOV AV. 1994. A new structure in pollen assigned to Striatopodocarpites Sedova 1956 and Protohaploxypinus Samoilovich emend. Morbey 1975, from the Late Permian (Tatarian) of the Russian Platform. J Australian Geol Geophys 15(2): 235-238.) in the constrainment of Protohaploxypinus versus Striatoabieites (most common Striatoabieites multistriatus (Balme & Hennelly) Hart 1964HART GF. 1964. A review of the classification and distribution of the Permian miospores: Disaccate Striatiti. Procceedings Compté Rendu 5° Congrès International de Stratigraphie et de Géologie du Carbonifère, Paris 1963 1: 1117-1129.) and Striatopodocarpites. Detailed descriptions and synonymy of genus and species treated in this contribution are mainly addressed by Balme (1970)BALME BE. 1970. Palynology of Permian and Triassic Strata in the Salt Range and Surghar Range, West Pakistan. In: Kummel B and Teichert C (Eds), Stratigraphic boundary problems: Permian and Triassic of West Pakistan. University of Kansas Special Publication 4: 305-453., Morbey (1975)MORBEY SJ. 1975. The palynostratigraphy of the Rhaetian Stage, Upper Triassic in the Delbachgraben, Austria. Palaeöntographica Abt B 152: 1-75., Foster (1979)FOSTER CB. 1979. Permian plant microfossils of the Blair Athol Coal Measures, Baralaba Coal Measures, and Basal Rewan Formation of Queensland. GSQ 372: 1-154., McRae (1988) and Utting (1994)UTTING J. 1994. Palynostratigraphy of Permian and Lower Triassic rocks, Sverdrup Basin, Canadian Arctic Archipielago. GSC 478: 1-107..

Protohaploxypinus bharadwajii Foster, 1979

HOLOTYPE. Foster (1979)FOSTER CB. 1979. Permian plant microfossils of the Blair Athol Coal Measures, Baralaba Coal Measures, and Basal Rewan Formation of Queensland. GSQ 372: 1-154., Pl. 29, fig. 10.

1965 P. microcorpus (Schaarschmidt) Clarke, p. 338, Pl. 41, Fig. 3.

DIAGNOSTIC CHARACTERS. Haplo- slight diploxylonoid subcircular to longitudinally oval amb, narrow lateral ridges somewhat present, well- defined vertical narrow corpus, cappa with parallel to wedge –shape striae in number of ca. 5 to 11, cappula lensoid to rectangular relatively broad, around 1/3 of the central body radius and folds associated to the distal roots. Total diameter 55-105 µm (from the diagnosis).

COMPARISON. We accept the combination of P. microcorpus proposed by Clarke based on his description. Clarke´s specimen illustrated in the plate 41.3 is transferred to P. bharadwajii Foster based on narrower t-a central body and striae lesser than 10-11. Although, both species are herein considered morphologically transitional and possibly extremes of a mopho-group (see Fig. 9, Table I).

Protohaploxypinus diagonalis Balme, 1970

For description see Balme (p. 364-365, Pl. 10, figs. 6-8).

Protohaploxypinus haigii Foster, 1979

REMARKS. This species is distinguished from others of the genus mostly due to its broad central body and cappula in horizontal direction (see description and comparisons in Foster 1979FOSTER CB. 1979. Permian plant microfossils of the Blair Athol Coal Measures, Baralaba Coal Measures, and Basal Rewan Formation of Queensland. GSQ 372: 1-154.).

Protohaploxypinus hartii Foster, 1979

Figures 2.2, 3.2, 4.4, 5.2, 5.4, 5.6, 7.2

HOLOTYPE. Foster 1979FOSTER CB. 1979. Permian plant microfossils of the Blair Athol Coal Measures, Baralaba Coal Measures, and Basal Rewan Formation of Queensland. GSQ 372: 1-154., Pl. 30, fig. 2.

1965 P. chaloneri Clarke, Pl. 42, Figs. 3 and 5.

REMARKS. This species bears a barely rectangular, haplo to slightly diploxylonoid amb, corpus vertically oval in shape, cappula broad rectangular and cappa with 5-8 striae, parallel to wedge- shape.

COMPARISON. This species is distinguished from P. limpidus due to the corpus shape and frequent wedge striae beyond parallel ones, although we consider both taxa as transitional and possibly part of a morpho-group addressed below. Main features of specimens illustrated as P. chalonerii Clarke agree with the morphology of P. hartii (see above) and they are junior synonyms.

Protohaploxypinus latissimus (Luber in Luber & Waltz) Samoilovich, 1953

Figures 2.6, 3.2, 4.9-10, 5.1, 5.5, 6.5, 6.6, 6.10, 7.2, 8.4, 8.5

BASIONYM. Pemphygaletes latissimus Luber in Luber & Waltz, 1941, Pl. 13, fig. 221 (see Samoilovich 1953SAMOILOVICH SR. 1953. Pollen and spores from the Permian deposits of the Cherdyn’ and Aktyubinsk areas, Cis-Urals. Trudy Vsesoiuznyi Nauchno-issledovatel’skii Geologo-razvedochnyi Institut, new series, Leningrad 75: 5-57. [in Russian]., Pl. 4, fig. 4).

1965 Protohaploxypinus chaloneri Clarke, p. 337, Pl. 42, Fig. 4 (holotype).

1967 Faunipollenites varius Bharadwaj (auct. non); Maheshwari, p. 275, Pl. 8, Fig. 62.

1970 Protohaploxypinus varius (Bharadwaj) Balme, p. 365, Pl. 10, figs. 4, 5.

2010 F. varius; Murthy et al., p. 707, Pl. 1, fig. 15.

2018 P. varius; di Pasquo et al., p. 154, fig. 11.D, p. 155, fig. 12.N.

2018 Protohaploxypinus sp. cf. goraiensis; di Pasquo et al., p. 156, fig. 13.B.

DIAGNOSTIC CHARACTERS. Haploxylonoid to slightly diploxylonoid, amb oval to subrectangular. Cappa bearing less than 12 striae, generally 8-12, subparallel and occasionally branched/wedged. Central body subcircular to slightly oval in transverse section barely distinct. Cappula rectangular ½ to 1/3. Lateral ridges somewhat present (see description in Utting 1994UTTING J. 1994. Palynostratigraphy of Permian and Lower Triassic rocks, Sverdrup Basin, Canadian Arctic Archipielago. GSC 478: 1-107.).

COMPARISON. This species bears morphological characters like a broad rectangular cappula, somewhat lateral ridges and subparallel to wedge – shape striae being transitional to P. limpidus and P. perfectus. The diagnostic features of P. chaloneri and its holotype illustrated by Clarke (1965, Pl. 42, Fig. 4) agree with the morphological description of P. latissimus and support our reassignment to this species. The same argument is applied to the specimens illustrated as P. varius by Balme (1970)BALME BE. 1970. Palynology of Permian and Triassic Strata in the Salt Range and Surghar Range, West Pakistan. In: Kummel B and Teichert C (Eds), Stratigraphic boundary problems: Permian and Triassic of West Pakistan. University of Kansas Special Publication 4: 305-453. and Protohaploxypinus sp. cf. P. goraiensis by di Pasquo et al. (2018, transitional to P. haigii). Hence, they are transferred to P. latissimus. Two specimens illustrated as P. latissimus by MacRae (1988, Pl. 25, figs. 13, 14) that are confirmed herein.

Protohaploxypinus limpidus (Balme & Hennelly) Balme & Playford, 1967

Figures 2.1, 3.5, 4.3, 5.9, 5.10, 5.15, 6.2, 6.7, 8.6

BASIONYM. Lueckisporites limpidus Balme & Hennelly, pars., p. 94, Pl. 3, fig. 29 (designated by Balme & Playford 1967BALME BE & PLAYFORD G. 1967. Late Permian plant microfossils from the Prince Charles Mountains, Antarctica. Rev Micropaléont 10(3): 179-192.).

1962 Faunipollenites varius Bharadwaj, p. 95, Pl. 18, fig. 230.

1964 Protohaploxypinus micros Hart, p. 29, text-fig. 60.

1965 P. jacobii Jansonius (auct. non); Clarke, p. 337, Pl. 41, fig. 1.

1972 Faunipollenites singrauliensis Sinha, p. 195, Pl.7, fig. 105.

1979 Protohaploxypinus jacobii Jansonius; Foster, p. 89, Pl. 30, Fig. 12.

1988 Protohaploxypinus suchonensis (Sedova) Hart (auct. non); MacRae, p. 63, Pl. 25, fig. 16.

(Auct. non) 2018 P. limpidus; di Pasquo et al., p. 155, Fig. 15.M.

See Balme (1970)BALME BE. 1970. Palynology of Permian and Triassic Strata in the Salt Range and Surghar Range, West Pakistan. In: Kummel B and Teichert C (Eds), Stratigraphic boundary problems: Permian and Triassic of West Pakistan. University of Kansas Special Publication 4: 305-453., Foster (1979)FOSTER CB. 1979. Permian plant microfossils of the Blair Athol Coal Measures, Baralaba Coal Measures, and Basal Rewan Formation of Queensland. GSQ 372: 1-154. and MacRae (1988)MACRAE CS. 1988. Palynostratigraphical correlation between the Lower Karoo sequence of the Waterburg and Pafuri coal basins and the Hammanskraal plant macrofossil locality, RSA. Memoirs Geological Survey South Africa 75: 1-217. for additional synonymy.

DIAGNOSTIC CHARACTERS. Bilaterally oval to subrectangular haploxylonoid amb, central body outline distinct ± subcircular to slightly oval in horizontal-vertical shape, cappa with more than 4 to 8 striae parallel, distal saccus attachment ill-defined occasionally with folds and a broad cappula bearing somewhat a median, vertical slit or groove can be present. Holotype measure 95 x 67 µm.

ORIGINAL DIAGNOSIS OF F. singrauliensis. Horizontally oval, bilateral pollen grains. Size 64 x 50 µm-100 x 70 µm. Central body ill defined, proximally bearing well defined horizontal striations, 5-10 in number. Sacci haploxylonoid, zone of distal saccus attachment diffused, a median vertical groove or slit present.

COMPARISON. The holotypes of F. varius (type species) and F. singrauliensis are synonymized with P. limpidus based on equal number of (5-10) striations, narrow cappula and occasional folds along the cappula. Ill- defined nature of central body of the holotype is not taken into consideration herein since the nature of central body is doubtful due to the treatment with HNO₃ applied to the Indian materials as it was explained. Size is also discarded to separate species of Protohaploxypinus. The specimens illustrated as P. jacobii by Foster (1979, Pl. 30, Fig. 12), as P. jacobii by Clarke (1965, Pl. 41 Fig. 1), and P. suchonensis (Sedova) Hart by MacRae (p. 63, Pl. 25, fig. 16) are relocated into P. limpidus based on sharing the same diagnostic features with this taxon described above.

Protohaploxypinus microcorpus (Schaarschmidt) Clarke, 1965

Figures 3.6, 6.3, 8.2

BASIONYM. Striatites microcorpus Schaarschmidt (p. 55, Pl. 14, figs. 6-7)

1979 Faunipollenites multistriatus Srivastava; p. 86, Pl. 3, Fig. 16.

For other synonymies see Foster (1979)FOSTER CB. 1979. Permian plant microfossils of the Blair Athol Coal Measures, Baralaba Coal Measures, and Basal Rewan Formation of Queensland. GSQ 372: 1-154., di Pasquo et al. (2018)DI PASQUO MM, SOUZA PA, KAVALI PS & FÉLIX C. 2018. Seasonally warmer and humid climates in a lower paleolatitude position of southern Brazil (Paraná Basin): new findings of the Lueckisporites virkkiae zone (late Cisuralian-Guadalupian) in the Serra do Rio do Rastro and neighboring localities. J South Am Earth Sci 82: 143-164..

DIAGNOSTIC CHARACTERS. Amb haplo- to slightly diploxylonoid, subrectangular to oval in shape, narrow lateral ridges occasionally present, corpus vertically oval, cappula rectangular to lensoid, variably broad, distal attachment usually with folds, cappa with more than 10 striae mostly parallel and lesser in wedge- shaped.

ORIGINAL DIAGNOSIS OF F. multistriatus. Size range 104-120 µm x 70-80 µm; central body obscure, circular to horizontally oval, 58-64 x 60-68 µm; horizontal striations 12-18 µm, branched, sulcus 20-24 µm wide.

ORIGINAL DESCRIPTION. Bilateral, disaccate diploxylonoid; body thin, intra-microreticulate; sacci hemispherical, attachment full, straight to convex, intra-reticulation fine.

REMARKS. This species is haplo to weakly diploxylonoid oval amb, central body indistinct, cappa with 10 to 20 taeniae commonly branched, discontinuous or wedge-shaped, sacci cappula ½-1/3 rectangular to lensoid. Total diameter 92-121 µm (after Foster, 1979).

COMPARISON. F. multistriatus is synonymized with P. microcorpus based on similarities in size, the number of striations (12-18) and lensoid cappula. Our morphological description and illustrations of P. microcorpus agrees with the one provided by Foster (1979, p. 91). Striatites richterii Jansonius (1962, Pl. 14, figs. 21-22) and Striatopodocarpites magnicorpus Bharadwaj & Tiwari (1964)BHARADWAJ DC & TIWARI RS. 1964. On two monosaccate genera from Barakar Stage of India. The Palaeobotanist 12: 139-146. illustrated by Murthy et al. (2010)MURTHY S, CHAKRABORTI B & ROY MD. 2010. Palynodating of subsurface sediments, Raniganj Coalfield, Damodar Basin, West Bengal. J Earth System Sci 119: 701-710., are quite diploxylonoid differing from the haplo- to slightly diploxylonoid amb of P. microcorpus. Hence, a reassignment to Striatopodocarpites is proposed. P. microcorpus is considered transitional to P. bharadwajii and P. pennatulus, and also, to Striatoabieites multistriatus in the case of specimens having a horizontally oval corpus and smaller sacci (see also P. sewardi after Hart 1964HART GF. 1964. A review of the classification and distribution of the Permian miospores: Disaccate Striatiti. Procceedings Compté Rendu 5° Congrès International de Stratigraphie et de Géologie du Carbonifère, Paris 1963 1: 1117-1129.).

Protohaploxypinus pennatulus (Andreyeva) Hart, 1964

Figures 2.7, 2.9, 3.3, 4.1, 4.2

BASIONYM. Coniferaletes pennatulus Andreyeva et al. (Lectotype selected by Hart 1964HART GF. 1964. A review of the classification and distribution of the Permian miospores: Disaccate Striatiti. Procceedings Compté Rendu 5° Congrès International de Stratigraphie et de Géologie du Carbonifère, Paris 1963 1: 1117-1129.).

1961 Lunatisporites goraiensis Potonié & LelePOTONIÉ R & LELE KM. 1961. Studies in the Talchir flora of India-Sporae Dispersae from the Talchir beds of South Rewa Gondwana basin. The Palaeobotanist 8: 22-37., p. 32, Pl. 3, figs. 70-72.

1964 P. kumaonensis (Lankhapal, Sah & Dube) Hart, p. 29, text-fig. 62.

1965 Faunipollenites copiosus Bharadwaj & Salujha, p. 36-37, Pl. 2, fig. 40.

1967 Faunipollenites bharadwajii Maheshwari, p. 275, Pl. 8, fig. 63 [auct. non P. bharadwajii Foster 1979FOSTER CB. 1979. Permian plant microfossils of the Blair Athol Coal Measures, Baralaba Coal Measures, and Basal Rewan Formation of Queensland. GSQ 372: 1-154.].

1988 P. goraiensis; MacRae, Pl. 25, figs. 17, 19, 25.

1991 P. amplus; Backhouse, Pl. 17, fig. 11.

DIAGNOSTIC CHARACTERS. Amb horizontal (transversely) oval to subcircular, haplo to slightly diploxylonoid, with narrow lateral ridges, little discernible central body of subcircular to slightly longitudinal oval form. Cappa with more than 8 well- marked taenia, parallel to occasionally wedge. Relatively narrow capula (1/5-1/10) with rectangular to irregular shape enlarging at the lateral ends (i.e. narrower in the central part, Table I).

ORIGINAL DIAGNOSIS OF F. copiosus. Central body vertically oval, outline hardly discernible, exine intra-microreticulate, horizontal striations 9-11, coarsely intra-reticulate bladders, 25-32 µm apart distally. Size 148-162 µm in width (Bharadwaj & Salujha 1965BHARADWAJ DC & SALUJHA SK. 1965. A sporological study of Seam VII (Jote Dhemo Colliery) in the Raniganj Coalfield, Bihar (India). The Palaeobotanist 13: 30-41., p. 36-37).

ORIGINAL DESCRIPTION. Bisaccate, bilateral, golden yellow pollen grains; holotype 162 µm. Central body vertically oval, 70-76 µm x 88-98 µm, slightly bigger in height than the bladders, outline hardly discernible, striation ends distinct. Exine intra-microreticulate, proximally bearing 9-11 horizontal striations without any vertical partitions. Bladders subspherical, coarsely intrareticulate attaching distally and leaving a 25-32 µm wide thin bladder free area (Bharadwaj & Salujha 1965BHARADWAJ DC & SALUJHA SK. 1965. A sporological study of Seam VII (Jote Dhemo Colliery) in the Raniganj Coalfield, Bihar (India). The Palaeobotanist 13: 30-41., p. 36-37).

ORIGINAL DIAGNOSIS OF F. bharadwajii. Pollen grains disaccate, bilateral, haploxylonoid, 152-180 µm long, holotype 179 μm. Central body ill-defined, proximally bearing 7-10 simple or forked striations, exine in between the striations intra-microreticulate. Sacci hemispherical, distally inclined leaving a narrow, ill-defined distal sulcus; saccus intra-reticulation coarse.

COMPARISON. Both Protohaploxypinus pennatulus and the holotype of Faunipollenites copiosus are characterized by haploxylonoid to slightly diploxylonoid saccus construction, circular to oval central body (oval in F. copiosus) with a slight transverse or longitudinal elongation with 8-12 striations and a thin straight narrow cappula. F. bharadwajii and Lunatisporites goraiensis are reassigned to P. pennatulus because of their similarity especially based on the number of striations (7-10 in F. bharadwajii) and narrow cappula. It is interesting to note that Foster (1979, page 87, pl. 29, figs. 6-10) erected a new species of Protohaploxypinus with the same epithet “bharadwajii”, but he did not mention that it was used by Maheshwari (1967)MAHESHWARI HK. 1967. Studies in the Glossopteris Flora of India - 29. Miospore assemblage from the Lower Gondwana exposures along Bansloi River in Rajmahal Hills, Bihar. The Palaeobotanist 15(3): 258-280. for a new species of Faunipollenites. The synonymization of F. bharadwajii to Protohaploxypinus pennatulus allows us to retain Foster´s epithet. Interestingly, Playford & Dino (2000)PLAYFORD G & DINO R. 2000. Palynostratigraphy of upper Palaeozoic strata (Tapajós Group), Amazonas Basin, Brazil: Part Two. Palaeöntographica Abt B 255: 87-145. among others that described P. bharadwajii Foster, did not notice this situation since some Permian papers published by Indian researchers were ignored due to inaccessibility or not considered by researchers outside India.

Protohaploxypinus perexiguus (Bharadwaj & Salujha) Foster, 1979

Figures 3.4, 4.8, 5.14, 6.4, 6.9, 6.12

BASIONYM. F. perexiguus Bharadwaj & Salujha, 1965.

See Foster (1979)FOSTER CB. 1979. Permian plant microfossils of the Blair Athol Coal Measures, Baralaba Coal Measures, and Basal Rewan Formation of Queensland. GSQ 372: 1-154. for additional synonymy.

1965 Faunipollenites parvus Tiwari, p. 200, Pl. 7, Fig. 158.

1965 Faunipollenites minor Salujha, p. 232, Pl. 2, Fig. 30.

1969 Protohaploxypinus rugatus SegrovesSEGROVES KL. 1969. Saccate plant microfossils from the Permian of Western Australia. Grana Palynologica 9: 174-227., p. 200-201, Pl. 7, Figs. G-H.

1979 P. rugatus Jansonius; Foster, p. 92, Pl. 33, figs. 5-6.

1988 P. limpidus; MacRae, p. 61, Pl. 25, fig. 7.

1988 P. goraiensis; MacRae, p. 64, Pl. 25, fig. 24.

1988 P. microcorpus; MacRae, p. 64, Pl. 25, figs. 21-22.

1989 Faunipollenites congoensis (Bose & Kar) Tiwari, Srivastava, Tripathi & Vijaya, p. 259, Text Fig. 9.

1989 Faunipollenites magnus (Bose & Kar) Tiwari, Srivastava, Tripathi & Vijaya, p. 259; Text Fig. 10.

1991 P. rugatus; Backhouse, p. 289, Pl. 17, figs. 1-5.

1994 Protohaploxypinus panaki Utting, p. 55, Pl. 6, figs. 30, 31, Pl. 7, figs. 1-3.

2010 P. singrauliensis; Murthy et al., p. 707, Pl. 1, fig. 14.

2012 P. rugatus; di Pasquo & Grader, Pl. 7, figs. 3, 5.

DIAGNOSTIC CHARACTERS. Amb haploxylonoid, subcircular to vertically oval, central body indistinct, circular- subcircular to vertically oval. Cappa with taenia not well- defined, more or less parallel to wedge/branching in shape into central part, sensu stricto than 8 in number and more distinct in the central part of the body in general. Cappula narrow (mostly 1/10), rectangular to weakly widening at the lateral ends, without distal folds or occasionally narrow folds. Sacci semicircular scarcely developed in longitudinal width, without lateral ridges. Total diameter 29-40 µm to 110 µm (Foster 1979FOSTER CB. 1979. Permian plant microfossils of the Blair Athol Coal Measures, Baralaba Coal Measures, and Basal Rewan Formation of Queensland. GSQ 372: 1-154.).

ORIGINAL DIAGNOSIS OF F. parvus. Small, bilaterally oval pollen grains, known size 53-70 µm x 33-51 µm. Central body small, thin, obscure, apparently subcircular; known number of horizontal striations 5-7, without vertical partitions; distal channel narrow poorly defined, 8-9 µm broad when slightly apparent; sacci ± hemispherical finely intrareticulate.

ORIGINAL DESCRIPTION. Holotype 60 x 37 µm. Pollen grains small bilaterally oval; central body almost ill-defined, apparently subcircular thin, finely intra-microreticulate. Horizontal striations unbranched without any vertical partitions. Distally, saccus free area thin, narrow and uniformly broad, only in few cases measurable, 8-9 µm broad. Sacci small, ± hemispherical, finely intrareticulate.

ORIGINAL DIAGNOSIS OF F. minor. Overall size 36-62 µm. Central body indistinct, exine intra-microreticulate; distal sulcus 8-12 µm broad, bladder intra-reticulation small sized.

ORIGINAL DESCRIPTION. Yellow, bisaccate, bilateral pollen grains, holotype 46 µm. Central body vertically oval, outline indistinct made out by striation ends; exine intra-microreticulate, proximally bearing 5-7 horizontal striations without any vertical partitions in between them. Bladders semi-spherical with small sized meshwork, attaching distally leaving 8-12 µm wide bladder free thin area.

COMPARISONS. The original diagnosis of the holotype of F. parvus states it is characterized by an obscure central body. However, the holotype does not support this view. Foster (1979)FOSTER CB. 1979. Permian plant microfossils of the Blair Athol Coal Measures, Baralaba Coal Measures, and Basal Rewan Formation of Queensland. GSQ 372: 1-154. considered F. parvus and F. minor as junior synonyms of P. limpidus and included small, distinctly haploxylonoid end members of P. limpidus and P. perexiguus under P. rugatus. He said the latter differs only on their limboid sacci margins and narrow cappulae slit like, rectangular or slightly expanded at lateral extremities. However, F. parvus and F. minor are closely similar to P. rugatus even in their small size. Tiwari et al. (1989)TIWARI RS, SRIVASTAVA SC, TRIPATHI A & VIJAYA. 1989. Morphographic study of Permian palynomorphs: Callumispora, Parasaccites, Crucisaccites and Faunipollenites. The Palaeobotanist 37(2): 215-266. synonymized F. parvus with F. perexiguus, whilst Millstead (1999) considered them as junior synonyms of P. limpidus. Balarino (2012)BALARINO ML. 2012. Palinología del Pérmico de la Cuenca Claromecó-Colorado, Argentina. Ameghiniana 49: 343-364. considered P. perexiguus and P. rugatus as junior synonyms of P. goraiensis also agreeing with the criteria of size stated by Lindström et al. (1997)LINDSTRÖM S, MCLOUGHLIN S & DRINNAN AN. 1997. Intraspecific variation of taeniate bisaccate pollen within Permian glossopterid sporangia, from the Prince Charles Mountains, Antarctica. Int. J Plant Sci 158: 673-684.. MacRae (1988)MACRAE CS. 1988. Palynostratigraphical correlation between the Lower Karoo sequence of the Waterburg and Pafuri coal basins and the Hammanskraal plant macrofossil locality, RSA. Memoirs Geological Survey South Africa 75: 1-217. proposed to maintain P. goraiensis separated from P. pennatulus because of a slight difference in the number of striae and size, which are here considered not enough arguments in agreement with Foster (1979)FOSTER CB. 1979. Permian plant microfossils of the Blair Athol Coal Measures, Baralaba Coal Measures, and Basal Rewan Formation of Queensland. GSQ 372: 1-154.. In this work we accept the validity of P. pennatulus as senior synonym of P. goraiensis (holotype) preventing its usage as a valid species (with priority), and we consider the holotypes of Faunipollenites parvus, F. minor, Protohaploxypinus rugatus and P. panaki as junior synonyms of P. perexiguus, because they are all morphologically similar to P. perexiguus beyond differences in size, in agreement with Lindström et al. (1997)LINDSTRÖM S, MCLOUGHLIN S & DRINNAN AN. 1997. Intraspecific variation of taeniate bisaccate pollen within Permian glossopterid sporangia, from the Prince Charles Mountains, Antarctica. Int. J Plant Sci 158: 673-684., who said it should not be used to differentiate species.

On the other hand, Tiwari et al. (1989)TIWARI RS, SRIVASTAVA SC, TRIPATHI A & VIJAYA. 1989. Morphographic study of Permian palynomorphs: Callumispora, Parasaccites, Crucisaccites and Faunipollenites. The Palaeobotanist 37(2): 215-266. made two new combinations, Faunipollenites congoensis and F. magnus. They transferred the species of Striatopiceites congoensis Bose & Kar 1966BOSE MN & KAR RK. 1966. Palaeozoic Sporae Dispersae from Congo. I-Kindu-Kalima and Walikale Regions. Annales de Musée Royal de l’Afrique Centrale, Série 8, Sciences géologiques 53: 1-238. to F. congoensis due to bearing ill- defined central body contrary to a well- defined central body that characterized the genus Striatopiceites. Nevertheless, when Foster (1979)FOSTER CB. 1979. Permian plant microfossils of the Blair Athol Coal Measures, Baralaba Coal Measures, and Basal Rewan Formation of Queensland. GSQ 372: 1-154. made a new combination of Protohaploxypinus perexiguus (Bharadwaj & Salujha), he stated that S. congoensis is undoubtedly closely related to P. perexiguus but is distinguished only by its larger size (154-192 µm) than P. perexiguus (57-110 µm). Further, the type specimens of P. perexiguus fall within the morphological limits of Striatopiceites Zoricheva & Sedova ex Sedova. Foster (1979)FOSTER CB. 1979. Permian plant microfossils of the Blair Athol Coal Measures, Baralaba Coal Measures, and Basal Rewan Formation of Queensland. GSQ 372: 1-154. reassigned Striatopiceites to Protohaploxypinus since it is a less restricted form genera. On these grounds, we re-assign Faunipollenites congoensis and also Faunipollenites magnus to Protohaploxypinus perexiguus.

From the above, it is evident that there are many difficulties to differentiate the mentioned species because of the overlapping of almost all their characters. Hence, this reinforces the proposal of considering P. perexiguus as senior synonym of those species with overlapping features such as the size. It is not discarded that some specimens can show intermediate morphologies with other species of Protohaploxypinus like P. diagonalis and P. pennatulus.

Protohaploxypinus perfectus (Naumova) Samoilovich, 1953

Figures 2.3, 2.5, 3.7, 3.8, 4.6, 4.7, 5.3, 5.16, 6.1, 6.8, 7.3, 8.1

LECTOTYPE. Samoilovich (1953, work translated in 1959 and edited in march 1961).

Additional synonymy see Utting (1994, p. 55) and Foster (1979, p. 86) as P. amplus.

1962 Striatites jacobii Jansonius, p. 67-68, Pl. 14, Figs. 16-17.

1965 Strotersporites indicus Tiwari, p. 199, Pl. 6, figs. 139-141 (holotype fig. 140).

1970 Protohaploxypinus goraiensis (Potonié & Lele) Balme, p. 362, Pl. 11, figs 1-3.

1979 Protohaploxypinus amplus; Foster, p. 86, Pl. 29, Figs. 11-13.

1979 Protohaploxypinus jacobii; Foster, p. 89, Pl. 30, Fig. 11.

1991 Protohaploxypinus amplus; Backhouse, Pl. 17, fig. 12.

1999 Protohaploxypinus goraiensis; MillsteedMILLSTEED BD. 1999. Palynology of the Early Permian coal-bearing deposits near Vereeniging, Free State, South Africa. Bulletin Council Geoscience South Africa 124: 1-81., Pl. 13, figs. 4, 10, 11.

2000 Protohaploxypinus amplus; Playford & Dino, p. 108, Pl. 10, figs. 1-4.

2012 Protohaploxypinus amplus; di Pasquo & GraderDI PASQUO MM & GRADER G. 2012. Palynology and paleoenvironment of the Asselian-?Artinskian Copacabana Formation at Apillapampa near Cochabamba, Bolivia. Palynology 36: 264-276., Pl. 6, fig. 15.

2015 Protohaploxypinus amplus; Stephenson, p. 224, Pl. 4, fig. 8.

DIAGNOSTIC CHARACTERS. Striate pollen bilaterally symmetrical, amb oval to sub-rectangular, haploxylonoid, lateral ridges are often present. Corpus circular to slightly elongate oval (in both ways), frequently distinct, cappa with 7–10 taeniae straight to occasionally branched. Cappula parallel-sided to oval- shaped, width ½ to ¼ of the corpus, distal intexinal folds generally present.

REMARKS. We follow the taxonomic status of P. perfectus by Utting (1994)UTTING J. 1994. Palynostratigraphy of Permian and Lower Triassic rocks, Sverdrup Basin, Canadian Arctic Archipielago. GSC 478: 1-107.. It seems that he cited or considered common species in palynofloras from Northern Hemisphere especially in Russia, as he did not mention P. amplus, widely known in Gondwana, even their close similarity as stated by Foster (1979, p. 87) that further support the synonymy list above.

COMPARISON. Protohaploxypinus baradwajii Foster 1979FOSTER CB. 1979. Permian plant microfossils of the Blair Athol Coal Measures, Baralaba Coal Measures, and Basal Rewan Formation of Queensland. GSQ 372: 1-154. differs from P. perfectus in its t-a elongated central body with rhomboidal amb. The holotypes of Protohaploxypinus jacobii and Strotersporites indicus Tiwari, 1965 are considered junior synonyms of P. perfectus; the same occurs with specimens illustrated in the literature revised (see list above), all because of their similarity. We accept herein the proposal of Hart (1964, p. 32), who considered the genus Strotersporites Wilson 1962WILSON LR. 1962. Permian plant microfossils from the Flowerpot Formation, Greer County, Oklahoma. OGS 49: 1-50. as a junior synonym of Striatopodocarpites Sedova emend. Hart 1964HART GF. 1964. A review of the classification and distribution of the Permian miospores: Disaccate Striatiti. Procceedings Compté Rendu 5° Congrès International de Stratigraphie et de Géologie du Carbonifère, Paris 1963 1: 1117-1129., and the combination of its type species S. communis (Wilson) Hart, based on the grounds of its diploxylonoid amb with striate cappa. However, we do not agree with the argument used by Stephenson (2015, p. 224) of having a monolete mark to maintain Strotersporites separated from Striatopodocarpites.

Finally, the specimen akin to P. jacobii by Foster (1979, Pl. 30, Fig. 12) is better relocated into P. limpidus, and the specimens illustrated as Striatopodocarpites rarus (Bharadwaj & Salujha) Balme (auct. non), by Foster (1979, Pl. 36, figs. 4-5) look morphologically transitional to P. perfectus.

Protohaploxypinus samoilovichii (Jansonius) Hart, 1964

Figures 2.4, 3.1, 6.11, 8.3

BASIONYM. Striatites samoilovichii Jansonius, p. 67, Pl. 14, figs. 9-11.

1991 P. amplus; Backhouse, Pl. 17, fig. 10.

2010 Crescentipollenites fuscus Bharadwaj et al. (auct. non); Murthy et al., p. 707, Pl. 1, fig. 16.

For additional synonym see Foster (1979)FOSTER CB. 1979. Permian plant microfossils of the Blair Athol Coal Measures, Baralaba Coal Measures, and Basal Rewan Formation of Queensland. GSQ 372: 1-154..

DIAGNOSTIC CHARACTERS. Amb slightly diploxylonoid, corpus subcircular to slightly oval in both directions, cappa with well- defined taenia and striae (6-8), cappula broad, rectangular to lensoid in shape, frequently with folds in saccus attachment.

COMPARISON. Crescentipollenites fuscus illustrated by Murthy et al. (2010)MURTHY S, CHAKRABORTI B & ROY MD. 2010. Palynodating of subsurface sediments, Raniganj Coalfield, Damodar Basin, West Bengal. J Earth System Sci 119: 701-710. and P. amplus in Backhouse (1991)BACKHOUSE J. 1991. Permian palynostratigraphy of the Collie Basin, Western Australia. Rev Palaeobot Palynol 67: 237-314. are reassigned to the current species due to its haploxylonoid amb, corpus vertically oval and lensoid cappula. It is noticed that the original diagnosis of Crescentipollenites includes diploxylonoid forms with folds in their attachment zones (Bharadwaj et al. 1974BHARADWAJ DC, TIWARI RS & KAR RK. 1974. Crescentipollenites gen. nov., a new name for hitherto known Lunatisporites Leschik (1955) from the Lower Gondwana. Geophytology 4(2): 141-146.) and it is a junior synonym of Striatopodocarpites (Foster 1979FOSTER CB. 1979. Permian plant microfossils of the Blair Athol Coal Measures, Baralaba Coal Measures, and Basal Rewan Formation of Queensland. GSQ 372: 1-154.).

Species and specimens rejected herein for Protohaploxypinus

• Faunipollenites enigmatusMaheshwari (1969)MAHESHWARI HK. 1969. Paleozoic sporae dispersae from Congo-X. Microfossils from a cliff section at the confluence of Lufupa and Mushyashya rivers, south Katanga. Annals Museum Royal African Center, Series 8o, Science geologist 63: 115-168., F. circumstriatites Maheshwari (1969)MAHESHWARI HK. 1969. Paleozoic sporae dispersae from Congo-X. Microfossils from a cliff section at the confluence of Lufupa and Mushyashya rivers, south Katanga. Annals Museum Royal African Center, Series 8o, Science geologist 63: 115-168., F. gopadensis Bharadawaj & Srivastava (1969) and the specimens illustrated as Protohaploxypinus suchonensis and P. limpidus by MacRae (1988, in Pl. 25, Figs. 15 and 12 respectively)

• Those species and specimens differ from species of Protohaploxypinus in having striations encircled by a marginal striae that is a main feature of Illinites, and they are likely similar to Illinites unicus.

• Striatites rugosusJansonius, 1962JANSONIUS J. 1962. Palynology of Permian and Triassic sediments, Peace River area, western Canada. Palaeöntographica Abt B 110: 35-98. (p. 69, Pl. 14, figs. 18-19). This species was combined without certainty with Protohaploxypinus rugosus (Jansonius) Foster (1979, Pl. 33, fig. 7). The description and the specimens illustrated support their reassignment to Striatopodocarpites due to a diploxylonoid amb and cappa with 4 taenia, although P. hartii could be a transitional form as well (see description in Foster 1979FOSTER CB. 1979. Permian plant microfossils of the Blair Athol Coal Measures, Baralaba Coal Measures, and Basal Rewan Formation of Queensland. GSQ 372: 1-154.).

• The specimen illustrated as P. limpidus by di Pasquo et al. (2018, p. 155, Fig. 15.M) is reassigned herein to Lunatisporites variesectus Archangelsky & Gamerro (1979).

• Protohaploxypinus suchonensis (Sedova) Hart (1964, text-fig. 54) is likely Striatoabieites multistriatus or a species of Vittatina.

DISCUSSION

Since the first palynological studies on Permian rocks began, many striate taxa have been instituted and their classification soon became unreasonable. Hart (1964)HART GF. 1964. A review of the classification and distribution of the Permian miospores: Disaccate Striatiti. Procceedings Compté Rendu 5° Congrès International de Stratigraphie et de Géologie du Carbonifère, Paris 1963 1: 1117-1129. proposed the first attempt to improve the Infraturma Striatiti Pant 1954PANT DD. 1954. Suggestions for the classification and nomenclature of fossil spores and pollen grains. Bot Rev 20: 33-60. in which many genera and species were reassigned to validly published taxa with priority over another and discarding nomen nudum taxa. This kind of taxonomic changes applied to morpho-taxa of fossil palynomorphs are in agreement with the rules recommended in the International Code of Botanical Nomenclature. This important morpho-taxonomic contribution was followed by other palynologists such as Balme (1970)BALME BE. 1970. Palynology of Permian and Triassic Strata in the Salt Range and Surghar Range, West Pakistan. In: Kummel B and Teichert C (Eds), Stratigraphic boundary problems: Permian and Triassic of West Pakistan. University of Kansas Special Publication 4: 305-453., Scheuring (1970)SCHEURING BW. 1970. Palynologische und palynostratigraphische Untersuchungen des Keupers im Bölchentunnel (Solothurner Jura). Schweiz Palaeontol Abh 88: 1-199., de Jersey (1972)DE JERSEY NJ. 1972. Triassic miospores from the Esk Beds.GSQ 357: 1-40., Morbey (1975)MORBEY SJ. 1975. The palynostratigraphy of the Rhaetian Stage, Upper Triassic in the Delbachgraben, Austria. Palaeöntographica Abt B 152: 1-75., amongst others. Foster (1979)FOSTER CB. 1979. Permian plant microfossils of the Blair Athol Coal Measures, Baralaba Coal Measures, and Basal Rewan Formation of Queensland. GSQ 372: 1-154. presented a taxonomic revision of striate pollen grains of genera like Protohaploxypinus, Lunatisporites and Striatopodocarpites including combinations and large synonymy lists. In those contributions, the synonymization of Faunipollenites Bharadwaj 1962BHARADWAJ DC. 1962. The miospore genera in the coals of Raniganj Stage (Upper Permian), India. The Palaeobotanist 9: 68-106. with Protohaploxypinus has been a subject of debate since it was initially proposed by Hart (1964)HART GF. 1964. A review of the classification and distribution of the Permian miospores: Disaccate Striatiti. Procceedings Compté Rendu 5° Congrès International de Stratigraphie et de Géologie du Carbonifère, Paris 1963 1: 1117-1129.. This synonymization was rejected and criticized by Indian workers such as Venkatachala & Kar (1968)VENKATACHALA BS & KAR RK. 1968. Palynology of the Karanpura Sedimentary Basin, Bihar, India-1. Barakar Stage at Badam. The Palaeobotanist 16(1): 56-90., Sinha (1972)SINHA V. 1972. Sporae dispersae from Jhingurdah Seam, Singrauli Coalfield (MP) India. The Palaeobotanist 19(2): 175-201., Tiwari (1974)TIWARI RS. 1974. Palynological succession in the Barakar type area. Geophytology 3: 166-183. and Tiwari et al. (1989)TIWARI RS, SRIVASTAVA SC, TRIPATHI A & VIJAYA. 1989. Morphographic study of Permian palynomorphs: Callumispora, Parasaccites, Crucisaccites and Faunipollenites. The Palaeobotanist 37(2): 215-266.. They argued that the genus Faunipollenites is distinguished from Protohaploxypinus by “the diffused nature of intra-microreticulate central body, common haploxylonoid organization of sacci, merging zones of sacci attachment on distal side, the free ends of striations simulating the body limit and the protosaccate nature of sacci” (Tiwari et al. 1989TIWARI RS, SRIVASTAVA SC, TRIPATHI A & VIJAYA. 1989. Morphographic study of Permian palynomorphs: Callumispora, Parasaccites, Crucisaccites and Faunipollenites. The Palaeobotanist 37(2): 215-266.). Therefore, they claimed that it cannot be synonymized with Protohaploxypinus at any level. Balme (1970)BALME BE. 1970. Palynology of Permian and Triassic Strata in the Salt Range and Surghar Range, West Pakistan. In: Kummel B and Teichert C (Eds), Stratigraphic boundary problems: Permian and Triassic of West Pakistan. University of Kansas Special Publication 4: 305-453. provisionally accepted Hart’s (1964) view since Faunipollenites by definition included haploxylonoid taeniate grains. Foster (1979)FOSTER CB. 1979. Permian plant microfossils of the Blair Athol Coal Measures, Baralaba Coal Measures, and Basal Rewan Formation of Queensland. GSQ 372: 1-154. further reviewed species of Faunipollenites that were synonymized to species of Protohaploxypinus with variable confidence. This contribution was the basement for most of palynologists especially working in the Permian of Gondwana.

Concerning morphologic variations found in species of this striate genus among others, Foster (1979)FOSTER CB. 1979. Permian plant microfossils of the Blair Athol Coal Measures, Baralaba Coal Measures, and Basal Rewan Formation of Queensland. GSQ 372: 1-154. explained that striations may have worked preventing their dehydration in response to environmental conditions. The shrinkage and expansion of the exine is favored by the harmomegathic mechanism protecting its distal germinal area (see also Crane 1986CRANE PR. 1986. Form and function in wind dispersed pollen. In: Blackmore S and Ferguson IK (Eds), Pollen and spores: form and function. Academic Press, London, p. 179-202., 1996CRANE PR. 1996. The fossil history of the Gnetales. Int J Plant Sci 157: 50-57., Vijaya 1990VIJAYA. 1990. Evolutionary pattern of striations and taeniae in the Indian Gondwana saccate pollen. The Paleobotanist 38: 83-91., Chaloner 2013CHALONER WG. 2013. Three palynological puzzles. Int J Plant Sci 174(3): 602-607.). Another interpretation was presented by Tiwari (1982)TIWARI RS. 1982. Nature of striations and taeniae in Gondwana saccate pollen. Geophytology 12: 125-127., who analyzed striate grains from the early Triassic under SEM, and found two different organizations described as follows: 1- taeniate grains with endexine widely exposed in the zone between the taeniae, called platea sensu Vijaya (1990, p. 86), and 2- the striate grains where the ectexine cover all the cappa, reduced in the narrow striae or grooves and without marked taeniae. These two features would have been related to the change in polarity of the germinal area, i.e, in the striate grains the germination occurred through the cappa, whereas in the taeniate ones, from its distal face. Tiwari (1982)TIWARI RS. 1982. Nature of striations and taeniae in Gondwana saccate pollen. Geophytology 12: 125-127. considered it was a global morphological event relatively short lasting up to the end of Triassic. Vijaya (1990)VIJAYA. 1990. Evolutionary pattern of striations and taeniae in the Indian Gondwana saccate pollen. The Paleobotanist 38: 83-91. proposed an evolutionary pattern of striation (striae and taeniae) through the Permian and Triassic succession in India, and discussed the climatic influence on their morphological variations and diversity. Even it is evident the morphologic distinction between those two groups, striate and taeniate taxa, we reinforce the necessity of constraining most of the generic taxa and species into well-recognizable entities to prevent superfluous proliferation such as many of the genera figured by Vijaya (1990, p. 87, text-fig. 2).

Protohaploxypinus is documented from Pennsylvanian (likely Moscovian) up to Triassic palynofloras of the World. Some discrepancies exist today about the function of both harmomegathy and polarity change of germinal area in morphological variations of striate pollen grains due to the lack of modern examples to help in its understanding. Harmomegathia could explain the frequent intermediate forms found between species of the same genus or between genera (Foster 1979FOSTER CB. 1979. Permian plant microfossils of the Blair Athol Coal Measures, Baralaba Coal Measures, and Basal Rewan Formation of Queensland. GSQ 372: 1-154.). This is also noticed in our analysis of species of Protohaploxypinus, thus, four morpho- groups of taxa supported by intermediate forms in between are proposed as follows: 1 – Protohaploxypinus perfectus - limpidus – latissimus – haigii, 2 - Protohaploxypinus perexiguus (P. rugatus) – diagonalis, 3 – Protohaploxypinus pennatulus – microcorpus - baradwajii, 4 – Protohaploxypinus hartii – samoilovichii. Other intermediate forms are also shared with other genera (e.g. Striatopodocarpites, Striatoabieites, Illinites, Lunatisporites).

On the other side, the polarity change of germinal area could have had importance especially in the first steps of the evolution of the gymnosperms during the latest Mississippian-earliest Pennsylvanian (e.g. Foster 1979FOSTER CB. 1979. Permian plant microfossils of the Blair Athol Coal Measures, Baralaba Coal Measures, and Basal Rewan Formation of Queensland. GSQ 372: 1-154., Ouyang 1996OUYANG S. 1996. On the first appearance of some gymnospermous pollen and GSPD assemblages in the sub-Angara, Euramerian and Cathaysia provinces. The Palaeobotanist 45: 20-32., Chaloner 2013CHALONER WG. 2013. Three palynological puzzles. Int J Plant Sci 174(3): 602-607.). In the Permian, the ever-present striate/taeniate taxa and their related floras are used in local to global scale correlations and attested warmer and semiarid climates especially in Gondwana proved by its movement to lower latitudes. In northern Hemisphere they are related to Coniferalean and Peltasperms and in southern Hemisphere to Coniferalean, Glossopterids and also, Peltasperms (Crane 1988CRANE PR. 1988. Major clades and relationships in the “higher” gymnosperms. In: Beck CB (Ed), Origin and Evolution of Gymnosperms. Columbia University Press, New York, p. 218-272., Zhou 1994ZHOU YX. 1994. Earliest pollen-dominated microfloras from the early Late Carboniferous of the Tian Shan Mountains, NW China: their significance for the origin of conifers and palaeophytogeography. Rev Palaeobot Palynol 81: 193-211., Balme 1995BALME BE. 1995. Fossil in situ spores and pollen grains: an annotated catalogue. Rev Palaeobot Palynol 87: 81-323., Césari & Gutiérrez 2001CÉSARI SN & GUTIÉRREZ PR. 2001. Palynostratigraphy of the Upper Paleozoic sequences in central-western Argentina. Palynology 24: 113-146., Zavialova et al. 2001ZAVIALOVA NE, MEYER-MELIKIAN NR & GOMANKOV AV. 2001. Ultrastructure of some Permian pollen grains from the Russian Platform. In: Goodman DK and Clarke RT (Eds), Proceedings of the IX International Palynological Congress, Houston (Texas, U.S.A., 1996). American Association of Stratigraphic Palynologists Foundation, p. 99-114., Tripathi 2002TRIPATHI A. 2002. Major palynological trends in relation to the development of Glossopteris Flora through Lower Gondwana of India. Bull Natl Mus Nat Sci Ser C 28: 1-8., Zhu et al. 2005ZHU HC, OUYANG S, ZHAN JZ & WANG Z. 2005. Comparison of Permian palynological assemblages from the Junggar and Tarim Basins and their phytoprovincial significance. Rev Palaeobot Palynol 136: 181-207., Chaloner 2013CHALONER WG. 2013. Three palynological puzzles. Int J Plant Sci 174(3): 602-607., Zavialova & Karasev 2015ZAVIALOVA N & KARASEV E. 2015. Exine ultrastructure of in situ Protohaploxypinus from a Permian peltasperm pollen organ, Russian Platform. Rev Palaeobot Palynol 213: 27-41.). Its diversity and abundance increased during the Permian, as most of taeniate forms (e.g. Lunatisporites, Lueckisporites) appeared in the early Cisuralian/ or latest Pennsylvanian and soon after, their diversity and abundance increased as well (e.g. Ravn 1986RAVN RL. 1986. Palynostratigraphy of the Lower and Middle Pennsylvanian coals of Iowa. Geological Survey of Iowa, Technical paper 7: 1-245., Zhou 1994ZHOU YX. 1994. Earliest pollen-dominated microfloras from the early Late Carboniferous of the Tian Shan Mountains, NW China: their significance for the origin of conifers and palaeophytogeography. Rev Palaeobot Palynol 81: 193-211., Jones & Truswell 1992JONES MJ & TRUSWELL EM. 1992. Late Carboniferous and Early Permian palynostratigraphy of the Joe Joe Group, southern Galilee Basin, Queensland, and implications for Gondwana stratigraphy. Bureau of Mines and Mineral Resources. J Australian Geol Geophys 13: 143-185., Utting 1994UTTING J. 1994. Palynostratigraphy of Permian and Lower Triassic rocks, Sverdrup Basin, Canadian Arctic Archipielago. GSC 478: 1-107., Loboziak et al. 1997LOBOZIAK S, MELO JHG, DINO R, VACHARD D & STREEL M. 1997. Earliest taeniate bisaccates from the Amazon Basin are not older than Westphalian. Geobios 30: 467-474., Playford & Dino 2000PLAYFORD G & DINO R. 2000. Palynostratigraphy of upper Palaeozoic strata (Tapajós Group), Amazonas Basin, Brazil: Part Two. Palaeöntographica Abt B 255: 87-145., Césari & Gutiérrez 2001CÉSARI SN & GUTIÉRREZ PR. 2001. Palynostratigraphy of the Upper Paleozoic sequences in central-western Argentina. Palynology 24: 113-146., Azcuy et al. 2002AZCUY CL, DI PASQUO M & VALDIVIA AMPUERO H. 2002. Late Carboniferous miospores from the Tarma Formation, Pongo de Mainique, Perú. Rev Palaeobot Palynol 118: 1-28., Souza 2006SOUZA PA. 2006. Late Carboniferous palynostratigraphy of the Itararé Subgroup, northeastern Paraná Basin, Brazil. Rev Palaeobot Palynol 138: 9-29., di Pasquo 2009DI PASQUO MM. 2009. The Pennsylvanian palynoflora of the Pando X-1 Borehole, northern Bolivia. Rev Palaeobot Palynol 157: 266-284., di Pasquo et al. 2015DI PASQUO MM, GRADER GW, ISAACSON P, SOUZA PA, IANNUZZI R & DÍAZ-MARTÍNEZ E. 2015. Global biostratigraphic comparison and correlation of an early Cisuralian palynoflora from Bolivia. Hist Biol 27(7): 868-897., Stephenson 2018STEPHENSON MH. 2018. Permian palynostratigraphy: a global overview. In: Lucas SG and Shen SZ (Eds), The Permian Timescale. Geol Soc Spec Publ 450: 321-347.). Studies of fructifications summarized by Balme (1995)BALME BE. 1995. Fossil in situ spores and pollen grains: an annotated catalogue. Rev Palaeobot Palynol 87: 81-323. show a great variety of striate pollen grain genera although few are related with Protohaploxypinus. Instead, synangia of a Permian glossopterid of Antarctica yielded several striate genera such as Protohaploxypinus, Striatopodocarpites, Striomonosaccites, Crustaesporites (Lindström et al. 1997LINDSTRÖM S, MCLOUGHLIN S & DRINNAN AN. 1997. Intraspecific variation of taeniate bisaccate pollen within Permian glossopterid sporangia, from the Prince Charles Mountains, Antarctica. Int. J Plant Sci 158: 673-684., McLoughlin et al. 1997MCLOUGHLIN S, LINDSTRÖM S & DRINNAN AN. 1997. Gondwanan floristic and sedimentological trends during the Permian–Triassic transition: new evidence from the Amery Group, northern Prince Charles Mountains, East Antarctica. Antarct Sci 9: 281-298.). Another example corresponds to Late Permian isolated synangia Permotheca striatifera (Peltasperm?) from Russia in which several morphotypes of Protohaploxypinus were found (Zavialova & Karasev 2015ZAVIALOVA N & KARASEV E. 2015. Exine ultrastructure of in situ Protohaploxypinus from a Permian peltasperm pollen organ, Russian Platform. Rev Palaeobot Palynol 213: 27-41.).

Those morphological variations could have been enhanced due to another interesting argument proposed by Crane (1986)CRANE PR. 1986. Form and function in wind dispersed pollen. In: Blackmore S and Ferguson IK (Eds), Pollen and spores: form and function. Academic Press, London, p. 179-202.. He referred to the possibility that some gymnosperms were wind- pollinated plants and not obligately anemophylous. Hence, many could have been facultatively entomophilous, as shown by the occurrence of large amounts of typically anemophilous pollen in bee loads. Krassilov et al. (1999)KRASSILOV VA, RASNITSYN AP & AFONIN SA. 1999. Pollen morphotypes from the intestine of a Permian booklouse. Rev Palaeobot Palynol 106: 89-96. analyzed and illustrated the pollen content in the gut of a primitive booklouse from the Urals. In the latter, striate pollen grains mainly of Protohaploxypinus and Lunatisporites genera were recovered, and some monosaccate pollen grains were scarcely found as well. Krassilov et al. (1999)KRASSILOV VA, RASNITSYN AP & AFONIN SA. 1999. Pollen morphotypes from the intestine of a Permian booklouse. Rev Palaeobot Palynol 106: 89-96. have found that several insects in the Permian of Urals may be pollinivores on Protohaploxypinus, Lunatisporites and Vittatina pollen grains that correspond to Ullmaniaceous- Peltasperm fossil plant community. Therefore, one possibility is that these pollinivorous insects could have acted as pollination vectors sensu stricto, and the striae-taeniae (bands) present on their proximal face have facilitated to be stuck to insects or directly, they fed on the pollen grains before being released from the fructification. Considering any of the possible interpretations, it is evident they could have contributed to preserve high amounts of pollen grains into the sediments (i.e., fecal pellets, insects buried into sediments). It follows from what has been said that intra and inter-generic and specific transitional morphologies documented in striate pollen grains could have been triggered either by paleoclimatic changes and/or co-evolution of insect-plants since their appearance (see Crane 1986CRANE PR. 1986. Form and function in wind dispersed pollen. In: Blackmore S and Ferguson IK (Eds), Pollen and spores: form and function. Academic Press, London, p. 179-202., Vijaya 1990VIJAYA. 1990. Evolutionary pattern of striations and taeniae in the Indian Gondwana saccate pollen. The Paleobotanist 38: 83-91., Krassilov et al. 1999KRASSILOV VA, RASNITSYN AP & AFONIN SA. 1999. Pollen morphotypes from the intestine of a Permian booklouse. Rev Palaeobot Palynol 106: 89-96.). These are also recognized between species of Protohaploxypinus as confirmed herein (Table I, Fig. 9) and strengthened the necessity of eliminating more superfluous species documented in the literature from India and elsewhere.

Future perspectives

We aim to emphasize the usage of Protohaploxypinus and its species instead of Faunipollenites in future studies of Permian and Triassic successions in India. The former is well- represented in the Permian as well as in the Pennsylvanian to Triassic of the World. In India, a wider revision of illustrated specimens assigned to Faunipollenites needs to be done. This will improve the stratigraphic ranges of species of Protohaploxypinus in India and globally and will contribute to the knowledge of paleobiogeographic distribution of Permian palynofloras in both northern and southern Hemispheres (e.g. Utting & Piasecki 1995UTTING J & PIASECKI S. 1995. Palynology of the Permian of northern continents: A review. In: Scholle PA, Peryt TM and Ulmer-Scholle PA (Eds), The Permian of Northern Pangea. Vol. 1: Palaeogeography, Palaeoclimates, Stratigraphy. Berlin, Springer-Verlag, p. 236-261., Tripathi 2002TRIPATHI A. 2002. Major palynological trends in relation to the development of Glossopteris Flora through Lower Gondwana of India. Bull Natl Mus Nat Sci Ser C 28: 1-8., Chaloner 2013CHALONER WG. 2013. Three palynological puzzles. Int J Plant Sci 174(3): 602-607., di Pasquo et al. 2015DI PASQUO MM, GRADER GW, ISAACSON P, SOUZA PA, IANNUZZI R & DÍAZ-MARTÍNEZ E. 2015. Global biostratigraphic comparison and correlation of an early Cisuralian palynoflora from Bolivia. Hist Biol 27(7): 868-897., Stephenson 2018STEPHENSON MH. 2018. Permian palynostratigraphy: a global overview. In: Lucas SG and Shen SZ (Eds), The Permian Timescale. Geol Soc Spec Publ 450: 321-347., and references therein). Another consequence of the present revision concerns to the updating of the epithet of palynozones such as the Cisuralian Protohaploxypinus goraiensis Subzone of the Vittatina costabilis Zone of Brazil (see Souza 2006SOUZA PA. 2006. Late Carboniferous palynostratigraphy of the Itararé Subgroup, northeastern Paraná Basin, Brazil. Rev Palaeobot Palynol 138: 9-29.), which should be named as Protohaploxypinus pennatulus. Although, this change does not affect its composition and age.

CONCLUSIONS

Specimens of both Faunipollenites and Protohaploxypinus obtained from palynoassemblages of India and Brazil, mounted after every stage of acid treatment (HCl, HF, HNO₃ between two hours and 24-28 hours and KOH), were carefully analyzed and compared under optic, CLSM and SEM microscopes. From this morpho- taxonomic evaluation, we affirm that a prolonged oxidation treatment (more than 24 hours) introduced morphological changes to the specimens leading to maintain Faunipollenites and Protohaploxypinus separated in Permian studies of India. Our study supported that Protohaploxypinus is the senior synonym of Faunipollenites. Species of Faunipollenites are synonymized with Protohaploxypinus species revised and confirmed herein (Table I, Fig. 9). Specimens bearing transitional features between species of Protohaploxypinus strengthen the necessity of eliminating superfluous species documented in the literature of India and elsewhere. Four morpho- groups of taxa are also recognized considering intermediate forms, joined as follows: 1 –Protohaploxypinus perfectus - P. limpidus – P. latissimus – P. haigii, 2- Protohaploxypinus perexiguus (P. rugatus) – P. diagonalis, 3 – Protohaploxypinus pennatulus – P. microcorpus - P. baradwajii, 4 – Protohaploxypinus hartii – P. samoilovichii. Other intermediate forms are also shared with other genera (e.g. Striatopodocarpites, Striatoabieites, Illinites, Lunatisporites). The future re-assigment of specimens in any of the species of Protohaploxypinus herein confirmed, illustrated in Pennsylvanian to Permo-Triassic palynnoassemblages of India and elsewhere will enable to refine intra-Gondwanan and global correlations and their palaeobiogeographic distribution in association to the respective megafloras.

ACKNOWLEDGMENTS

The authors are grateful to the authorities and heads of BSIP Lucknow, India and, UERJ, and CENPES (Petrobras), Rio de Janeiro, IGc/USP, São Paulo, Brazil, and CICYTTP-CONICET-ER-UADER, Entre Ríos, Argentina, for giving permission to carry out this work in their respective institutions. We are also grateful to the reviewers, Drs. M. Zooba and Lucía Balarino for their useful suggestions and corrections that allowed us to improve our manuscript. To Dr. Peter Isaacson for his revision of the present manuscript to improve the language. This contribution is part of a current major project of collaboration between India, Brazil and Argentina. This work was supported by research grants from Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) of the science without frontiers program to PSK as Bolsa Jovens Talentos Scholarship (No. 300578/2015-6); MS as Post doctorate Junior (No. 150574/2015-0), to MECBO as Research productivity Scholarship (No. 304978/2013-2) and to PIP CONICET 0812-2015-2017.

REFERENCES

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