Faunipollenites Bharadwaj 1962 and Protohaploxypinus Samoilovich 1953 emend. Morbey 1975: morphologic comparison of oxidized and non- oxidized specimens from India and Brazil, and its taxonomic importance

: 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 defi ned corpus and absence of folds in distal attachment. In India, a standard method is applied including the oxidization with HNO 3 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 HNO 3 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 confi rmed. 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.


INTRODUCTION
Faunipollenites was erected by Bharadwaj (1962) to include haploxylonoid striate bisaccate pollen grains bearing a central body with an ill defi ned 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 1964, Balme 1970, Foster 1979, except for Indian workers that are still using it. On the other hand, Protohaploxypinus defi ned by Samoilovich 1953 was widely discussed by several authors especially Hart (1964), who formally emended it to exclude non taeniate pollen grains. Balme (1970) proposed the combination of the type species Faunipollenites varius Bharadwaj 1962 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) 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 1975, MacRae 1988, Utting 1994, Stephenson 2015) and among them, Foster (1979) 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 & Waltz) Samoilovich 1953 although he did not fi nd it in Australia. Instead, this species was described and illustrated by MacRae (1988, Pl. 25, fi gs. 13, 14) and Utting (1994, Pl. 6, fi gs. 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) also discussed the taxonomic status of Faunipollenites and tentatively assigned it as a junior synonym of Protohaploxypinus Samoilovich 1953emend. Morbey 1975. Indian workers (Venkatachala & Kar 1968, Sinha 1972, Tiwari 1974, and especially Tiwari et al. (1989), who treated Faunipollenites in detail, strongly rejected the taxonomic changes proposed for this genus and its type species, on the grounds of an ill defi ned central body outline. As this taxonomic controversy still requires a clarification, specimens attributed to Faunipollenites in palynological material from the Permian of India  and  are examined and compared using optical light microscope , 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 co r re la t i o n s a n d pale o b i o ge o g ra p h i c reconstructions.

Procedence of samples
The studied specimens were recorded from the Damodar and Godavari basins in India  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 2014), 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. 2012). Palynological materials from Brazil were obtained from surface samples of the Paraná Basin at two locations ( Fig. 1e-

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 HNO 3 (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)  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 HNO 3 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 HNO 3 and 2' of KOH  are maintained whereas morphological differences are noticed after the  application of HNO 3 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 1953 andMorbey 1975) and Faunipollenites (Bharadwaj 1962 andTiwari et al. 1989) are characterized by haploxylonoid to slightly diploxylonoid saccus construction. This feature did not suffer distortions after different processes applied and confi rms it is of generic importance.

MERCEDES DI PASQUO et al.
Faunipollenites AND Protohaploxypinus (INDIA, BRAZIL) An Acad Bras Cienc (2021) 93(1) e20190094 8 | 25 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 (6)(7)(8) after each treatment with HCl, HF, and 2 hours of HNO 3 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 confi rm 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) 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 2000 and references). The presence or absence of intexinal folds in Protohaploxypinus is not considered a generic feature (see Foster 1979). 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) did not mention folds in the cb-sacci attachment used to maintain both genera separated (Tiwari et al. 1989).
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) 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 1962) and the emended diagnosis states 6-20 (Tiwari et al. 1989). 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. 1997) 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 1967).

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.   (1)  REMARKS. We agree with Foster , see also Foster & Gomankov 1994 in the constrainment of Protohaploxypinus versus Striatoabieites (most common Striatoabieites multistriatus (Balme & Hennelly) Hart 1964) and Striatopodocarpites. Detailed descriptions and synonymy of genus and species treated in this contribution are mainly addressed by Balme (1970), Morbey (1975), Foster (1979), McRae (1988) and Utting (1994). Protohaploxypinus bharadwajii Foster, 1979HOLOTYPE. Foster (1979, Pl. 29, fig. 10. 1965 P. microcorpus (Schaarschmidt) Clarke, p. 338, Pl. 41, Fig. 3.

F. bharadwajii
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 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.
COM PA R I S O N . T h i s s p e c i e s b ea rs 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)  See Balme (1970), Foster (1979) and MacRae (1988) 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.
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 Table I Fig. 16.
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.
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) (2000) 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.

. D I A G N O S T I C C H A R A C T E R S .
A m b haploxylonoid, subcircular to vertically oval, central body indistinct, circular-subcircular to vertically oval. Cappa with taenia not welldefined, 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 1979).
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) 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) synonymized F. parvus with F. perexiguus, whilst Millstead (1999 considered them as junior synonyms of P. limpidus. Balarino (2012) 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). MacRae (1988) 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). 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), who said it should not be used to differentiate species.
On the other hand, Tiwari et al.  (1979) 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) 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.
REMARKS. We follow the taxonomic status of P. perfectus by Utting (1994). 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 1979 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 1962 as a junior synonym of Striatopodocarpites Sedova emend. Hart 1964, 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.

DISCUSSION
Since the first palynological studies on Permian rocks began, many striate taxa have been instituted and their classification soon became unreasonable. Hart (1964) proposed the first attempt to improve the Infraturma Striatiti Pant 1954 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), Scheuring (1970( ), de Jersey (1972, Morbey (1975), amongst others. Foster (1979) (1979) 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 1986, Vijaya 1990, Chaloner 2013. Another interpretation was presented by Tiwari (1982), 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) considered it was a global morphological event relatively short lasting up to the end of Triassic. Vijaya (1990) 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 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 Mississippianearliest Pennsylvanian (e.g. Foster 1979, Ouyang 1996, Chaloner 2013. In the Permian, the everpresent 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 1988, Zhou 1994, Balme 1995, Césari & Gutiérrez 2001, Zavialova et al. 2001, Tripathi 2002, Zhu et al. 2005, Chaloner 2013, Zavialova & Karasev 2015. 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 1986, Zhou 1994, Jones & Truswell 1992, Utting 1994, Loboziak et al. 1997, Playford & Dino 2000, Césari & Gutiérrez 2001, Azcuy et al. 2002, Souza 2006, di Pasquo 2009, di Pasquo et al. 2015, Stephenson 2018. Studies of fructifications summarized by Balme (1995) 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. 1997. Another example corresponds to Late Permian isolated synangia Permotheca striatifera (Peltasperm?) from Russia in which several morphotypes of Protohaploxypinus were found (Zavialova & Karasev 2015).
Those morphological variations could have been enhanced due to another interesting argument proposed by Crane (1986). 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) 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) 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 intergeneric 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 1986, Vijaya 1990, Krassilov et al. 1999. 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 1995, Tripathi 2002, Chaloner 2013, di Pasquo et al. 2015, Stephenson 2018. 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 2006), which should be named as Protohaploxypinus pennatulus. Although, this change does not affect its composition and age.