ISSN 00310301, Paleontological Journal, 2015, Vol. 49, No. 6, pp. 584–594. © Pleiades Publishing, Ltd., 2015. Original Russian Text © T.B. Leonova, M.S. Boiko, 2015, published in Paleontologicheskii Zhurnal, 2015, No. 6, pp. 26–37.
Phylogeny of the Late Paleozoic Superfamily Adrianitoidea Schindewolf (Ammonoidea) T. B. Leonova and M. S. Boiko Borissiak Paleontological Institute, Russian Academy of Sciences, Profsoyuznaya ul. 123, Moscow, 117997 Russia email:
[email protected],
[email protected] Received November 25, 2014
Abstract—The current state of knowledge on the Late Paleozoic ammonoid superfamily Adrianitoidea is reviewed. The section on Adrianitoidea in the revised edition of the Treatise on Invertebrate Paleontology (TIP) on Carboniferous and Permian ammonoids (Furnish et al., 2009) is discussed. Major phylogenetic trends in this superfamily are reconstructed. New data on the morphology of members of the subfamilies Adrianitinae, Emilitinae, and Pamiritellinae allow convincing hypotheses of the early evolution, diversifica tion, and decline in each of the above groups. Keywords: ammonoids, evolution, morphogenesis, phylogenetic scheme, Late Paleozoic DOI: 10.1134/S0031030115060064
INTRODUCTION Adrianitoidea is a taxonomically distinct group of Late Paleozoic ammonoids with a specific mode of sutural ontogeny, with the initial formula (V1V1)LU:U1ID. In contrast to other goniatitids, new umbilical lobes were formed on the top of the umbili cal saddle and, alternating, were shifted to the inner and outer sides of the whorl, respectively following the formula (V1V1)LU:ID → (V1V1)LUU2:U1ID → (V1V1)LUU2Un + 1:UnU1ID. The external lateral lobe did not divide. This mode of sutural ontogeny is unique to Adrianitoidea, and it was proposed to con sider these ammonoids as a suborder Adrianitina within the order Goniatitida (Leonova, 2002). In all Adrianitina, the suture is “Adrianitinalike,” with entire, narrow, deep lobes, the number of which can vary from 10 to 34. The shell shape and ornamentation are very diverse, but pachyconic and spheroconic involute shells with reticulate ornamentation are par ticularly common. Shigeta et al., 2001 showed that Adrianitoidea, like many other taxa of Permian goni atitids, have an ellipsoidal caecum, a short prosiphon, and a ventral siphuncle beginning from the first whorl. Our study corroborated these findings. Members of the superfamily Adrianitoidea composed a consider able part of the Late Paleozoic ammonoid biota. Their occurrences are known from Late Carboniferous and mostly Permian beds of various parts of Asia, Europe, North and Central Americas, and Australia. Their maximum taxonomic and morphological diversity is observed in the Middle Permian of Sicily and Timor. Numerous taxa of adrianitids are found in the Early and Middle Permian of Pamir, Crimea, and Kurdis tan, and they are somewhat less abundant in the
Early–Middle Permian series of North America (Nevada, Texas, and Mexico) and China; three genera are established in the Urals and only one species in each of the following three regions: Transcaucasia, Far East, and Australia. In our interpretation, the superfamily Adriani toidea includes two families: Adrianitidae Schinde wolf, 1931 and Palermoceratidae Zhou et Glenister, 2009 (=Hoffmanniidae Plummer et Scott, 1937). These families are different in size: the first unites four subfamilies, including 21 genera and 78 species, and the second includes only one genus with two species. A relatively complete history of the study of Adri anitoidea was provided by Leonova and Bogoslovskaya (1990). In this paper we will only discuss those issues that have not been discussed previously. This is mainly a result of the authors of the revised TIP on Carbonif erous and Permian ammonoids (Furnish et al., 2009) accepting a concept of the superfamily that included only one family Adrianitidae and four subfamilies: Adrianitinae Schindewolf, 1931, Dunbaritinae Miller et Furnish et Schindewolf, 1957, Palermoceratinae Zhou et Glenister, 2009, and Texoceratinae Ruzhencev et Bogoslovskaya, 1978. In our opinion, such a system of Adrianitoidea is not based on phylogeny. Unfortunately, the authors of the review did not take into account the studies of Ruzhencev (1974), which shed light on the previously uncertain origin of the genus Dunbarites Miller and Furnish, 1940 and the monotypic family Dunbaritidae. It should be said that the authors of the genus, even when they established it, were in doubt as to whether it was close to Emilites and Adrianitidae. They suggested that Dunbarites was “possibly more closely related to Emilites than to any
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(b)
(a) Fig. 1. Sutures: (a) Dunbarites rectilateralis (Miller), (b) Emilites incertus (Böse); both at Dm = about 10 mm, ×6 (after Miller and Furnish, 1940a, textfig. 4).
other genus, and they occur at the same general hori zon and in the same general area” (Miller and Fur nish, 1940a, p. 532), which certainly not sufficient to assign them to a family. Dunbarites and Emilites are very different in shell morphology, whereas their sim ple tenlobed sutures are only remotely similar (Fig. 1). Ruzhencev (1974) gave a detailed account of the history of studies of this genus and its assignments to various families. He substantiated its assignment to a separate family and showed its fundamental differ ence from Emilites, as well as the impossibility of its assignment to Adrianitidae. He suggested that the ori gin of Dunbarites should be linked to the genus Pseudoparalegoceras Miller, 1934 from the family Pseudoparalegoceratidae Librovitch, 1957 superfam ily Somoholitoidea Ruzhencev, 1938. Ruzhencev (1974) described a new species of Dunbarites (D. larus). Previously, only the type species D. rectilat eralis (Miller, 1930) was known, described based on only two specimens. The additional material allowed a better substantiation of the position of the genus in the system of Paleozoic ammonoids. In a later work (Ruzhencev and Bogoslovskaya, 1978), the family Dunbaritidae with the single genus Dunbarites was assigned to the superfamily Somoholitoidea. This remains the best supported position. The second problem with the ammonoid systemat ics in the revised edition of TIP is that some wellbased phylogenetic lineages (Adrianitinae, Emilitinae, and Pamiritellinae) recognized within this group (Leonova and Bogoslovskaya, 1990; Leonova, 2002) (each with a separate evolutionary mode) did not receive subfam ily status, whereas the Texoceratinae (which includes a single genus Texoceras) was recognized as a subfamily. Since the TIP is a reference book, explanations for the taxonomic assignments are not included, and this lack of explanation leads to some complications. PALEONTOLOGICAL JOURNAL
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The subfamily Texoceratinae differs from other groups of adrianitids by the way in which new lobes shift from the umbilical zone, with the lobes moving first to the external side, and then to the internal side. Therefore Texoceratinae have one lobe more on the external side than on the internal. Only two genera of adrianitids, Metacrimites Ruzhencev, 1950 and Texo ceras Miller et Furnish, 1940 show this feature. Metac rimites currently includes nine species continuing over a long span of time (Kungurian–Capitanian ages) (Leonova, 2002). This genus is probably connected in its origin with Crimites. It is distinguished by the devel opment of a fourth lobe on the external flank of the shell. On the internal side, three lobes remained as in Crimites, an example of deviation of the order of new lobes displaced from the umbilical region. The umbil ical region of the suture was composed of one relatively deep lobe or several shallow lobes, as in Crimites. By the beginning of the Wordian, the ventral lobe (prima rily the ventral prongs) somewhat narrowed in the course of the evolution, and this feature remained throughout the entire time of existence of this genus. The shell shape changed, from the narrow umbilicate subspherical shape inherited from the ancestor to pachyconic. As in some species of Crimites, Metacrim ites has a very distinct, dominant spiral ornamenta tion. Metacrimites most likely branched off at the end of the Artinskian age because its earliest known species M. newelli, of Kungurian age has clearly defined fea tures of the genus. In Texoceras represented by a single species from the Roadian of Texas, the suture is prim itive, with a moderately narrow ventral lobe with nar row prongs rounded at the base, with three lobes on the external flank and two on the inner side, and a pachyconic shell with nodes in the umbilical region and prominent reticulate ornamentation (Fig. 2). The origin of Texoceras, is apparently related to one of the Metacrimites species sharing the same mode of sutural
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Capitanian
Pamiritellinae
Metacrimites dunbari
Wordian
Pseudagathiceras spinosum
Doryceras fimbriatum
Epadrianites beyrichi Adrianites elegans Sosiocrimites insignis
Roadian
Palermites distefanoi
Texoceras texanum
Texoceratinae Kungurian
Neocrimites pavlovi Pseudoemilites asianus
Pamiritella vinogradovi Neocrimites dutkevitchi
Artinskian
Veruzhites pamiricus
Adrianitinae Neocrimites fredericksi
Sakmarian
Crimites shyndensis
Emilitinae Asselian
Emilites prosperus
Gzhelian
Fig. 2. Phylogenetic reconstruction of the family Adrianitidae. Explanations: arrows pointing up show accelerations, arrows pointing down show retardations, twoheaded arrows show mosaic forms.
ontogeny (unpaired number of the external and inner lateral lobes). Apparently, Texoceras represented a paedomorphic, deadend branch of Metacrimites, occurring as a result of the regressive development of the septum: disappearance of one pair of lobes. Cantú Chapa (1997) studying Permian ammonoids of Mexico, commented on the taxonomic position of the species which we assigned to Metacrimites. He assigned several species M. newelli (Miller et Furnish, 1940), M. defordi (Miller et Furnish, 1940), M. plum meri (Miller, 1944), and M. dunbari (Miller et Fur nish, 1940) from the Lower and Middle Permian beds of Texas and Mexico to a new genus Millerites Cantú Chapa, 1997. Apparently this author was not familiar with Ruzhencev’s (1950) paper, in which he assigned the above species to the genus Metacrimites, because there is no mention of this in his monograph (Cantú Chapa, 1997). Irrespective of that, based on some characters (number of lobes, a generally straight shape of the external portion of the suture) Cantú Chapa separated them, in contrast to Miller and Furnish (1940b), from Adrianites, and also from Neocrimites, and placed them in a separate genusgroup taxon. In
the revised TIP (Furnish et al., 2009), the genus Metacrimites was synonymized under Neocrimites without taking into consideration the essential differ ences in the sutural ontogeny. It appears that the data on adrianitoids were not comprehensively analyzed when that revision was prepared. Apart from the above, in the TIP, the name Hoff manniidae Plummer et Scott, 1937 was replaced by Palermoceratinae Zhou et Glenister, 2009, while the genus Hoffmannia Gemmellaro, 1887 was replaced by Palermoceras Zhou et Glenister, 2009, and the family and subfamily were also renamed. This procedure was legitimate, within the rules of the International Code of Zoological Nomenclature (1999, Art. 52.2, Russian translation, 2004). The name Hoffmannia had been preoccupied twice, by Hoffmannia Heinemann et Wocke, 1877 (Insecta), and Hoffmannia Forcart, 1953 (Gastropoda) (Furnish et al., 2009). In our opinion, this taxon should be assigned to a separate family Pal ermoceratidae, even though the family is monotypical (Leonova, 2002), because it differs profoundly from all other Adrianitoidea by the ophioconic shell and coarse dichotomous ribbing. In addition, the suture is PALEONTOLOGICAL JOURNAL
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typically adrianitid, moderately complex, with four lobes on the flanks allowing its positive assignment to Adrianitoidea. We had at our disposal large collections of adrianit ids from the Permian of the Urals (PIN, coll. no. 317) and Pamir (PIN, coll. no. 3951), and we reexamined members of three major subfamilies Emilitinae, Adri anitinae, and Pamiritellinae. The results of this study are discussed below. The distribution of adrianitids in different regions of the world are discussed in the following works: Urals (Ruzhencev, 1950, 1956, 1974; Bogoslovskaya, 1962), Crimea (Toumanskaya, 1931, 1937), Sicily (Gemmel laro, 1887), Oman (Bledinger et al., 1992), Afghani stan and Iraq (Termier et al., 1972; Vaší cek and Kull mann, 1988), Pamir (Leonova and Dmitriev, 1989; Leonova, 2011), Timor (Haniel, 1915; Smith, 1927), USA, Canada and Mexico (Miller and Furnish, 1940a, b; Nassichuk and Henderson, 1986; Mapes and Boardman, 1988; Schiappa et al., 1995; Cantú Chapa, 1997), Australia (Glenister and Furnish, 1961; Armstrong et al., 1967), China and Malaysia (Liang Xiluo, 1981, 1982; Sheng, 1984, 1988), Japan (Ehiro and Araki, 1997). ˆ
RESULTS AND DISCUSSION Subfamily Emilitinae Leonova et Bogoslovskaya, 1990 The diagnosis of the subfamily is given (Leonova and Bogoslovskaya, 1990; Leonova, 2002). In its com position we discuss six genera from the Kasimovian– Wordian. The species composition of these genera is discussed below: Genus Emilites Ruzhencev, 1939 (=Plummerites Miller et Furnish, 1940). Altogether seven species, three of which are in open nomenclature. The type species E. incertus (Bose, 1917) comes from the Gzhe lian of Texas (Gaptank Formation); E. bennisoni Mapes et Boardman, 1988 from the Kasimovian of Oklahoma (Yola Formation), E. brownwoodi Mapes et Boardman, 1988 from the Gzhelian of Texas (Caddo Creek Formation); E. plummeri Ruzh., 1941 from the upper part of the Gzhelian of the Urals; E. prosperus Ruzh., 1978 from the upper part of the Asselian–Sak marian of Pamir (Tashkazyk Formation) and South China (Maping Formation); and also ?E. sp. from the upper part of the Moscovian (Des Moines) of North America; E. sp. from the Gzhelian of Central Asia and E. sp. from Asselian of Arctic Canada (lower part of the Belcher Channel). Genus Crimites Toumanskaya, 1937. The type spe cies C. pamiricus Toumanskaya, 1937 from the Kun gurian (Bolorian) of Pamir (Kochusu Formation); C. elcoensis Miller and Furnish et Clark, 1944 from the Sakmarian of Texas (Wolfcamp Formation; C. glomu lus Ruzhencev, 1952 from the Sakmarian of the Urals (Sterlitamakian); C. subkrotowi Ruzhencev, 1938 from the Sakmarian and Artinskian of the Urals (Sterlita PALEONTOLOGICAL JOURNAL
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makian–Baigendzhinian); C. krotowi Karpinsky, 1889 from the Artinskian of the Urals (Aktastynian– Baigendzhinian); C. singularis Bogoslovskaya, 1962 from the Artinskian of the Urals (Baigendzhinian); C. doliaris Leonova, 1988 from the Artinskian and Kungurian (Yakhtashian–Bolorian) stages of Pamir (Chelamchi and Kochusui formations); C. murgaben sis Pavlov, 1967, C. smithi Toumanskaya, 1937, C. spathi Toumanskaya, 1937 from the Kungurian (Bolorian) of Pamir (Kochusu Formation); C. shyn densis Leonova, 1988 from the Kungurian (Bolorian) of Pamir (Kochusu and Shindy formations); C. hanieli Toumanskaya, 1931 and C. gemmellaroi Toumanskaya, 1931 from the Kungurian of the Crimea (Soraman Beds); C. sp. from the Sakmarian of Pamir (Khoridzh Formation). Genus Nevadoceras Schiappa et Spinosa, 1995; only the type species N. steelei Schiappa et Spinosa, 1995 from the Sakmarian–Artinskian of Nevada (Por tuguese Spring Formation). Genus Istycoceras Pavlov, 1967; only the type spe cies I. bodylevskyi Pavlov, 1967 in the Kungurian (Bolorian) of Pamir (Kochusu Formation). Genus Veruzhites Leonova, 1988; only the type species V. pamiricus Leonova, 1988 in the Artin skian–Kungurian (Yakhtashian–Bolorian) of Pamir (Chelamchi–Kochusu formations). Genus Pseudoemilites Leonova, 1988; only the type species P. asianus Leonova, 1988 in the Kun gurian (Bolorian) of Pamir (Kochusu Formation). Representatives of the subfamily Emilitinae are characterized by a spheroconic, subspheroconic or pachyconic involute shell, relatively simple septum that changes insignificantly throughout growth. The number of lobes remained constant for most above genera (Table 1). The shell ornamentation became very diverse, although in each generic group, the major directions in its evolution were more or less main tained. Emilitins, like other adrianitids, typically have a very low projection of transverse ornamentation (lamellae, constrictions), or complete absence of cur vature on the venter. The peak of the radiation of the subfamily was in the Artinskian–Kungurian (Fig. 3). The main region for this group was apparently the upper part of the pelagial, as they can be assigned to the “Plankton 2 group,” based on their septal mor phology, following the classification used by Barskov et al. (2008). The earliest representative of the subfamily and the entire suborder was certainly Emilites (Figs. 1, 2). It had a subspheroconic shell and a 10lobe suture (V1V1) LU:U1ID. Emilites gave rise to Crimites, a large and very long surviving genus, which gave rise to almost all lineages of adrianitids. Crimites inherited the ancestral subspheroconic shell covered by festoon ornamenta tion at early stages of its development and a morpho logically very similar suture including one pair of lobes too many. In Crimites, as in Emilites the number of
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LEONOVA, BOIKO 35 30 25 20
4 3
15
2 1
10 5
M
os co vi K an as im ov ia n G zh el ia n As se lia n Sa km ar ia n Ar tin sk ia n K un gu ria n Ro ad ia n W or di an C ap ita W ni uc an hi ap in gi an
0
Fig. 3. Dynamics of species diversity of adrianitids by subfamilies. Explanations: (1) Emilitinae, (2) Adrianitinae, (3) Texocera tinae, (4) Pamiritellinae.
major lobes on the flank portion of the suture and cor responding lobes of the inner portion remained unchanged throughout its existence. All species of Crimites typically have a relatively shallow ventral lobe with narrow branches, separated by a relatively low saddle (SH/LW from 0.45 to 0.70). Compared to the ancestral genus, the external lobes became deeper and narrower, with a pointed base and constricted anterior portion, i.e., acquired a specific “adrianitid” shape. In many species of Crimites, a series of small lobes devel ops in the umbilical zone the number of which varies even within the same species. Retaining the general involute shape, the shell varied reasonably widely: from spheroconic to subspheroconic or pachyconic with various shapes of whorl cross section. The orna mentation was both festoonlike and reticulate, with either longitudinal or transverse elements dominating. Transverse elements formed a ventral projection. Crimites was certainly linked to Nevadoceras and Isty coceras. The American endemic Nevadoceras (Sak marian–Artinskian) is similar to Crimites in the sutural morphology, but is distinguished by a larger ventral lobe with a relatively high median saddle (HS/LD=0.7). Apart from that, it acquired a wider umbilicus, a narrower shell with the thin transverse ornamentation, rectiradiate on the venter (Schiappa et al., 1995). The Pamir endemic genus Istycoceras apparently branched off at the end of the Artinskian. It has a considerably narrower umbilicus, pachyconic shell shape, and no spiral ornamentation, like Neva doceras, which retained a suture very similar to the
ancestral kind. These two genera may be considered as examples of close parallel development through suc cessive iterations, based on genetic relationship and similar habitats. The main phylogenetic lineage within the subfam ily Emilitinae is the regressive succession Crimites → Veruzhites → Pseudoemilites (Fig. 2), developing at the end of the early Permian, the main trends in its devel opment in the reverse succession repeat the evolution of the orthogenetic lineage Emilites → Crimites. Dur ing the transition from Crimites to Veruzhites in the Artinskian, the sutural outline changed in that the ventral prongs became extremely wide and the height of the median saddle increased (HS/LD = 0.85), whereas the number of the main elements did not change. The ornamentation showed only thin, distinct transverse lamellae. The constrictions form, as in the ancestor, a pronounced ventral projection. The shell is pachyconic, its width varying within certain limits (WW/Dm from 0.57 to 0.67 at a shell diameter over 15 mm and from 0.64 to 0.74 at Dm < 15 mm), which is particularly well outlined on a large number of mea sured specimens. However this is not sufficient to establish species groups, because all these specimens form continuous successions with gradual morpholog ical changes. In turn, Veruzhites gave rise to Pseudoe milites, which existed in the Kungurian. In this genus, the ventral prongs and lateral lobes retained a parallel sided shape but unlike their ancestor, their number decreased to two, whereas the height of the median saddle increased even more (SH/LD = 0.94). As in PALEONTOLOGICAL JOURNAL
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Pachycone
Pachycone
Pachycone
Subspherocone
Spherocone– fusiform
Spherocone
Pachycone
Spherocone–sub Very narrow spherocone
Cadicone
Istycoceras
Veruzhites
Pseudoemilites
Neocrimites
Aricoceras
Adrianites
Sosiocrimites
Epadrianites
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Vol. 49
Adrianitinae
Pamiritellinae
Pachycone
Pachycone
Doryceras
Discocone
Pamiritella
Sizilites
Narrow
Discocone
Pamirioceras
Mediumsized
Mediumsized
Mediumsized
Moderately narrow
Moderately wide
Pachycone
Straight
Straight
Straight
Straight
Straight
Straight
Arched
Arched
Reticulate with spiral Arched
Reticulate
Reticulate
Straight or arched
Straight
Transverse (lamellae, Straight costellae)
Reticulate with trans Straight verse
Reticulate with trans Straight or arched verse
Transverse
Transverse
Reticulate with spiral, Straight spines
3
4
5
6
5
4
4–5
5–8
6–7
4–7
4–6
2
3
3
3
3
2
The outline Number of the external of external lobes portion of the suture
Narrow or moderate Reticulate with spiral Straight or arched ly narrow
Moderately narrow
Closed
Pseudagathiceras Subcadicone– discocone
Palermites
Transverse
Transverse
Transverse
Transverse
Festoonshaped reticulate
Festoonshaped
Ornamentation
Very narrow or mod Reticulate with spiral Straight or arched erately narrow
Closed
Closed
Very narrow
Narrow
Very narrow or nar row
Nevadoceras
Umbilicus
Spherocone– pachycone
Shell shape
Crimites
Genus
Spherocone–sub Very narrow spherocone
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Emilitinae
Table 1. Major diagnostic features of the members of the subfamilies Adrianitinae, Emilitinae, and Pamiritellinae
Narrow
Narrow
Narrow
Narrow
Narrow
Medium width
Medium width
Narrow
Narrow
Narrow
Narrow
Wide
Wide
Narrow
Medium width
Medium width
Medium width
Ventral prong width
Low
Low
Low
Low
Mediumsized
Mediumsized
Mediumsized
Low
Low
Low
Low
High
High
Mediumsized
High
Mediumsized
Mediumsized
Median saddle height PHYLOGENY OF THE LATE PALEOZOIC SUPERFAMILY ADRIANITOIDEA 589
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Veruzhites, Pseudoemilites has only transverse orna mentation represented by lamellae, forming a weak ventral projection. As a result, it became extremely similar morphologically to the Early Permian Emilites, i.e., showing features of retroconvergence (Fig. 2). Subfamily Adrianitinae Schindewolf, 1931 The diagnosis of the subfamily was given in previ ous papers (Leonova and Bogoslovskaya, 1990; Leonova, 2002). In this subfamily we include seven genera from the Artinskian–Amarassian stages. Below is the species composition of these genera. Genus Adrianites Gemmellaro, 1887. The type species A. elegans Gemmellaro, 1887 from the Word ian of Sicily (Sosio Beds), Oman (Hamrat Duru For mation), Croatia (MrzlaVodica Beds) and the ?Roa dian of Kurdistan; A. isomorphus Gemmellaro, 1887 from the Wordian of Sicily (Sosio Beds) and Croatia (MrzlaVodica Beds). Genus Neocrimites Ruzhencev, 1940. The type species N. fredericksi Emeliancev, 1929 and N. stuck enbergi Karpinsky, 1889 from the Artinskian of the Urals (Baigendzhinian Substage); N. dutkevichi Pav lov, 1972 from the Artinskian and Kungurian (Yakhtashian and Bolorian) stages of Pamir (Chelam chi and Kochusu formations); N. nalivkini Touman skaya,1949 and N. pavlovi Leonova, 1988 from the Kungurian (Bolorian) of Pamir (Kochusu Forma tion); N. oyensi (Haniel, 1915) from the Sakmarian and Kungurian of Timor (Atsabe and Bitauni beds); N. rothpletzi (Haniel, 1915) from the Kungurian of Timor (Bitauni Beds); N. guangxiensis Chao et Liang, 1974 from the Kungurian of South China and Malay sia (Lee Mine Beds–Kungurian–Wordian?); N. sp. from the Kungurian of Western Australia (Coyrie For mation). Genus Sosiocrimites Ruzhencev, 1950 (=Subcrim ites Liang, 1982). The type species S. insignis (Gem mellaro, 1887) from the Wordian of Sicily (Sosio Beds), Oman (Hamrat Duru Formation), Kurdistan (Kulkula Formation); S. biassalensis (Toumanskaya, 1931) from the Roadian of the Crimea (Burnian); S. compressus (Liang, 1982) from the Wordian of Northwestern China (Fanjiatun Formation); S. dar vasicus Leonova, 1992 from the Artinskian (Yakhtash ian) of Pamir (Chelamchi Formation); S. sp. from the Roadian (Kubergandian) of Pamir (Leonova, 2011); S. jintaensis (Liang, 1981) from the Roadian of North western China (Shuangputang Formation); S. jialaen sis (Sheng et Liu, 1983) from the Wordian of Tibet (Jiala Formation); ?S. minuta Sheng, 1988 from Kun gurian–Roadian of Tibet (Urulun Formation); ?S. nitidus Sheng, 1984 from the Capitanian of Tibet (Xiukang Formation). Genus Aricoceras Ruzhencev, 1950. The type spe cies A. ensifer (Gemmellaro, 1887) from the Wordian of Sicily (Sosio Beds), and also Oman (Hamrat Duru Formation) and Kurdistan (Qulqula Formation);
A. meridionale (Teichert et Fletcher, 1943) from the Kungurian of Eastern Australia (Middle Jimpy For mation, Elderslie); A. cancellatum (Haniel, 1915) from the Kungurian–Wordian stages of Timor (Bitauni Beds, Tae Wei, Basleo); A. discoidale (Haniel, 1915) from the Wordian of Timor (Basleo Beds), A. warreni (Miller et Crockford, 1936) from the Word ian British Columbia (Cache Creek Formation). Genus Neoaricoceras Ruzhencev, 1950; only the type species N. kingi (Gemmellaro, 1887) from the Wordian of Sicily (Sosio Beds). Genus Epadrianites Schindewolf, 1931 (=Basleo ceras Ruzhencev, 1950). The type species E. timorensis (Boehm, 1907) from the Capitanian–Dzhulfian of Timor (Amarassi Beds), and also Tibet (Lansuo For mation); E. beyrichi (Haniel, 1915) from the Kun gurian–Wordian of Timor (Bitauni Beds, Basleo) and Oman (Hamrat Duru Formation); E. haueri (Gem mellaro, 1888) from the Wordian of Sicily (Sosio Beds) and Croatia (MrzlaVodica Beds); E. stenosella tus Liang, 1982 from the Wordian of Northwestern China (Fanjiatun Formation); E. involutus (Haniel, 1915) from the Capitanian–Dzhulfian of Timor (Amarassi Beds); E. kotljarae (Yu. Zakharov, 1983) from the Dzhulfian of Transcaucasia. Genus Pseudagathiceras Schindewolf, 1931. The type species P. wichmanni (Haniel, 1915) from the Wordian of Timor (Basleo Beds); P. difuntense Miller, 1944 and P. spinosum Miller, 1944 from the Wordian of Mexico (Beds with Waagenoceras); P. ornatum Ehiro et Araki, 1997 from the Capitanian of Japan (Ochia Formation). This group includes adrianitids with a very variable septum and external shell morphology, as well as wide variations in size. Members of this genus, during its relatively short time existence, showed progressive increase in the number of lateral lobes, whereas the external portion of the suture was frequently arched. The shell shape was also very changeable (from spheroconic and completely involute to fusiform or pachyconic and cadiconic with the umbilicus from closed to moderately wide), and character of orna mentation (reticulate, with predominant spirals ele ments, sometimes with spines). The maximum species diversity was in the Wordian (Fig. 3). The Adrianitina mainly inhabited the pelagial; and most of them belonged to the “plankton 1” living form, some to “plankton 2,” like emilitines, whereas some taxa (e.g., Epadrianites) belonged to the benthopelagic living form (Barskov et al., 2008). The genus Neocrimites was the initial in the sub family. It branched off Crimites in the Artinskian due to the increase in the shell width and acquisition of addi tional pairs of lateral lobes. That genus gave rise to sev eral branches. Evolution in the lineage Neocrimites was connected with increase of shell width up to acquisi tion of a barrelshape or fusiform shape (N. pavlovi) (Fig. 4). It should be noted that the assignment of some pachyconic taxa to this genus is erroneous, PALEONTOLOGICAL JOURNAL
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PHYLOGENY OF THE LATE PALEOZOIC SUPERFAMILY ADRIANITOIDEA N. fredericksi
N. nalivkini
(a)
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N. pavlovi
(d)
(g)
(b)
(e)
(h)
(c)
(f)
(i)
Fig. 4. Shells, cross sections, and sutures of Neocrimites species: (a–c) N. fredericksi, specimen no. 14103 at WW/Dm = 1.05 (after Ruzhencev, 1956); (d–f) N. nalivkini, specimen no. 3591/359 at WW/Dm = 1.25; (g–i) N. pavlovi, specimen no. 3591/360 at WW/Dm = 1.53, scale is the same for all shells.
because the main diagnostic character of the genus is the large whorl width often exceeding its diameter. In the type species N. fredericksi, WW/Dm varies from 0.95 to 1.19 (Ruzhencev, 1956); in N. stuckenbergi it varies from 1.11–1.15 (Bogoslovskaya, 1962); N. nalivkini from 1.01 to 1.08 and N. pavlovi from 1.40 to 1.78 (Leonova and Dmitriev, 1989). The number of lobes increased from four in the Artinskian specimens from the Urals to six in the Kungurian (Bolorian) Pamir species. The external portion of the suture retained the straight shape. Glenister (pers. comm. and unpublished manuscript) suggested that the barrel shaped and fusiform members of Neocrimites should be assigned to a separate genus, an opinion that we do not share. The above figures show that the transversely elongated shell shape in this genus is a characteristic feature, whereas in N. pavlovi, this is a marginal mem ber of this lineage. A few transversely elongated speci mens found in collections from the Sosio Beds and Nevada fit well into the general lineage of evolution of the shell shape in Neocrimites. The ornamentation throughout the duration of the genus remained reticu late with dominant spiral elements. The genus Sosioc rimites separated from Neocrimites shortly after its branching off in the Artinskian (S. darvasicus). Both these species have a general trend towards increased complexity due to the increased number of lobes, but differ in the accompanying shell shape (com bination of superimposed ontogeny of the suture while PALEONTOLOGICAL JOURNAL
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the shell shape deviated and ornamentation remained unchanged). In Sosiocrimites the number of lateral lobes in the course of evolution of ontogeny also increased to seven or eight, but the whorl width remained moderate (the shell shape changed from subspherical to pachy conic), whereas the external portion of the suture became arched. In the reticulate ornamentation like in Neocrimites, spiral ornamentation remained dominant. The genus Adrianites with the pachyconic shell, a relatively wide umbilicus, distinct reticulate ornamen tation and arched suture with seven lateral lobes also apparently related to Sosiocrimites. Adrianites inher ited from its ancestor a complex, multielement suture, the arched curvature of which in the region between the ventral lobe to the umbilical margin was a consequence of the change in the shell shape from subspheroconic to pachyconic. This phylogenetic branch was very conservative: the first representatives (Wordian) were morphologically similar to the latest (Amarassian). In another branch, the initial genus of which was Epadrianites, there were the following characters: shell shape from inflated to barrelshaped, moderately involute with a relatively wide umbilicus and spiral (closely spaced or widely spaced) ornamentation. The suture has four or five lobes on the external flank. These characters suggest that Epadrianites was sepa rated from Neocrimites at a relatively early stage by widening of the umbilicus, increased spiral elements
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of ornamentation and a slight change in the shape of the lobes. Species of Epadrianites during its entire time of existence maintained a moderately complex suture (deviations in the shell morphology and ornamenta tion and conservatism in the sutural ontogeny). In turn, Epadrianites could give rise to a peculiar genus Pseudagathiceras with a widely umbilicate, pachyconic shell, with spiral ornamentation. Some species (P. spinosum) have ornamentation with ventral spines (Fig. 2). Until now there is no certainty about the taxonomy of Ruzhencev’s (1950) genus Aricoceras with the sub genera Aricoceras, Neoaricoceras, and Metaricoceras. The first two are known from the Wordian of Sicily and Timor, and can probably, be considered as separate genera. The general shell morphology and arched suture they are more likely related to Sosiocrimites. The suture of Aricoceras is less complex than in Sosioc rimites, with four or five lobes on the flanks. In Neoar icoceras there are seven lobes on the flank. We prefer to assign the Timor species A. discoidale (Haniel, 1915), recognized by Ruzhencev as a separate subgenus Metaricoceras, to the genus Aricoceras until more comprehensive data are obtained. Subfamily Pamiritellinae Ruzhencev et Bogoslovskaya, 1978 The diagnosis of the subfamily is given in the previous studies (Leonova and Bogoslovskaya, 1990; Leonova, 2002). We assign to this genus six genera from the Kungurian–Wordian. Below we list the species com position of these genera. Genus Pamiritella Toumanskaya, 1963; only the type species P. vinogradovi Toumanskaya, 1963 from the Kungurian (Bolorian) of Pamir (Kochusu Forma tion). Genus Pamirioceras Pavlov, 1967; only the type species P. markovskii Pavlov, 1967 from the Kungurian (Bolorian) of Pamir (Kochusu Formation). Genus Palermites Toumanskaya, 1937. The type species P. distefanoi (Gemmellaro, 1887) from the Wordian of Sicily (Sosio Beds) and Kurdistan (Qulqula Formation); P. minor (Toumanskaya, 1931) from the Roadian of the Crimea (Burnian Beds); P. sp. from the Roadian (Kubergandian) of Pamir (Leonova, 2011); P. reticulatus Liang, 1982 from the Wordian of Northwestern China (Fanjiatun Formation). Genus Sizilites Toumanskaya, 1937. The type spe cies S. affinis (Gemmellaro, 1888) from the Wordian of Sicily (Sosio Beds) and Kurdistan (Qulqula Forma tion), and the Roadian of Crimea (Burnian Beds); S. craticulatus (Gemmellaro, 1888) from the Wordian of Sicily (Sosio Beds) and Kurdistan (Qulqula Forma tion); S. kurdistanensis Vaší cek et Kullmann, 1988 from the Wordian of Kurdistan (Qulqula Formation). Genus Doryceras Gemmellaro, 1887. The type species D. fimbriatum Gemmellaro, 1887 and D. stuck
enbergi Gemmellaro, 1888 from the Wordian of Sicily (Sosio Beds). Genus Lingzhouceras Sheng, 1988. The type spe cies L. ornatum Sheng, 1988 and L. galeatum Sheng, 1988 from the Kungurian–Roadian of Tibet (Urulun Formation). This compact (occurring in three stages) (Fig. 3) group of adrianitids is distinct from other groups in the shell shape. The major ontogenetic trend was the shell “uncoiling,” decrease in its width, increased complex ity of transverse ornamentation due to the develop ment of constrictions, fascicles of lamellae and umbil ical nodes. No consistent increase in sutural complex ity is observed in Pamiritellinae. Quite opposite—a regression is observed (decrease in the number of ele ments). For example, in the Kungurian Pamirioceras and Pamiritella there are five and six lateral lobes respectively, whereas the Wordian Palermites, Sizilites, and Doryceras have five, four, and three, respectively (Fig. 2). In contrast to other adrianitids, that inhabited the pelagial and were more or less incapable of active swimming, Pamiritellinae included species that can be assigned to the nektobenthic living form (Pamiritella, Pamirioceras), and also included “plankton 1” (Sizilites, Doryceras) (Barskov et al., 2008). The question of the group’s origin again brings us to the genus Crimites, the most likely ancestor of the first known representative of Pamiritellinae (Pamirio ceras). Apparently, in the Late Artinskian time, Crim ites, with a moderately wide shell and transverse orna mentation (e.g., C. pamiricus) gave rise to taxa with a wider umbilicus, narrowed venter and more complex suture. Such characters are more typical of the earliest known Pamiritellinae (Pamir species Pamirioceras and Pamiritella.) However no intermediate forms between these taxa and Crimites have yet been found. These can only be assumed, or it is necessary to suggest a salta tory evolution of these forms from Crimites as a result of the early ontogenetic change (archallaxis). It is cer tain that Pamirioceras and Pamiritella are genetically related. This is supported by the similar ventral lobe outline, not known in other adrianitids, and the large depth of the first external lateral lobe, making it differ ent from other lobes. Morphologically the genus Pamirioceras is close to Crimites, and the present state of knowledge suggests that this genus is the initial genus of the subfamily. Compared to Crimites, it has a slightly more widely opened umbilicus, whereas the shell and especially the venter became narrower, and the number of the lateral lobes increased to five. The very peculiar genus Pamiritella gave rise to an entire group of widely umbilicate taxa, with prominent transverse ornamentation and a simplified suture. Together with these taxa, the genus Pamiritella consti tutes the major phylogenetic lineage in the subfamily, the development of which was retrogressive. Pamiri tella has a small moderately involute shell, with rela tively narrow and low whorls and a comparatively complex suture with a relatively shallow ventral lobe PALEONTOLOGICAL JOURNAL
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and six lateral lobes. In this phylogenetic lineage the development proceeded toward the uncoiling of the shell, increase in transverse ornamentation, and sim plification of the suture, and terminated in the appear ance of three genera with signs of degradation. The genus Palermites has a shell with a relatively wide umbilicus and evolute whorls, with a distinct trans verse ornamentation (lamellae). The arched external portion of the suture is composed of only five lateral lobes. Pamiritella is apparently connected with the genera Sizilites, Doryceras, and Lingzhouceras. All these are characterized by a moderately evolute shell with narrow whorls and a simplified suture. Represen tatives of the genus Sizilites have four lateral lobes in the external, arched portion of the suture, whereas Doryceras has only three such lobes. This group possi bly includes the Tibetian endemic Lingzhouceras (the genus is not sufficiently described) with two lobes on the external side. CONCLUSIONS In general, the evolution of the subfamilies under consideration occurred in similar ways: the stage of early evolution after the establishment of the initial taxon was followed by a relatively short stage of pro gressive development and the group entered the diver sification stage with many mosaic taxa combining conservative and advanced characters. This was fol lowed by a decline phase with features of regressive development up to the formation of paedomorphs and a sharp decrease in diversity (Fig. 2). This agrees well with the patterns observed in the phylogeny of suprageneric taxa of Paleozoic ammonoids (Leonova, 2014, 2015). ACKNOWLEDGMENTS The paper is supported by the Program of the Pre sidium of the Russian Academy of Sciences “Evolu tion of the Organic World and Planetary Processes” (Subprogram 2). REFERENCES Armstrong, I.D., Dear, I.F., and Runnegar, B., Permian ammonoids from eastern Australia, J. Geol. Soc. Aust., 1967, vol. 14, no. 1, pp. 87–98. Barskov, I.S., Boiko, M.S., Konovalova, V.A., Leonova, T.B., and Nikolaeva, S.V., Cephalopods in the marine ecosys tems of the Paleozoic, Paleontol. J., 2008, vol. 42, no. 11, pp. 1168–1284. Blendinger, W., Furnish, B., and Glenister, B.F., Permian cephlopod limestones, Oman mountains: evidence for a Permian seaway along the northern margin of Gondwana, Palaeogeogr., Palaeoclimatol., Palaeoecol., 1992, vol. 93, nos. 1–2, pp. 13–20. Bogoslovskaya, M.F., Artinskie ammonoidei Srednego Urala (Artinskian Ammonoidea of the Middle Ural Mountains), PALEONTOLOGICAL JOURNAL
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Tr. Paleontol. Inst. Akad. Nauk SSSR, vol. 87, Moscow: Akad. Nauk SSSR, 1962. Cantú Chapa, A., Los cefalópodos del Paleozoico de Méx ico, Geociencias, instituto Politécnico Nacional, 1997, no. 1, pp. 1–130. Ehiro Masayuki and Araki Hideo, Permian cephalopods of Kurosawa, Kesennuma City in the Southern Kitakami Massif, northeast Japan, Paleontol. Res., 1997, vol. 1, no. 1, pp. 55–66. Furnish, W.M., Glenister, B.F., Kullmann, J., and Zhou Zuren, Treatise on Invertebrate Paleontology: Part L. Mol lusca 4. Revised, vol. 2: Carboniferous and Permian Ammonoidea (Goniatitida and Prolecanitida), Selden, P.A., Ed., Boulder, CO–Lawrence KS: Geol. Soc. Am.–Univ. Kansas Press, 2009, pp. 125–135. Gemmellaro, G.G., La fauna dei calcari con fusulina della valle del fiume Sosio nella provincia di Palermo, Giorn. Sci. Nat. Econ., 1887, vol. 19, pp. 1–106. Glenister, B.F. and Furnish, W.M., The Permian ammonoids of Australia, J. Paleontol., 1961, vol. 35, no. 4, pp. 673–736. Haniel, C.A., Die Cephalopoden der Dyas von Timor, Palä ontol. von Timor, vol. 3, no. 6, Stuttgart: E. Schweizerbart, Nägele und Dr. Sprosser, 1915. Leonova, T.B. and Dmitriev, V.Yu., Rannepermskie ammonoidei YugoVostochnogo Pamira (Early Permian Ammonoidea of the Southeastern Pamirs), Tr. Paleontol. Inst. Akad. Nauk SSSR, vol. 235, Moscow: Nauka, 1989. Leonova, T.B. and Bogoslovskaya, M.F., Phylogenetic rela tionships in the superfamily Adrianitaceae, in Iskopaemye tsefalopody (Fossil Cephalopoda), Moscow: Nauka, 1990, pp. 87–97. Leonova, T.B., Permian ammonoids: Classification and phylogeny, Paleontol. J., 2002, vol. 36, no. Suppl. 1, pp. 1– 114. Leonova, T.B., The Kubergandian (Middle Permian) ammonoid assemblage of the southeastern Pamirs (Tajiki stan), Byull. Mosk. Ova Ispyt. Prir., Otd. Geol., 2011, vol. 86, no. 3, pp. 21–31. Leonova, T.B., Heterochronies in the evolution of Paleo zoic Ammonoidea, Sb. mater. konf. “Morfogenez v individ ual’nom i istoricheskom razvitii: geterokhronii, geterotopii i allometriya”, Moskva, 16–18 aprelya 2014 g. (Abstracts of the Conference “Morphogenesis in Individual and Histori cal Development: Heterochronies, Heterotopias, and Allometry,” Moscow, April 16–18, 2014), Moscow: Pale ontol. Inst. Akad. Nauk SSSR, 2014, pp. 111–123. Leonova, T.B., Role of heterochronies in the the morpho genesis of Paleozoic ammonoids, Paleontol. J., 2015, vol. 49, no. 14 (in press). Liang Xiluo, Early Permian cephalopods from northwest ern Gansu and western Nei Mongol, Acta Palaeontol. Sin., 1981, vol. 20, no. 6, pp. 485–500. Liang Xiluo, Some Early Permian ammonoids from Jilin and Nei Mongol, Acta Palaeontol. Sin., 1982, vol. 21, no. 6, pp. 645–657. Mapes, R.H. and Boardman, D.R., The Upper Paleozoic (Pennsylvanian–Permian) ammonoid Emilites, J. Paleon tol., 1988, vol. 62, no. 1, pp. 69–75. Mezhdunarodnyi kodeks zoologicheskoi nomenklatury. Izd. chetvertoe (The International Code of Zoological Nomen clature: Fourth Edition), Moscow: KMK, 2004.
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Miller, A.K. and Furnish, W.M., Studies of Carboniferous ammonoids: parts 5–7, J. Paleontol., 1940a, vol. 14, no. 6, pp. 521–543. Miller, A.K. and Furnish, W.M., Permian ammonoids of the Guadalupe Mountain Region and Adjacent Areas, Spec. Pap. Geol. Soc. Am., no. 26, New York: Geol. Soc. Am., 1940b. Nassichuk, W.W. and Henderson, Ch.M., Lower Permian (Asselian) ammonoids and conodonts from the Belcher Channel Formation, southwestern Ellesmere Island, Curr. Res. Geol. Surv. Can., 1986, Pap. 861B, pp. 411–416. Ruzhencev, V.E., Verkhnekamennougol’nye ammonity Urala (Upper Carboniferous Ammonites of the Urals), Tr. Pale ontol. Inst. Akad. Nauk SSSR, vol. 29, Moscow: Akad. Nauk SSSR, 1950. Ruzhencev, V.E., Nizhnepermskie ammonity Urala. Kn. 2. Ammonity artinskogo yarusa (Lower Permian Ammonites of the Urals: 2. Ammonites of the Artinskian Stage), Orlov, Yu.A., Ed., Tr. Paleontol. Inst. Akad. Nauk SSSR, vol. 60, Moscow: Akad. Nauk SSSR, 1956. Ruzhencev, V.E., Late Carboniferous ammonoids of the Russian Platformy and Cisuralia, Paleontol. Zh., 1974, no. 3, pp. 32–46. Ruzhencev, V.E. and Bogoslovskaya, M.F., Namyurskii etap v evolyutsii ammonoidei. Pozdnenamyurskie ammonoidei (The Namurian Stage in the Evolution of Ammonoids: Late Namurian Ammonoids), Tr. Paleontol. Inst. Akad. Nauk SSSR, vol. 167, Moscow: Nauka, 1978. Schiappa, T., Spinosa, C., and Snyder, W., Nevadoceras, a new Early Permian adrianitid (Ammonoidea) from Nevada, J. Paleontol., 1995, vol. 69, no. 6, pp. 1073–1079.
Sheng Huaibin, The Early Permian ammonoid facies stratigraphy in Yarlung Zangbo area, Xizang (Tibet), Prof. Pap. Stratigr. Palaeontol., 1984, no. 17, pp. 129–142. Sheng Huaibin, Early Lower Permian ammonoids of the Urulung Formation from Luobadui, Xizang (Tibet), in Tec tonic Evolution of the Lithosphere of the Himalayas, Beijing: Geol. Publ. House, 1988, pp. 123–148. Shigeta Yasunari, Zakharov, Yu.D., and Mapes, R.H., Ori gin of the Ceratitida (Ammonoidea) inferred from the early internal shell features, Paleontol. Res., 2001, vol. 5, no. 3, pp. 201–213. Smith, J.P., Permian ammonoids of Timor, Jb. Mijnw. Nederl.Indie, 1927, vol. 1, pp. 1–58. Termier, H., Termier, G., Desparmet, R., and Montenat, Ch., Les ammonoides du Permien (Kubergandien) de Tezak (Afganistan central), Ann. Soc. Géol. Nord, 1972, vol. 92, no. 3, pp. 105–115. The International Code of Zoological Nomenclature: Fourth Edition London: International Trust for Zoological Nomenclature, 1999. Toumanskaya, O.G., Permokarbonovye otlozheniya Kryma. Cephalopoda (Permian–Carboniferous Sediments of the Crimean Peninsula: Cephalopoda), Leningrad: Geol. Izd. Gl. Geol.Razved. Upr., 1931. Toumanskaya, O.G., On representatives of a new genus, Crimites, from Permian sediments, Ezhegodn. Vseross. Pale ontol. Ova, 1937, vol. 11, pp. 146–147. Vašíc ek, Z. and Kullmann, J., Eine AmmonoideenFauna vom SosioTyp (Guadalupian, OberPerm) aus Kurdistan (Irak), Palaeontogr. A, 1988, vol. 204, nos. 1–3, pp. 95–115. ˆ
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