TREATISE ONLINE Number 111

Part M, Chapter 23C: Systematic Descriptions: Phragmoteuthida

Dirk Fuchs and Desmond Donovan 2018

Lawrence, Kansas, USA ISSN 2153-4012 paleo.ku.edu/treatiseonline

PART M, CHAPTER 23C: SYSTEMATIC DESCRIPTIONS: PHRAGMOTEUTHIDA Dirk Fuchs1 and Desmond Donovan2 [1SNSB-Bayerische Staatssammlung für Paläontologie und Geologie, Munich, Germany; 2Wells, Somerset, UK]

INTRODUCTION pro-ostracum consists of a central median field with parabolar growth increments and The Phragmoteuthida Mojsisovics, 1882, likewise parabolar lateral fields. A narrow is a small and rare belemnoid order that area with backwards (hyperbolar) curved includes one family with two unambiguous or dense longitudinal growth lines—vari- genera and four, supposed phragmoteuthid- ously called “hyperbolar zone,” “arcuated like forms. It is regarded as being morpho- zone,” “notch,” or “reentrant”—is usually logically and phylogenetically transitional intercalated between the median field and between the Paleozoic coleoids (Hematititda the lateral fields. This tripartite construc- Doguzhaeva, Mapes, & Mutvei, 2002; tion is well known from fossil gladii (see Donovaniconida Doguzhaeva, Mapes, & Treatise Online, Part M, chapter 9B), and it Mutvei, 2007; Aulacoceratida Stolley, is why many authorities agreed that Meso- 1919) and the Mesozoic Belemnitida Zittel, zoic gladius-bearing coleoids originated 1895. Mojsisovics (1882) first recognized within the Phragmoteuthida (e.g., Jeletzky, the uniqueness of the family Phragmoteu- 1966; Doyle, Donovan, & Nixon, 1994; thidae Mojsisovics, 1882, which Jeletzky Donovan, 2006; Fuchs, 2006a, 2006b; (1965) later elevated to ordinal rank. System- Fuchs & Weis, 2008; Schweigert & Fuchs, atics for this group are shaped by the charac- 2012). For this reason, the Phragmoteu- teristically wide and three-lobed (tripartite) thida appears to be the root of two signifi- pro-ostracum, which is significantly different cantly different evolutionary strategies: one from the spatulate pro-ostracum of the retaining a calcareous phragmocone (Belem- Belemnitida and Diplobelida Jeletzky, nitida) and one entirely losing its buoyancy 1965 (e.g., Jeletzky, 1966; Donovan, 2006; device through decalcification (Octobrachia Doguzhaeva & others, 2007). According to Haeckel, 1866). A monophyletical status for the common idea, a pro-ostracum developed the Phragmoteuthida is therefore difficult through gradual reduction of the ventral to establish. body chamber (see Treatise Online, Part M, The stratigraphical occurrence of the chapter 8A). The phragmoteuthid type of Phragmoteuthida started with certainty in pro-ostracum encompasses about three- the Middle Triassic (Anisian) with Brevi- quarters of the phragmocone circumference conoteuthis Rieber, 1973. Although the and, hence, represents an initial state of Late Permian Permoteuthis Rosenkrantz, pro-ostracum formation. The free space of 1946, has frequently been regarded as the the formerly tubular body chamber has most earliest phragmoteuthid, the single, frag- probably been replaced by muscular mantle, mentary specimen lacks clear evidence of a which is supposed to be attached to the tripartite pro-ostracum (Jeletzky, 1966). lateral margins of the (weakly mineralized) Permoteuthis will therefore be treated as pro-ostracum (Fuchs & others, 2016). The a putative phragmoteuthid (Fig. 1). The

© 2018, The University of Kansas, Paleontological Institute, ISSN (online) 2153-4012

Fuchs, Dirk, & Desmond Donovan. 2018. Part M, Chapter 23C: Systematic Descriptions: Phragmo- teuthida. Treatise Online 111:1–7, 4 fig. 2 Treatise Online, number 111

150 Tithonian Kimmeridgian Late Oxfordian Callovian Bathonian ddl e 170 Bajocian leriteuthis Mi Aalenian

Toarcian Wink JURASSIC y

Pliensbachian Phragmoteuthis

190 Earl Sinemurian Hettangian ? Rhaetian ? 210

Norian Late unzoteuthis onoteuthis 230 L Carnian TRIASSIC Zugmontites Ladinian Brevic

iddle Anisian ? M ermoteuthis

Olenekian P 250 Early Induan Changhsingian

opingia n Wuchiapingian L e Capitanian

270 Wordian putativ Phragmoteuthidae

Roadian Phragmoteuthida Guadalupian Kungurian PERMIAN Artiskian 290

Cisuralian Sakmarian Asselian

Fig. 1. Stratigraphical occurrences of phragmoteuthid taxa.

Early Jurassic “Phragmoteuthis” montefiorei 2013, are confirmed, the order would have (Buckman, 1880) (Callovian) and “Phrag- disappeared by the end of the Jurassic. moteuthis” conocauda (Quenstedt, 1849 The assumption of ten-armed Phrag- in 1845–1849) (Toarcian) were known moteuthida was exclusively based on the for a long time as the youngest occur- well-preserved, hook-bearing arm crowns rences of phragmoteuthids. However, of Clarkeiteuthis (“Phragmoteuthis”) monte- both species have been reassigned to the fiorei and Clarkeiteuthis (“Phragmoteuthis”) (putatively diplobelid) genus Clarkeiteuthis conocauda. The reassignment of the latter Fuchs, Donovan, & Keupp, 2013, owing species strongly influenced our knowl- to the presence of a distinctly narrow pro- edge: a complete arm crown and, hence, ostracum. Therefore, the Sinemurian Phrag- the true number of arms is still unknown moteuthis huxleyi from the UK remains from unambiguous phragmoteuthids. The the only unambiguous, post-Triassic only complete arm crown consisting of phragmoteuthid (Donovan, 2006). If ten arms was recorded from a specimen phragmoteuthid affinities of the Tithonian determined as ?Phragmoteuthis ticinensis Winkleriteuthis Fuchs, Heyng, & Keupp, (Anisian, Monte San Giorgio), but it still Systematic Descriptions: Phragmoteuthida 3 lacks confirmation of a tripartite pro- hooks, ×2 (new; photo by H. Summesberger).—— ostracum (Rieber, 1970). Fig. 2,1c–f, P. huxleyi Donovan, upper Sinemurian, Charmouth Mudstone Formation, Dorset, UK; c, paratype, BMNH 83963, ventral view, ×0.4 Order PHRAGMOTEUTHIDA (Donovan, 2006, fig. 4A);d, 3-D reconstruction, Mojsisovics, 1882 ventral view (new); e, paratype, BMNH 83963, dorsolateral view, ×0.4 (Donovan, 2006, fig. 4B); [nom. transl. Jeletzky in Sweet, 1964, p. 12, ex Phragmoteuthidae Mojsisovics, 1882 p. 304; nom. correct. Jeletzky, 1965, p. 73, pro Phrag- f, 3-D reconstruction, dorsal view (new). moteuthidida Jeletzky in Sweet, 1964, p. 12] [=Phragmoteuthidoidea Breviconoteuthis Rieber, 1973, p. 7 [*Atractites brevi- Starobogatov, 1983, p. 7] conus Reis, 1907, p. 148; M]. Phragmocone very small to small, brevi- to slightly cyrtoconic, circular Phragmocone ortho-, brevi-, or slightly in cross section, apical angle 38°–40°; ratio of cyrtoconic, very small to medium-sized; chamber length to diameter about 0.15; pro- number of chambers moderate to low, ostracum three-lobed, longer than phragmocone, chamber length short; mural parts of septa anterior median field rounded; siphuncle marginal; rostrum unknown. Middle Triassic (upper Anisian): 30 to 50 percent of chamber length; pro- Switzerland (Canton Ticino), Austria.——Fig. ostracum weakly mineralized, wide, encom- 2,2a–b. *B. breviconus (Reis), upper Anisian, Grenz- passing three-quarters of phragmocone bitumenzone, Monte San Giorgio, Switzerland; a, circumference, three-lobed, subdivided into specimen PIZ M/21, dorsal view showing three- lobed pro-ostracum, ×1.5; b, specimen PIZ M/12, median field, hyperbolar zones, and lateral dorsal view showing breviconic phragmocone, ×1.5 fields; lateral fields slightly shorter than (new; originals of Rieber, 1974, fig. 1–4). median field; protoconch unknown; rostrum aragonitic, investment-like; number of arms PUTATIVE PHRAGMOTEUTHIDA uncertain; arm hooks present, possibly bise- Lunzoteuthis Doguzhaeva, Summesberger, & Mutvei, rial, shape variable. Middle Triassic (upper 2006, p. 71 [*L. schindelbergensis; OD]. Phragmo- Anisian)–Lower Jurassic (Sinemurian): cone very small, breviconic, ratio of chamber length to diameter about 0.33; pro-ostracum with northern Italy, Austria, Switzerland, UK, median field and arcuated zones. Upper Triassic ?Germany, ?Vietnam. (lower Carnian): Austria.——Fig. 3,1. *L. schin- delbergensis, lower Carnian, Schindelberg, Lunz, Family PHRAGMOTEUTHIDAE Austria, holotype, NHMW2005z0005/1, showing Mojsisovics, 1882 dorsal fragments of phragmocone with anteriorly projecting growth lirae of the median field cf( ), [Phragmoteuthidae Mojsisovics, 1882 p. 304] [=Phragmoteuthidae Naef, ×7 (Doguzhaeva, Summesberger, & Mutvei, 2006, 1921a, p. 534; =Phragmoteuthididae Jeletzky, 1966, p. 37] pl. 1A). Phragmoteuthis Mojsisovics, 1882, p. 304 [*Belemno- Permoteuthis Rosenkrantz, 1946, p. 161 [*P. groen- teuthis bisinuata Bronn, 1859, p. 44; OD]. Phrag- landica; M]. Known only from a small piece of shell mocone medium sized, orthoconic, apical angle that appears to represent a fragment of a three- 25°–30°; chamber length short; sutures simple; lobed pro-ostracum. [The name Permoteuthis was septal necks orthochoanitic, connecting rings applied by Rosenkrantz (1946) to several fossils extremely long and superimposing 5–6 previous that may or may not belong to the same species. It rings; siphuncle marginal; pro-ostracum length was restricted by Jeletzky (1966, p. 38) by designa- equals phragmocone length, anterior median field tion of the pro-ostracum fragment.] Upper Permian weakly acute; lateral field slightly shorter than (Changhsingian): eastern Greenland.——Fig. 3,2. median field, relative length variable; hooks vari- *P. groenlandica; line drawing of a putative, frag- able, stylet-like or with strongly curved uncinus. mentary pro-ostracum, ×0.5 (new). ?Middle Triassic (upper Anisian), Upper Triassic Winkleriteuthis Fuchs, Heyng, & Keupp, 2013, p. 242 (lower Carnian)–Lower Jurassic (upper Sinemu- [*Acanthoteuthis problematica Naef, 1922, p. 183; rian): northern Italy, Austria, UK, Switzerland, OD]. Phragmocone small, breviconic, otherwise ?Germany, ?Vietnam.——Fig. 2,1a–b. *P. bisinuata poorly known; lateral fields of the pro-ostracum (Bronn), Carnian, Cave de Predil (Raibl), northern well developed and anteriorly (parabolar) projected; Italy; a, specimen NHMW 1864/52/47 (original onychites uniform, biserial, base short, shaft slightly of Mojsisovics, 1882, pl. 94,3), showing later- inclined, uncinus weakly curved. Upper Jurassic ally compacted, three-lobed pro-ostracum, ×1.6 (lower Tithonian): southern Germany.——Fig. (Doguzhaeva & others 2007; photo by H. Summes- 3,3a–e. *W. problematica (Naef), lower Tithonian, berger); b, specimen showing differentiation of arm Solnhofen Formation, Eichstätt region, Germany; 4 Treatise Online, number 111

1c 1d

1b 1a 1e 1f1f Phragmoteuthis

Breviconoteuthis

2a 2b

Fig. 2. Family Phragmoteuthidae (p. 3). Systematic Descriptions: Phragmoteuthida 5

1 Lunzoteuthis 2 Permoteuthis

Winkleriteuthis

3e

3a 3b 3c 3d

Fig. 3. Putative Phragmoteuthida (p. 3–5).

a–b, paratype, BSPG MC-22; a, entire specimen, REFERENCES ×1.2; b, close-up view of hooks, ×16 (Fuchs, Heyng, & Keupp, 2013, fig. 2);c–d, reconstruction of the Bronn, H. G. 1859. Nachtrag über die Trias-Fauna von shell, ventral (c) and dorsal (d ) views (new); e, 3-D Raibl. Neues Jahrbuch für Mineralogie, Geologie, reconstruction of a hook (new). und Paläontologie 1859:39–45. Zugmontites Reis, 1907, p. 148 [*Z. mojsisovicsi Reis, Buckman, J. 1880. On the Belemnoteuthis Montefiorei. 1907, p. 148; SD Bülow-Trummer, 1920, p. 75]. Proceedings of the Dorset Natural History and An- Phragmocone very small to small, cyrtoconic, laterally tiquarian Field Club 3:141–143. compressed; apical angle about 38°; ratio of chamber Bülow-Trummer, E. von. 1920. Cephalopoda Dibran- length to diameter about 0.30; siphuncle marginal; pro- chiata. Fossilium Catalogus, 1: Animalia, Pars 11. W. ostracum unknown; rostrum unknown. Middle Triassic Junk. Berlin. 313 p. (upper Anisian): Austria.——Fig. 4a–c. *Z. mojsisovicsi, Doguzhaeva, L. A., R. H. Mapes, & Harry Mutvei. 2002. upper Anisian, Wettersteinkalk, Ehrwald, Austria, Shell morphology and ultrastructure of the Early holotype, BSPG 1901 II 508; ventral (a), lateral (b), Carboniferous coleoid Hematites Flower & Gordon and dorsal (c) views, ×2 (new; photo by A. Nützel). 1959 (Hematitida ord. nov.) from the Midcontinent 6 Treatise Online, number 111

a b c abZugmontites c

Fig. 4. Putative Phragmoteuthida (p. 5).

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