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Fossil Record 13 (1) 2010, 5 –12 / DOI 10.1002/mmng.

200900008

A key for the description of Palaeozoic ammonoids
Dieter Korn
Museum fr Naturkunde Berlin, Invalidenstraße 43, 10115 Berlin, Germany. E-mail: dieter.korn@mfn-berlin.de

Abstract
Received 4 June 2009 A key for the description of Palaeozoic ammonoids is presented. It contains a catalo-
Accepted 12 September 2009 gue of descriptive terms to characterise conch morphology, ornament, and suture line
Published 19 February 2010 particularly of Carboniferous ammonoids. A number of examples of tables and illustra-
tions are given for the description of conch characters, their ontogeny and intraspecific
variability.
Key Words
Ammonoidea
Carboniferous
morphology
ontogeny
taxonomy

Introduction the conch width index and umbilical width index and
separated between 35 possible morphs. He named these
Palaeozoic ammonoids have been described for almost morphs either according to their shape (e.g. ‘sferokon’,
200 years. Since the first nomination of the Carbonifer- ‘kadikon’) or after ammonoid genera (e.g. ‘oppellikon’,
ous ammonoid “Ammonites Listeri” by Sowerby (1812), ‘daktilikon’); name pairs such as ‘ofikon’ (for very
styles and contents of descriptions have significantly slender forms) and ‘subofikon’ (for moderately slender
been modified and completed. The numbers of conch forms) lead to a reduction of descriptive terms.
characteristics taken into account have increased consid- The description key outlined here can be seen as a
erably, and external as well as internal characters were further development of the schemes proposed by Ruz-
evaluated to form a balance. While the early authors hencev and Kutygin. This key is specifically coined for
mainly described more or less mature stages of single in- the Early Carboniferous ammonoids from Algeria,
dividuals, later authors also included the ontogeny of which will be described in monographs of this volume
specimens and or the variability within populations. (Bockwinkel et al. 2010; Ebbighausen et al. 2010; Korn
Although there is a sort of consensus among ammo- et al. 2010a, 2010b, 2010c); it was developed mainly
noid workers about the use of the various conch charac- for Carboniferous ammonoids of the predominant sub-
ters and their importance for ammonoid description and order Goniatitina, which has a wide morphological
classification, there is no agreement on how species range but very similar suture lines.
should be characterised in terms of diagnoses and illus-
trations. It was particularly Ruzhencev (e.g. 1956, 1960),
who in a series of monographs, developed a standard for
Conch characteristics
ammonoid descriptions accompanied by illustrations of
Conch geometry
representative specimens on plates, well-drawn suture
lines of mature and often of juvenile stages, and occa- Descriptive terms of the conch characters are mainly
sionally also conch cross sections. His diagnoses and de- adopted, with some modifications, from Korn & Klug
scriptions of species followed a strict scheme, in which (2003, 2007), who outlined the morphological spectrum
all the important characters were outlined. for Devonian ammonoids. Cross-sections have the highest
Based on the study of Permian forms, Kutygin potential for providing a large data set for morphometric
(1998) has proposed a scheme for the description of analysis of ammonoid conchs. With a reasonable number
ammonoid conchs. In a bivariate diagram he showed of cross-sections, ontogenetic development as well as in-

# 2010 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim

85 globular 0.de # 2010 WILEY-VCH Verlag GmbH & Co. see Fig. Classification of the general conch shape (conch width index. 1): Using these basic parameters.10 zoic ammonoids. Descriptive terms for conch morphology of Palaeo. Classification of the maximum conch diameters of individual specimens. ww1 and ww2 as above) cies or complete assemblages. Bivariate plot illustrating the terminology of the conch width index (ww/dm) on the x-axis and umbilical width index (uw/dm) on the y-axis.85 –1. traspecific variability can be analysed for a number of spe. ww/dm). D. descriptive term maximum diameter very small <25 mm Small 25 – 50 mm moderate 50 – 100 mm Large 100– 200 mm very large > 200 mm Table 2. secondary parameters can easily be computed:  conch diameter (dm. dm1 is the largest diameter in a cross section and dm2 is the diameter exactly half a  umbilical width (uw) ¼ dm1 –– wh1 –– wh2 whorl earlier)  aperture height (ah) ¼ dm1 –– dm2 museum-fossilrecord. Figure 2.: Key for description of Palaeozoic ammonoids Table 1.35 discoidal 0.35 –0. wh1 and wh2 as above) the cross-sections.6 Korn. KGaA. spindle-shaped > 1. descriptive term ww/dm extremely discoidal < 0.60 –0.wiley-vch. Weinheim .10 Figure 1.60 pachyconic 0. 2. three of the basic conch parameters can immediately be obtained for each half volution (Fig. From computer drawings of  whorl height (wh.  whorl width (ww.

# 2010 WILEY-VCH Verlag GmbH & Co.45 –0.15 –0.50 (whorl width index. (dm1 –– wh1 –– wh2)/dm1  whorl expansion rate (WER) ¼ (dm1/dm2)2 or [dm1/(dm1 –– ah)]2 Table 5.50 –3.30 –0. angular. ww/wh).00 –2. 3.00 strongly embracing 0.30 –0.75 rowly rounded. nar.00 –1.45 (conch subevolute)  whorl width index (WWI) ¼ ww1/wh1 wide 0. IZR).50 –1.wiley-vch. see Fig. strongly compressed < 0.00 descriptive term IZR weakly depressed 1. see Fig.45 Figure 3.50 moderately depressed 1.15 (conch involute) narrow 0.15 –0.50 wall can be thus described as broadly rounded.60 (conch very evolute). the arrangement and the shape of the flanks.Fossil Record 13 (1) 2010.30 (conch subinvolute)  conch width index (CWI) ¼ ww1/dm1 moderate 0. 4). Classification of the coiling rate of the conch  imprint zone rate (IZR) ¼ wh1  ah/wh1 or (whorl expansion rate. low 1.00 high 2.15 strongly depressed 2. Classification of the whorl cross section shape extremely high > 2.00 very strongly embracing > 0.00 –2. 5 –12 7  imprint zone width (iz) ¼ wh1 –– ah or Table 4.45 extremely depressed > 3. moderate 1. KGaA. 3.75 –2. Classification of the umbilical width (umbilical wh1 –– (dm1 –– dm2) width index.50 –2. Classification of the whorl overlap rate (imprint zone rate.de . 2.50 –1. descriptive term ww/wh Table 6. subangular. WER). Conch proportions and expansion rates (growth rates) descriptive term uw/wh were calculated in the following way by using the three basic conch parameters: very narrow < 0.60 (conch evolute)  umbilical width index (UWI) ¼ uw/dm1 or very wide > 0.50 moderately embracing 0. see Fig. uw/dm). Weinheim museum-fossilrecord. the umbilical margin and the umbilical very low < 1. Bivariate plot illustrating the terminology of the imprint zone rate (IZR) on the x-axis and whorl width in- dex (ww/wh) on the y-axis.50 Table 3.20 very high 2.30 very strongly depressed 2. and acute (Fig.25 –2. –– (wh1 (dm1 –– dm2))/wh1 descriptive term WER The shape of the venter.00 weakly embracing < 0.50 weakly compressed 0.

umbilical margin sub- angular. they can be smooth.de # 2010 WILEY-VCH Verlag GmbH & Co. venter broadly rounded. Conch thinly pachyconic. flattened. 5). from being little eral as well as a ventral sinus more than strengthened growth lines to prominent ra- museum-fossilrecord. a projection at the boundary between flank counted from umbilicus to umbilicus. flanks con- verging. Shell ornament and steinkern Table 7. flanks slightly converging. G. Conch thinly globular. subevolute. umbili- cal margin subangular. ranging from extremely fine to linear without remarkable projections and sinuses coarse with different spacing.: Key for description of Palaeozoic ammonoids Figure 4. venter broadly rounded. flanks subparallel (conch widest in the midflank area). subinvolute. ribs. involute. D. with bifur- biconvex with two projections on the flank (dorsolat- cation and/or with intercalation of secondary ribs). I. umbilical wall convex. flanks subparallel. the three ornament ele- rursiradiate with a general backward direction ments can be characterised by a uniform terminology. venter very broadly rounded. subevolute. venter broadly rounded. flanks strongly converging. umbilical wall flattened. is often a criter- and venter (ventrolateral projection). Conch thickly pachyconic. Conch thickly discoidal. Strong crenulation may result in a (lateral projection) and a ventral sinus spiral ornament. umbilical wall rounded. umbilical margin rounded. A. Therefore. umbilical margin rounded. Conch thinly pachyconic. D. umbilical margin rounded. from fine to coarse. venter rounded.8 Korn. in which the spiral lines can range concavo-convex with a sinus on the midflank (lateral sinus). Weinheim . umbilical margin rounded. in their strength. eral and ventrolateral projection) and a lat- They can range. E. Conch thickly pachyconic. and constrictions can firstly be char- acterised by their direction and course across flanks descriptive term characteristics and venter (Tab. ventral sinus Ribs may be simple or more complex (i. um- bilical wall convex. evolute. involute. Classification of the direction and course of the growth lines. convex with only one wide projection on the flank rough or crenulated. umbilical margin rounded. flanks strongly converging. venter narrowly rounded. umbilical margin subangular. B. subevolute. C. J. Growth lines. Conch thickly discoidal. F. and constrictions (Fig. subevolute. Conch thickly discoidal. umbilical wall flat. The number of spiral lines. umbilical margin subangular. umbilical wall flattened. venter broadly rounded. venter rounded. ribs. Conch thinly discoidal.e. H. venter very broadly rounded. rectiradiate with a general radial direction Growth lines can occur in a wide range of strengths prorsiradiate with a general forward direction and consistencies. umbilical wall convex. flanks converging (conch widest near the umbilicus).wiley-vch. umbilical wall oblique. umbilical wall convex. flanks strongly converging. involute. umbilical wall flattened. KGaA. involute. flanks subparallel. and a ion for the separation of species. umbilical margin subangular. venter narrowly rounded. 7). Various shapes of ammonoid conchs. Conch thinly discoidal.

D. B. Biconvex with low dorsolateral projection. with two kern constrictions) or on the shell surface as well as the strongly different flanks). acute. G. Suture line Sutural terminology (Fig. Convex with shallow ventral sinus. E. Its general shape can Rare cases of a midventral groove are known particu. Prolecanitid am- 1988). I. scribed as grooves. Biconvex with high dorsolateral projection. Concavo-convex with low ventrolateral projection and deep ventral sinus (direction rectiradiate). Constrictions can occur on the internal mould when The general shape of the ventrolateral saddle can they are caused by internal shell thickenings (= stein. can be described (1) by the ratio of the width (at half depth) of the external Figure 6. (2003):  external lobe (E lobe). dial sculptures. Biconvex with high dorsolateral projection. Their direction broadly rounded to narrowly rounded.de . Weinheim museum-fossilrecord.  lateral lobe (L lobe). Concavo-convex with very low ventrolateral projection and very low ventral sinus (direction rectiradiate). low ventro- lateral projection and deep ventral sinus (direction rursiradiate). Their direction and course is described depth) and total depth of the external lobe (EL/h as as in the growth lines. Convex with extremely shallow ventral sinus.  umbilical lobe (U lobe). A. straight. J. Goniatitid ammonoid. Descriptive terms for sutural morphology of Palaeo- lobe and the adventive lobe (E/A as used by Korn zoic ammonoids. low ventrolateral projection and shallow ventral sinus (direction rectiradiate). between adventive lobe and lateral lobe). L.  internal lobe (I lobe).g.  dorsolateral saddle (A/L saddle. (deeper than the external lobe). K. used by Korn 1997 or EL w/d as used here). 5 –12 9 Figure 5.e. larly from the Prolecanitida.  adventive lobe (A lobe). moderately deep the Goniatitina. in some groups within not as deep as the external lobe). B. in the ventrolateral area and are visible (about the same depth as the external lobe) or deep on the internal mould as well as on the shell surface. Concavo-convex with high ventrolateral projection and deep ventral sinus (di- rection prorsiradiate). KGaA. The adventive lobe can be described as shallow (i. subacute or and course is described as in the growth lines. it can range from being internal mould (= shell constrictions). and for the Prolecanitida according to Korn et al. Con- cavo-convex with high ventrolateral projection and very shallow ventral sinus (direction prorsiradiate). # 2010 WILEY-VCH Verlag GmbH & Co. high ventrolateral projection and deep ventral sinus (direction rectiradiate). Biconvex with low dorsolateral projection. They occur. Linear. or (2) by the ratio of width (measured at half monoid. be symmetric or asymmetric with incurved (concave). A. high ventrolateral projection and deep ventral sinus (direction prorsiradiate). or convex flanks.Fossil Record 13 (1) 2010.wiley-vch.  median saddle (M saddle). Longitudinal (spirally arranged) constrictions are de. The width of the external lobe.  ventrolateral saddle (E/A saddle. Convex with deep ventral sinus. Various modes of constrictions. either be symmetric or asymmetric (e. which is an important criterion for the separation of species and genera parti- cularly within the Goniatitina. F. C. 6) for the Goniatitida is ac- cording to Wedekind (1918). H. between external lobe and adventive lobe).

35 Y-shaped with flanks parallel in the lower part.50 width of the adventive lobe. pays attention to characters of conch morphology and acters but much more frequently by a combination of ontogeny. calculated by its width in narrow 0. 13).75 extremely wide > 1. B. Weinheim . KGaA. Shape of the external lobe as an important criterion for distinguishing major ammonoid taxa. museum-fossilrecord. Whorl expansion rate (WER) of all available specimens of a distinct species.00– 1. Descriptions of ammonoid species quantitative characters.de # 2010 WILEY-VCH Verlag GmbH & Co.wiley-vch. Table 10. A.75 Table 8. moderately narrow 0. Classification of the width of the external lobe (EL further characteristics are demonstrated.25 wide 1. moderately low 0. calculated by its width descriptive term EL w/d in comparison with the external lobe (EL/VLS) or its width/height proportion (VLS w/h). and (2) the relative very narrow < 0.25 – 1. Example of an illustration of the ontogenetic development of the conch in ammonoids. D.20 – 0.00 moderately wide 1. Therefore. Classification of the height of the median saddle descriptive term characteristics (MS h).50 – 1. ornament.80 Figure 7.50 – 0. C.80 but narrower in the upper part. very high > 0. rectangular with largely parallel and almost uncurved flanks descriptive term MS h pouched or inflated with more or less parallel and convexly curved flanks very low < 0.50 very wide 1.20 V-shaped with diverging and almost straight flanks low 0. width of the ventrolateral saddle. Table 9.75 – 1. D.10 Korn. Umbilical width index (uw/dm).65 bottle-shaped with flanks parallel in the lower part. diagnoses must often be arranged in a matrix-like matter.65 – 0.75 comparison with the external lobe (EL/AL). Whorl width index (ww/wh).: Key for description of Palaeozoic ammonoids In the tables with the suture line proportions (Tab. the diagnosis characterised by a single or by a few apomorphic char. high 0. (1) the relative w/d). Usually.35 – 0.50 but diverging in the upper part moderately high 0. composed of a set Diagnosis of characters that allow the comparison between the Most of the Carboniferous ammonoid species are not species within distinct genera. and suture line.50 – 0. Conch width index (ww/dm).

e.85.38) (ww/wh ¼ 2.8 23.8 3.90 – 2. whorl width index and the imprint zone rate on the y-axis (Fig.84 0.45 – 0.55 – 1. IZR ¼ 0. In these.45) (WER ¼ 2.15) Table 12.51 0.47 paratype MB.7 12. uw/dm ¼ 0. Korn & Bockwinkel.75) 10 mm thickly pachyconic. subinvolute moderately depressed. KGaA.60 – 2.75 – 0.50. IZR ¼ 0. characters.10 –0.40 – 0.1 0.91 0.50 – 0.5 26.80 – 1.10) 60 mm thinly to thickly discoidal.10) (ww/wh ¼ 0.02 0. the conch diameter is logarithmically represented on the presentative specimens.09 2.40) (WER ¼ 1. uw/dm ¼ 0.C.6 18. Traditionally.6 0.46 holotype MB. IZR ¼ 0.Fossil Record 13 (1) 2010.8 3. 3). strongly embracing high (ww/dm ¼ 0. The first of these dia- stages is much more informative.C. dm ww wh uw ah ww/dm ww/wh uw/dm WER IZR paratype MB.75 – 0.08 2.40 – 0.75) 30 mm thickly discoidal.35 – 0.de .1 27. IZR ¼ 0.45) (WER ¼ 1.C.60.9 3. describing the conch morphology and its variation in four distinctive growth stages.05 –0.42 # 2010 WILEY-VCH Verlag GmbH & Co.47 0.50.2 51. Weinheim museum-fossilrecord. uw/dm ¼ 0. bivariate plots for distinct netic trajectories and intraspecific variability as well as pairs of characters were developed. conch width index (ww/ second illustrates the whorl width index on the x-axis dm).30 –0.3 15.56 1. Example of a table to illustrate the conch dimensions and proportions for reference specimens of a Carboniferous ammonoid species. specimens and rich representation by cross sections These diagrams allow only the illustration of single showing the conch ontogeny. In these.18610.25.wiley-vch.20) (ww/wh ¼ 1.00 0.7 32. strongly embracing moderate to high (ww/dm ¼ 0. Example of a bivariate plot of the ontogenetic trajectories of an Early Carboniferous ammonoid (Hammatocyclus pollex Ebbighausen.8 3.03 0.00.20 –0.00 – 2.5 20.44 paratype MB.00. The the main conch characters i.06 2.12 38.1 45.0 27. 2).51 0. umbilical width index (uw/dm). In cases with a large number of x-axis and the ratio or expansion rate is on the y-axis.94 0.7 21.50) (WER ¼ 1. dm conch shape whorl cross section shape aperture 4 mm thickly pachyconic.12 0. Example of a table to illustrate the conch ontogeny of a Carboniferous ammonoid species.18610. very strongly embracing low (ww/dm ¼ 0. strongly embracing low (ww/dm ¼ 0. grams shows the conch width index on the x-axis and Ontogenetic trajectories and intraspecific variability of the umbilical width index on the y-axis (Fig.80. involute to subinvolute weakly depressed.06 2.10 0. 5 –12 11 Figure 8. subevolute strongly depressed. descriptions are based on a number of re. and therefore.5 13. two related tabular depiction with the focus on selected growth characters can be visualised.C. and whorl expansion rate (WER) can be visua- lised in simple semi-logarithmic plots (Fig.55 – 1.00 – 2. Description (ww/wh).10 59. involute weakly compressed.00 – 1.5 25. uw/dm ¼ 0.1 0. 2010). 7).45 – 0. the illustration of ontoge.18610.18610.3 0. The Table 11.30) (ww/wh ¼ 1.

Die Goniatiten des Kulmplattenkalkes (Cephalopoda.10 54.C.7 mm 0. In the monographs published parallel to this account Korn. sistema i filogeniya pa- Early Visan. and Mode of Life of the Frasnian (Late De- vonian) Ammonoid Manticoceras from Coumiac (Montagne diameters of the specimens by large dots (Fig. Mineral Conchology of Great Britain. (Potsdam) for proofreading the manuscript and Christian Klug (Z. Ebbighausen et al.. Printsipy sistematiki. 8). C.wiley-vch. The A- (Bockwinkel et al. V. Dordrecht: pp. D.52 0. Springer. Using Noire. Korn mode sutural ontogeny in prolecanitid ammonoids. The Palaeozoic ammonoids of the South Portuguese Wedekind. – Fossil Record 13 (1): 153–202. D. 2003.. The ammonoids Instituta Akademiya Nauk SSSR 6: 1–275.41 0. – Paleontologicheskiy Zhurnal 1998 (1): Bockwinkel. me I. Gourara. J.39 0. R. V. Gourara.79 A lobe asymmetric ontogenetic stage is illustrated by the size of the dots. 1960. ontogenetic traits can be illustrated and (eds).12 Korn. E.18610. from the Dalle  Merocanites of Timimoun (Late Tournaisian– Ruzhencev. Morphological pathways in the evolution nitidae. In Landman.. – Trudy Paleontologicheskogo Ebbighausen. [EL w/d ¼ width of the external lobe in relation to its depth. – Fossil Record 13 (1): 13– 34.18610. & Klug. J. & Klug. & Bockwinkel. measured at half depth. J. Mor- with small stages represented by small dots and large phological Disparity. C. D. Algeria). J. of Early and Middle Devonian ammonoids. Korn.00 0. V. Weinheim . 2010b. References Kutygin.5 mm 0. spectives. R. Conch Form Analysis. Glyphioceratidae und Proleca- Korn.C. R. D.79 0. Cephalopods Present and Past: New Insights and Fresh Per- help for the separation of species. I also thank Sonny Walton 214. Ammonoids from Acknowledgements the Dalle des Iridet of the Mouydir and Ahnet (Central Sahara) and the Formation d’Hassi Sguilma of the Saoura Valley (Late I am indebted to Jrgen Bockwinkel and Volker Ebbighausen in parti. D. Unterkarbon. a tabular description of ogy 46 (6): 1123–1132. D. specimen at dm EL w/d EL/VLS EL/AL MS h VLS w/h remarks paratype MB. 2010a. D. & Bockwinkel. II. Gourara. 1956. KGaA.. VLS w/h ¼ width of the ventrolateral saddle in relation to its height. 1998. Korn. D. from the Argiles de Timimoun of Timimoun (Early and Middle Ruzhencev. 2010. The ammonoids from the Grs du Kahla suprieur of Timimoun (Middle–early 13) was given priority over narrative descriptions. 2010c. Mit Ausschluß der Mimoceratidae. N.18610. 2010b. Ebbighausen. – Geo. D. these diagrams. 1812–1814. 2010c). V. H. J.50 1. V. Zone. E. Ammonity Artinskogo yarusa. Davis.: Key for description of Palaeozoic ammonoids Table 13. 2010a. H. 57–85. volu- logie und Palontologie in Westfalen 11: 1–293. Algeria). Algeria). Nizhnepermskiye ammonity Yuzhnogo Urala. 2010. Bockwinkel. Korn. Ammonoidea.. The ammonoids 20–31. & Klug. – Fossil Record 13 (1): 203– cular for the discussion of the manuscript. V. Rheinisches Schiefergebirge).8 mm 0. – Palaeontol- et al. EL/VLS ¼ width of the external lobe in relation to the ventrolateral saddle. Variability.. D.89 A lobe with slightly curved flanks paratype MB. & Ebbighausen.4 32.1 44. V. V.85 A lobe with almost straight flanks holotype MB. 1997. 2003. – Memrias do Instituto Geolgico e Mineiro 33: 1–131. measured at half depth]. – Paleobiology 29: 329–348. France).18 0.. – Fossil Record 13 (1): 35–152. Ebbighausen. V.91 0.24 0. Algeria). Korn.21 0. & Mapes. A. Korn.de # 2010 WILEY-VCH Verlag GmbH & Co. R. – Fossil Record 13 (1): 215–278. D. biometric characters of conch and suture line (Tabs 11– Korn. Sowerby. MS h ¼ height of the median saddle in relation to the depth of the external lobe. – Palaeontographica 62: 85–184. & Ebbighausen. 1988. Die Genera der Palaeoammonoidea (Goniatiten). Ebbighausen.. Late Tournaisian. J. Korn. 2010. The ammo- rich) as well as an anonymous reviewer for a review of the article. – Trudy Paleontologicheskogo Instituta Korn.43 0. Mouydir. EL/AL ¼ width of the external lobe in relation to the adventive lobe. J. C. 234 pp. Example of a table to illustrate the suture line proportions and characteristics of a Carboniferous ammonoid spe- cies. Forma rakovin permskikh ammonoidei se- vero-vostoka Rossii.42 0. 1918. 2010. noids from the Argiles de Teguentour of Oued Temertasset (early Late Tournaisian. leozoyskikh ammonoidey. museum-fossilrecord. & Bockwinkel. 2007. Akademiya Nauk SSSR 133: 1–331. Bockwinkel.C. Tournaisian–Early Visan. Visan. Algeria).