692


Journal of Paleontology 91(4):685–714


Table 1. Discrete morphological characters and related character states used for the cladistic analysis. The third column refers to specimens illustrated in Figure 5.


Characters


Thecal size (height) Number of circlets


constituting the theca States


Small theca<4.74cm Large theca ≥4.74cm


6 circlets 7 circlets 8 circlets 9 circlets 10 circlets >10 circlets


Number of radial circlets Additional plates between


infralaterals and laterals


Additional plates between laterals and radials


Rows of additional plates between laterals


Radials shape Type of ornamentation


≤3 circlets 4 circlets


Absent Absent Isolated or incomplete circlet


One complete circlet (with or without 2nd incomplete circlet) Two complete circlets or more


Isolated or incomplete circlet Complete circlet


Absent Present


Ornamentation folds (radiating vermiculate ridges) Nipple-shaped umbo Granulations


Spine-shaped Absent


Convex Conical Globular


Basal plate with vermiculate ornamentation folds


Spine-shaped umbos Absent


Present


1773a, b, 2367a–d, g, h,M2367a–i;Destombes collection). Some other historical samples are registered in the collections of the Muséum d’Histoire naturelle de Nantes, France (samples 2367 SR1–8; Régnault collection) and of the Natural History Museum of Oslo, Norway (samplesA 22122, A 29116, A 29120,A29122, A 29124, A 29132, A 29134; Spjeldnaes collection). The new material includes 119 samples (containing about 400 individuals), the bulk of which are deposited in the paleontological collections of the Université Cadi-Ayyad deMarrakech (samples AA.TAM. OI.1–80, AA.TISa.OI.1–27; Lefebvre and RALI collections), Marrakesh,Morocco; some other specimens are registered in the collections of theMusée des Confluences, Lyon, France (samples


Specimens in Figure 5 3–13, 14, 17–20


1, 2, 16 8, 13


6, 7, 9, 11, 14, 17, 19 5, 20


4, 10, 18 3


1, 2, 16


1, 2, 3–8, 13, 18–20 9-12, 14, 17


9, 13, 14, 17 6, 7, 11


5, 8, 10, 12, 19, 20 3, 4, 18


6, 7, 9, 11, 13, 14, 17, 19, 20 3-5, 8, 12, 18 10


4, 6–9, 11, 13, 17, 19, 20 3, 5, 12, 18


13 17


18–20 14 15


2, 16


14, 17 18–20


ML 20.269.379–382; Reboul collection) and of the Université Claude Bernard-Lyon 1, Villeurbanne, France (samples UCBL- FSL 711.159–711.163, UCBL-FSL 711.650, UCBL-FSL 712.044, 712.045; Reboul collection).


Morphological disparity and species differentiation within the genus Rhopalocystis


Thecal features.—The unrooted cladogram resulting from the cladistic analysis of 58 Rhophalocystis specimens based on nine discrete morphological characters suggests the distinction of


Figure 5. Illustration of the different states associated with the characters used for the cladistic analysis (see Table 1). (1–13) Drawings realized under a stereomicroscope with a camera lucida; (14–20) photographs of latex casts dusted with ammonium chloride. (1) Rhopalocystis grandis Chauvel, 1971 (holotype 1725a), large-sized theca partially complete, a radius including three globular-shaped radials is visible. (2) R. havliceki (AA-TISa-OI-22j), large-sized well-preserved theca; two radii are visible; the radials are spine-shaped; the infralaterals and some laterals have a spine-shaped umbo more or less elongated. (3) R. zagoraensis (holotype 1725g), elongated theca with three circlets of additional secondary plates between infralaterals and laterals and one incomplete circlet between laterals and radials. (4) R. zagoraensis (AA-TISa-OI-13a), theca with two irregular circlets of additional plates between infralaterals and laterals; the suture between the two visible infralaterals is not vertical, probably as a result of a growth abnormality. (5) R. zagoraensis (2367d1), theca showing one circlet of small secondary plates between infralaterals and laterals; a row of these additional plates is also visible between the laterals. (6) R. zagoraensis (AA-TISa-OI-20a), small theca with an incomplete circlet of additional secondary plates between infralaterals and laterals. (7) R. zagoraensis (AA-TISa-OI-21b), small theca with isolated small secondary plates inserted between infralaterals and laterals; the globular shape of the radials is clearly visible. (8) R. zagoraensis (AA-TISa-OI-16c), small theca with one complete circlet of additional plates between infralaterals and laterals; the basal is dislocated. (9) R. destombesi (ML 20.269.380a); theca constituted of seven well-organized circlets, no additional secondary plate, conical radials. (10) R. destombesi (2367a1), theca showing two additional circlets of secondary plates, one located between infralaterals and laterals, and another one between laterals and infralaterals, conical radials. (11) R. destombesi (2367 SR5), theca with one circlet of small secondary plates between infralaterals and laterals. (12) R. zagoraensis (2367c2), aboral part of theca with one complete circlet of additional plates between infralaterals and laterals, and a row of additional plates inserted between laterals; three radii can be observed, each composed of one column of four globular-shaped radials. (13) Rhopalocystis fraga Chauvel, 1971 (1725g'1), theca composed of six circlets of plates including three circlets of convex radials. (14) R. destombesi (1157e1, right side, and 1157e2, left side), specimens having nipple-shaped umbos centered in the middle of most of their thecal plates. (15) R. grandis (AA-TISa-OI-5), fragment of theca constituted of irregular plates covered with small granulations. (16) R. havliceki (AA-TAM-OI-9), large-sized theca; the aboral extremity is missing; numerous laterals exhibit a spine-shaped relatively elongated umbo. (17) R. destombesi (2367b1), unornamented specimen with smooth thecal plates. (18–20) R. zagoraensis, thecae showing more or less continuous ornamentation folds, basals with vermiculated folds. (18) UCBL-FSL 712.044a. (19) AA-TISa-OI-16a. (20) AA-TISa-OI-17e. (1, 3, 13, 14) Locality 809 (=1157 = 1725), late Tremadocian (A. murrayi Zone); (2, 4, 6–8, 15, 18–20) locality Z-F12c, late Tremadocian (A. murrayi Zone); (9) locality Z-F25, middle Floian (?B. jacksoni Zone); (5, 10–12, 17) locality 2367, late Tremadocian (H. copiosus Zone); (16) locality Z-F1, late Tremadocian (A. murrayi Zone). Scale bars = 1 cm.


1, 3–8, 12, 18–20 2


9–11, 14, 17


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