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Novitates PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y

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Novitates PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y AMERICAN MUSEUM Novitates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2998, 7 pp., 2 illustrations March 6, 1991 Morocconites Struve, 1989, A Devonian Acastine Trilobite (Calmoniidae: Acastinae) GREGORY D. EDGECOMBE1 ABSTRACT A well-preserved dorsal exoskeleton of Moroc- toides Delo, 1935, with unique modifications of conites malladoides Struve, 1989, from the De- the cephalic border and pygidium. Morocconi- vonian ofMorocco, allows a complete description tinae Struve, 1989, is a subjective synonym ofthe ofthe type species ofthis monotypic genus. Moroc- calmoniid subfamily Acastinae Delo, 1935. conites is most closely related to the acastine Acas- INTRODUCTION Morocconites malladoides Struve, 1989, me to examine a superbly preserved acastine was based on a mostly exfoliated dorsal exo- from Devonian strata at Alnif in the Mo- skeleton from Devonian strata in the Alnif roccan North Sahara. It was determined that area, west of Erfoud, Maider, Morocco. The the outstretched dorsal exoskeleton described species was interpreted as "possibly rooting in this paper is conspecific with the holotype in dalmanitinine dalmanitaceans" and with ofMorocconites malladoides Struve. The new "zeliszkelline/dalmanitinine descent ... ob- specimen provides strong evidence for clos- vious" (Struve, 1989: 567). An independent est relations to taxa classified in the calmoni- genus and subfamily (Morocconitinae Struve, id subfamily Acastinae (notably Acastoides 1989) were established with the hypothesis Delo, 1935). Accordingly, the subfamily Mo- that a "mixture of distinctly divergent char- rocconitinae is removed from the Dalmani- acters of different, clearly separated phyletic tidae, and is regarded as a synonym ofAcas- branches ... seems to root deep in the early tinae (Calmoniidae). New morphological history of the dalmanitids" (p. 568). These information afforded by fine preservation, as ancient (Ordovician) affinities are intriguing, well as necessity for differentia from similar given the Emsian or Eifelian age attributed acastines, have stimulated a revised diag- to Morocconites. nosis of Morocconites. Simon Harrison ofBath, England, allowed Descriptive terminology used herein is Department of Invertebrates, American Museum of Natural History; Department of Geological Sciences, Co- lumbia University. Copyright © American Museum of Natural History 1991 ISSN 0003-0082 / Price $1.30 2 AMERICAN MUSEUM NOVITATES NO. 2998 generally that of Harrington et al. (1959). Form and incision of the lateral glabellar Points of curvature on the facial suture are furrows (e.g., evenly curved S3) are very sim- lettered as by Richter and Richter (1949: fig. ilar in Morocconites malladoides and Acas- 1). Lateral glabellar lobes and furrows are toidespaeckelmanni (cf. Richter and Richter, denoted following Jaanusson (1956: 37), with 1952: pl. 2) and inA. constricta (Salter) (Rich- the prefix L- or S-, numbered 0-4 from rear ter and Richter, 1954: pl. 3, fig. 42). A short- (occipital) to front. Pseudo-articulating half ened (exsag.) LI groups Morocconites with ring is as suggested by Holloway (1981: 695). the clade Acastinae + Acastavinae + Astero- The term epipleuralfurrow is introduced for pyginae (Destombes and Henry, 1987: 136), transverse grooves on the pygidial pleural ribs in contrast to primitive Dalmanitidae and between the pleural and interpleural furrows. Calmoniinae. This synapomorphy excludes These are observed in the Acastinae (Mor- certain taxa previously assigned to Acastinae, occonites), Phacopidellinae (e.g., Chlupac, which have plesiomorphic conditions of 1977: pl. 1, fig. 11), and Dalmanitidae (e.g., longer (exsag.) L1, with S1 inclined abaxially, Destombes, 1972: pl. 4, fig. 7). and bifurcate proximally (e.g., Andinacaste Eldredge and Branisa, 1980; "Acaste" bir- minghamensis Norford, 1972; Juyuyops Bal- SYSTEMATIC PALEONTOLOGY dis and Blasco in Baldis et al., 1976). Acas- FAMILY CALMONIIDAE tinae s.s. (e.g., Acaste Goldfuss; Acastoides DELO, 1935 Delo + Morocconites Struve; Acastocephala SUBFAMILY ACASTINAE Shergold) have S2 approximately transverse DELO, 1935 (although distinctly convex forward). Andi- nacaste and "Acaste" birminghamensis re- Morocconites Struve, 1989 tain the primitive state of S2 inclined abax- TYPE SPECIES: Morocconites malladoides ially (shared with primitive Calmoniinae and Struve, 1989. By original designation. Baniaspis Destombes, 1972). DIAGNosIs: Acastinae with anterior border Likewise, the auxilliary impression system having very long, upcurved median spine. in the frontal glabellar lobe ofMorrocconites Cephalic lateral border broadened against an- is that diagnostic of post-Llandovery Acas- terior genal region; lateral margins subpar- tacea (Eldredge, 1979), a triangular pattern allel between border swelling and genal angle. ofdiscrete radiating rows ofpits (fig. 1). Llan- Pygidial margin entire; transverse apodemes dovery "Acaste" birminghamensis Norford in anterior five ring furrows; ring furrows resembles Ordovician Acastacea (Kloucekia shallow medially. Anterior pleural furrow Delo; Phacopidina Bancroft; Baniaspis Des- narrow but moderately deep, succeeding tombes) in the presence of muscle scars in pleural furrows shallow, subequal to inter- the median region of the impression system pleural furrows; shallow epipleural furrows (Norford, 1972: pl. 1, fig. 13). Glabellar mor- bisect anterior and posterior pleural bands. phology of Morocconites bears resemblance DISCUSSION: Morocconites is maintained as to that ofthe Emsian asteropygine Kayserops a monotypic genus, but its diagnosis is re- kerfornei (Pichard) (see Morzadec, 1983: pls. vised to distinguish it from other genera of 12, 13), as does the long cephalic antero- Acastinae. The conspicuous cephalic anterior median spine; however, the smooth-mar- border spine is peculiar for an acastine, al- gined pygidium and rounded genal angle of though similar to Calmoniinae such as Morocconites are acastine features which pre- Schizostylus Delo and the acastavine Cen- clude assignment to Asteropyginae. Structure tauropyge Haas, 1968. The entire-margined, of the thoracic pleurae is also comparable to semicircular pygidium of Morocconites with that ofAcastinae, e.g., the gently sinuous epi- a distinct border furrow is most typically facetal pleural furrow, and blunt distal pro- acastine [although, as noted by Struve (1989), jection from behind the articulating facet. In pygidial form is superficially similar to that addition to similarities in glabellar furrows, of Phacopidellinae. This applies particularly Acastoides Delo, 1935, is most comparable to species with shallow epipleural furrows, to Morocconites in the relatively short (exsag.) e.g., Phacopidella glockeri (Barrande)]. eye with a low visual field, and strong arching 1991 EDGECOMBE: ACASTINE TRILOBITE 3 (tr.) ofthe cephalon, i.e., steep librigenal field. Shergold (1966: 200) cited a subpentagonal cephalic outline and eyes set high up on the genae as characteristic ofAcastoides; both of these features are shared with Morocconites. The relatively deep incision of the cephalic lateral border furrow in Devonian species such as Acastoides paeckelmanni (see Haas, 1968: pl. 32) is peculiar among Acastinae, but is shared with Morocconites. Among spe- cies assigned to Acastoides, the pygidium of A. (?Acastoides) aspidium Haas, 1968, most - closely resembles that of Morocconites mal- pgf likewise has the axis ladoides. This species Fig. 1. Camera lucida drawing of glabella in elongated behind the anterior three rings, with anterodorsal view, with auxiliary impression sys- less marked anteroposterior differentiation tem (shallow pits in cuticular surface) illustrated than other Acastoides. Devonian species from on left side offrontal lobe. pgf, preglabellar furrow. the Malvinokaffric Realm formerly assigned to Acastoides are regarded as Calmoniinae (cf. Eldredge and Branisa, 1980), and have been reassigned to Pennaia Clarke (see Edge- ward opposite L3; axial furrow shallowest op- combe, in press). posite L2 and posterior part of L3, narrow With this hypothesis of relationships, the but distinctly incised opposite anterior part recognition of a monotypic subfamily Mo- of L3/posterolateral edge of frontal lobe and rocconitinae would render Acastinae a para- Ll/LO. Preglabellar furrow moderately deep phyletic group (since Morocconites is, cladis- (sag.), shallowing abaxially; cranidial anterior tically, a modified acastine). The genus is border short, moderately declined forward, accordingly classified in Acastinae, with which longest sagittally in dorsal orientation but Morocconitinae is placed in subjective syn- visible across width. Cephalic anterior border onymy. Affinities to Siluro-Devonian Acas- spine gently curved, tip steepened slightly be- toides are also more compatible with the yond vertical, raised strongly above glabella; stratigraphic occurrence ofMorocconites than spine triangular, gently tapering in dorsal Struve's (1989) contention that the genus sig- view. Anterior branch of facial suture with nals early (i.e., Ordovician) dalmanitid his- nearly straight anteroventral course between tory. -y and just outside anterolateral region of frontal glabellar lobe; cranidial anteromedian Morocconites malladoides margin arched (tr.) dorsally. Posterior branch Struve, 1989 offacial suture gently curved, convex forward Figures 1, 2 (in dorsal orientation) between e and w, set in shallow, narrow genal sulcus; suture sharp- Morocconites malladoides
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