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A Small Antiarch, Minicrania Lirouyii Gen. Et Sp. Nov., from the Early

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A Small Antiarch, Minicrania Lirouyii Gen. Et Sp. Nov., from the Early A Small Antiarch, Minicrania lirouyii Gen. et sp. nov., from the Early Devonian of Qujing, Yunnan (China), with Remarks on Antiarch Phylogeny Author(s): Min Zhu and Philippe Janvier Reviewed work(s): Source: Journal of Vertebrate Paleontology, Vol. 16, No. 1 (Mar. 19, 1996), pp. 1-15 Published by: Taylor & Francis, Ltd. on behalf of The Society of Vertebrate Paleontology Stable URL: http://www.jstor.org/stable/4523686 . Accessed: 22/04/2012 21:43 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. The Society of Vertebrate Paleontology and Taylor & Francis, Ltd. are collaborating with JSTOR to digitize, preserve and extend access to Journal of Vertebrate Paleontology. http://www.jstor.org Journal of VertebratePaleontology 16(1):1-15, March 1996 C 1996 by the Society of VertebratePaleontology A SMALL ANTIARCH, MINICRANIA LIROUYII GEN. ET SP. NOV., FROM THE EARLY DEVONIAN OF QUJING, YUNNAN (CHINA), WITH REMARKS ON ANTIARCH PHYLOGENY MIN ZHU' and PHILIPPE JANVIER2 'Institute of Vertebrate Paleontology and Paleoanthropology (IVPP), Academia Sinica, P.O. Box 643, Beijing 100044, China 2U.R.A.12 du C.N.R.S., Laboratoire de Paleontologie, Mus6um National d'Histoire Naturelle, 8 rue Buffon, 75005 Paris, France ABSTRACT-A new Early Devonian (Lochkovian) yunnanolepiform-like antiarch Minicrania lirouyii gen. et sp. nov. is describedfrom Qujing, Yunnan,southeastern China. This tiny antiarchpossesses both a preorbitaldepression and a preorbitalrecess. It is characterizedby a long and narrow skull-roof, short obstantic margin, large orbital fenestra, fairly shortdorsal wall of the trunk-shieldand a close fusion of the dermalplates. It shows some neurocranialstructures, observed here for the first time in an antiarch.A growth series of the skull-roofshows a slight allometricgrowth. The phylogeny of antiarchsis discussed on the basis of new charactercombinations observed in this new form. It is suggested that Minicraniais the sister-groupof euantiarchsplus sinolepids, sharing with them the preorbitalrecess. The Yunnanolepidaewould be the sister-groupof the Chuchinolepidae(i.e., "procondylolepids"),and the Yunnano- lepiformesare re-definedhere to include only these two groups. INTRODUCTION one (supposedly juvenile) is shorter than 2.2 mm in length (that is, the smallest fossil vertebrate head found so far!). Liu Y.-H. (1963) erected the antiarch genus Yunnanolepis on The specimens described below are housed in the Institute the basis of a skull-roof from the Early Devonian of Qujing, of Vertebrate Paleontology and Paleoanthropology (IVPP), Ac- Yunnan, China. At that time, Yunnanolepis, first assigned to the ademia Sinica, Beijing. Asterolepiformes by Liu Y.-H. (1963), was the earliest known antiarch record. Subsequently, more detailed works (Chang K.- SYSTEMATIC DESCRIPTION J., 1978; P'an and Wang, 1978; Zhang G.-R., 1978; Zhang M.- PLACODERMI 1848 M., 1980) showed that Yunnanolepis and its relatives represent M'Coy, the most primitive antiarchs, lacking a complex dermal pectoral ANTIARCHACope, 1885 fin a new order G.- joint. Therefore, Yunnanolepiformes (Zhang MINICRANIAgen. nov. R., 1978) was erected to distinguish them from the other an- tiarchs, although the monophyly of this taxon is poorly sup- Diagnosis---as for the type species (by monotypy). ported (Janvier and Pan, 1982). Since the Yunnanolepidae (in- Type species-Minicrania lirouyii gen. et sp. nov. cluding Yunnanolepis) is the best known family of the Yunnan- Etymology-Mini, from minus (lat.) = small; crania, from olepiformes, it has been chosen as the root in the phylogenetic cranium (Lat.) = skull. analysis of antiarchs (Zhang and Young, 1992). However, as Remarks-Minicrania lirouyii, the only species of the ge- discussed below, yunnanolepids are relatively specialized in nus, is the smallest antiarch recorded so far. The entire dermal several characters, and their selection as the root for antiarch armour (skull-roof and trunk-shield) in the presumed adult is phylogeny would result in biases in the assessment of character less than 20 mm in length. polarities. Minicrania is very suggestive of the Yunnanolepiformes by In 1988, one of us (Z.M.) collected abundant early vertebrate its simple pectoral fin articulation and triangular preorbital de- it differs from the fossils (galeaspids, acanthodians, sarcopterygians and placo- pression. However, typical Yunnanolepidae in the characters: the skull-roof more or less derms) from the Lochkovian Xishancun Formation, in Qujing, following 1) being with the ratio Yunnan (Zhu, 1991, 1992, in press). Among this material, more square-shaped breadth/length approximately 1.0, the of a recess, the short obstantic than 40 specimens belong to a new, very small-sized antiarch, 2) presence preorbital 3) 4) the fused skull-roof and trunk-shield bones, Minicrania lirouyii gen. et sp. nov., described below. This spe- margin, closely cies is of the its 5) the anterior median dorsal plate roughly hexagonal in shape very suggestive Yunnanolepidae by simple with a broad anterior the median dorsal fin but it looks less than the latter in margin, 6) posterior pectoral joint, specialized and and the crista transversalis in- several a number of as to plate fairly large broad, 7) respects, thereby raising questions terna to the ventral antiarch All the material of this new form is posterior extending laterally posterior pro- phylogeny. pre- cess of the PMD The latter character is also found in the served as internal and external molds in a sedi- plate. fine-grained yunnanolepiform Zhanjilepis. ment, without any trace of bone. Therefore, it was studied es- sentially on the basis of elastomere casts. MINICRANIA nov. The growth series of the antiarch Bothriolepis canadensis LIROUYII sp. (Figs. 1-11) and its significance to antiarch evolution has been studied by Werdelin and Long (1986), and yet another example is given Diagnosis-Very small-sized antiarch, with closely fused here. The skull-roofs of M. lirouyii of various sizes are sup- skull-roof and trunk-shield bones; dermal bones ornamented posed here to form a relatively complete growth series. The with comparatively large, closely-set tubercles; obstantic mar- largest skull-roof is about 6 mm in length while the smallest gin short; postpineal plate convex and short; very large preor- 1 2 JOURNAL OF VERTEBRATE PALEONTOLOGY, VOL. 16, NO. 1, 1996 D Plesiotype--an external mold of the skull-roof associated with the ventral wall of the trunk-shield of a juvenile individual, io V10505.2 (Fig. 5C). Type Locality-Liaoguoshan, Qujing, Yunnan, China. Type Horizon-Xishancun Formation, Cuifengshan Group (Lochkovian, Early Devonian). Etymology-A species dedicated to Mr. Li Rou-yi (Qujing, Yunnan) who kindly provided help in collecting the specimens. Referred Material-skull-roofs (external or internal molds), so 10505.3-13, 46; trunk-shields, V10505.14-45. A Description B0 B' Measurements-Two systems of measurements of the an- ,, tiarchs are mentioned in the literature (Young, 1988). One is CIO that of Stensii (1948), followed and modified by Miles (1968) and Young (1988), and which suits best disarticulated speci- mens. The other is that of Werdelin and Long (1986) which is appropriate for articulated specimens. However, for articulated specimens whose bone sutures could not be observed, as in the present case, the system of Werdelin and Long has its limits. dend mpl In fact, a third system (Zhang G.-R., 1978) has been ignored. In the case of articulated skull-roofs, especially when no suture FIGURE 1. Skull-roof of Minicrania lirouyii (gen. et sp. nov.), show- can be seen, the system of Zhang G.-R. (1978) is more con- the landmarks and distances used in measurements. Landmarks ing of venient and depicts better the overall shape of the skull-roof. the A = anterolateral of anterior end skull-roof: (A') angle skull-roof, In this paper, Zhang's system is modified to accord with the of lateral skull-roof margin; B (B') = posterolateral angle (preobstantic current method of of end of lateral skull-roof C morphometrics (Fig. 1). corner) skull-roof, posterior margin; (C') Ten distances were measured on eleven skull-roofs of Mini- corner of skull-roof, end of obstantic - postobstantic posterior margin; crania and indices were calculated D = rostral angle of skull-roof; E = middle point of orbital fenestra lirouyii (Table 1), eight (Ta- margin of premedian plate; F = middle point of orbital fenestra margin ble 2). For comparison, the table of Zhang G.-R. (1978) for of postpineal plate; G = posterior angle of skull-roof; H (H') = lateral skull-roofs of Yunnanolepis chii was modified into Tables 3 and point of orbital fenestra. Distances. AA' = rostral width of skull-roof; 4. Since there are individual variations in skull-roofs, these BB' = total width of skull-roof; CC' = posterior width of skull-roof; kinds of indices only provide information about their shape and = = HH' width of rostral margin; DE length of premedian plate; pre- facilitate comparisons. orbital length of skull-roof; EF = length of orbital fenestra; FG = post- Skull-roof-The skull-roof of Minicrania is known orbital of DG = DE EF + of skull- lirouyii length skull-roof; + FG, length from fourteen of various either in external or AB = of lateral skull-roof BC = specimens sizes, roof; (A'B') length margin; (B'C') internal molds. In the skull-roof in either adult length of obstantic margin of skull-roof. dend, opening of the endolym- general, plates or assumed on the basis of their relative phatic duct; io, infraorbital sensory-line groove; mpl, "middle pit-line"; juvenile specimens (as so, supraorbital sensory-line groove. size), are tightly fused and no suture could be observed.
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