Component Interaction of ESCRT Complexes Is Essential for Endocytosis-Dependent Growth, Reproduction, DON Production and Full Virulence in Fusarium graminearum

Multivesicular bodies (MVBs) are critical intermediates in the trafficking of ubiquitinated endocytosed surface proteins to the lysosome/vacuole for destruction. Recognizing and packaging ubiquitin modified cargoes to the MVB pathway require ESCRT (Endosomal sorting complexes required for transport) machinery, which consists of four core subcomplexes, ESCRT-0, ESCRT-I, ESCRT-II, and ESCRT-III. Fusarium graminearum is an important plant pathogen that causes head blight of major cereal crops. Our previous results showed that ESCRT-0 is essential for fungal development and pathogenicity in Fusarium graminearum. We then, in this study, systemically studied the protein-protein interactions within F. graminearum ESCRT-I, -II or -III complex, as well as between ESCRT-0 and ESCRT-I, ESCRT-I and ESCRT-II, and ESCRT-II and ESCRT-III complexes and found that loss of any ESCRT component resulted in abnormal function in endocytosis. In addition, ESCRT deletion mutants displayed severe defects in growth, deoxynivalenol (DON) production, virulence, sexual, and asexual reproduction. Importantly genetic complementation with corresponding ESCRT genes fully rescued all these defective phenotypes, indicating the essential role of ESCRT machinery in fungal development and plant infection in F. graminearum. Taken together, the protein-protein interactome and biological functions of the ESCRT machinery is first profoundly characterized in F. graminearum, providing a foundation for further exploration of ESCRT machinery in filamentous fungi

Observations on the Hipparion Red Clays of the Loess Plateau

We discuss the history of exploration of Red Clay vertebrate fossils of the Loess Plateau, and record observations on “Hipparion Red Clay” localities of Shanxi and Gansu provinces. Red clay is widespread across the Loess Plateau, but misleading as a descriptive term because many deposits are neither red nor dominated by clay-size sediment. Many red clay sequences contain paleosols, but also water-laid deposits. Our survey includes well-known localities of Yushe Basin, Renjiagou and Leijiahe near Lingtai, and sites of Qingyang and Qin’an. We discuss fossils found at Renjiagou, a new discovery of micromammals (Pliopentalagus) from Yucun, and Pliocene burrows observed at Hu Jia Cun, both latter localities northeast of Lingtai. Observations are consistent with a high rate of supply of clay sized particles, likely air-fall origin, throughout the Loess Plateau during the late Miocene. In some areas where fluvial or lake processes dominate, red clay particles are replaced in part or completely by water-laid coarse-grained deposits. We saw no evidence for a dry Late Miocene-Early Pliocene, but rather hypothesize well-watered habitat of high productivity. Locally diverse vertebrate fossils attest to high terrestrial biomass for Hipparion faunas.Human Evolutionary Biolog

Two Rab5 Homologs Are Essential for the Development and Pathogenicity of the Rice Blast Fungus Magnaporthe oryzae

The rice blast fungus, Magnaporthe oryzae, infects many economically important cereal crops, particularly rice. It has emerged as an important model organism for studying the growth, development, and pathogenesis of filamentous fungi. RabGTPases are important molecular switches in regulation of intracellular membrane trafficking in all eukaryotes. MoRab5A and MoRab5B are Rab5 homologs in M. oryzae, but their functions in the fungal development and pathogenicity are unknown. In this study, we have employed a genetic approach and demonstrated that both MoRab5A and MoRab5B are crucial for vegetative growth and development, conidiogenesis, melanin synthesis, vacuole fusion, endocytosis, sexual reproduction, and plant pathogenesis in M. oryzae. Moreover, both MoRab5A and MoRab5B show similar localization in hyphae and conidia. To further investigate possible functional redundancy between MoRab5A and MoRab5B, we overexpressed MoRAB5A and MoRAB5B, respectively, in MoRab5B:RNAi and MoRab5A:RNAi strains, but neither could rescue each other’s defects caused by the RNAi. Taken together, we conclude that both MoRab5A and MoRab5B are necessary for the development and pathogenesis of the rice blast fungus, while they may function independently

Retromer Is Essential for Autophagy-Dependent Plant Infection by the Rice Blast Fungus

We thank Dr. Yizhen Deng at the Temasek Life sciences Laboratory (TLL) for providing the RFP-MoAtg8 plasmid. We would like to thank Drs. Zhenbiao Yang (University of California, Riverside) and Xianying Dou (Fujian Agriculture and Forestry University) for helpful discussions.Author Summary The rice blast fungus Magnaporthe oryzae utilizes key infection structures, called appressoria, elaborated at the tips of the conidial germ tubes to gain entry into the host tissue. Development of the appressorium is accompanied with autophagy in the conidium leading to programmed cell death. This work highlights the significance of the Vps35/retromer membrane-trafficking machinery in the regulation of autophagy during appressorium-mediated host penetration, and thus sheds light on a novel molecular mechanism underlying autophagy-based membrane trafficking events during pathogen-host interaction in rice blast disease. Our findings provide the first genetic evidence that the retromer controls the initiation of autophagy in filamentous fungi.Yeshttp://www.plosgenetics.org/static/editorial#pee

Figure 8. IVPP V15726 in A new species of crown-antlered deer Stephanocemas (Artiodactyla, Cervidae) from the middle Miocene of Qaidam Basin, northern Tibetan Plateau, China, and a preliminary evaluation of its phylogeny

Figure 8. IVPP V15726, Stephanocemas sp. from IVPP locality CD0406. A, dorsal, and B, ventral views of antler fragment. Scale is for both views.Published as part of <i>Wang, Xiaoming, Xie, Guangpu & Dong, Wei, 2009, A new species of crown-antlered deer Stephanocemas (Artiodactyla, Cervidae) from the middle Miocene of Qaidam Basin, northern Tibetan Plateau, China, and a preliminary evaluation of its phylogeny, pp. 680-695 in Zoological Journal of the Linnean Society 156 (3)</i> on page 689, DOI: 10.1111/j.1096-3642.2008.00491.x, <a href="http://zenodo.org/record/10114804">http://zenodo.org/record/10114804</a&gt

Stephanocemas PALMATUS 2009, SP. NOV.

STEPHANOCEMAS PALMATUS SP. NOV. (FIGS 3–6) <p> <i>Holotype:</i> IVPP V15722, nearly complete, shed left antler without pedicel (Figs 3, 4). Collected by Gary T. Takeuchi and Jack Tseng on 13 August 2006.</p> <p> <i>Etymology:</i> <i>palmatus</i>, Latin, webbed, palmate; in allusion to palmate antler of the species.</p> <p> <i>Type locality:</i> IVPP locality CD0626 (N37° 06′ 12.6″, E97° 16′ 33.4″), north limb of Barun Yawula anticline, 30 km south-west of Delingha, Haixi Mongolian Autonomous Prefecture, Qinghai Province, northern Tibetan Plateau, China (Figs 1, 2).</p> <p> <i>Referred specimens:</i> IVPP V15723, palm part of antler with only one medial tine preserved (Fig. 5), from IVPP locality CD0630 (N37° 06′ 24.3″, E97° 15′ 59.7″); IVPP V15724, posterior half of a small juvenile antler fragment (Fig. 6), from IVPP CD0630.</p> <p> <i>Geology and age:</i> IVPP V15722 (from IVPP CD 0626) was produced from a sequence of rusty yellow to greenish cross-bedded sandstones and fine gravels along a north–south orientated wash on the western end of the north limb of the Barun Yawula anticline (Fig. 2). IVPP V15723 and V15724 (IVPP CD 0630) are also from the same rusty yellow sandstone layers but are stratigraphically ~ 115 m above IVPP CD0626. The <i>Stephanocemas</i> -producing sandstone, dipping toward the north, is more than 600 m above the axis of the Barun Yawula anticline, where reddish mudstones are exposed as the stratigraphically lowest exposure. Fifty kilometres north-west of the Barun Yawula anticline is the north limb of the Keluke (Kurliq) anticline, where a sequence of ~ 5000 m of Neogene sediments is exposed. Known as the Huaitoutala section, this long sequence contains vertebrate faunas ranging from the middle Miocene to early Pliocene age (Wang <i>et al.</i>, 2007) and palaeomagnetically dated to 15.7–1.8 Mya (Fang <i>et al.</i>, 2007). Lithologically the <i>Stephanocemas</i> -producing strata in Barun Yawula are approximately comparable to the greenish sandstones in the Huaitoutala section, where the middle Miocene Olongbuluk Fauna is found. The main difference seems to be that the Huaitoutala section is a much thicker sequence (the Olongbuluk Fauna alone spans more than 1000 m), whereas the same section in Barun Yawula is represented by substantially thinner beds, possibly indicating a lower rate of deposition.</p> <p> Faunally, the <i>Stephanocemas</i> -producing sandstones in Barun Yawula (localities CD 0626, 0630, 0633, 0634; see Fig. 2) have so far not yielded other agediagnostic taxa beside a few non- <i>Stephanocemas</i> cervid antler fragments, as well as fragments of turtles and teleost fishes. However, in the eastern end of the Barun Yawula anticline, ~ 6 km to the southeast of CD0626, a small assemblage of fossil mammals was collected. Of these, the most agediagnostic include an astragalus of hipparionine horse (IVPP CD 0618), indicating a late Miocene age. Based on correlations by satellite image (Fig. 2), the CD0618 locality is stratigraphically close to or slightly above the highest <i>Stephanocemas</i> -producing horizon (CD 0634), although such correlations are not without difficulties, particularly in the eastern end of the Barun Yawula anticline where multiple faults have horizontal offsets of up to several hundred metres. To the best of our knowledge (see further discussion in ‘Problems with Bohlin’s Qinghai Materials’ below), <i>Stephanocemas</i> is restricted to the middle Miocene strata (the Olongbuluk Fauna) in the eastern Qaidam Basin, i.e. no lagomerycine deer has been collected in known late Miocene faunas (Wang & Wang, 2001; Deng & Wang, 2004a, b; Wang <i>et al.</i>, 2007). We thus conclude that <i>Stephanocemas palmatus</i> probably represents the latest appearance of the lineage in the Tibetan Plateau in the latest middle Miocene, although the possibility exists that its upper-most occurrence (CD 0634) is earliest late Miocene.</p> <p> <i>Diagnosis: Stephanocemas palmatus</i> is easily distinguished from species of <i>Paradicrocerus</i> (<i>Paradicrocerus flerovi</i>, <i>Paradicrocerus elegantulus</i>, and <i>Paradicrocerus brevistephanos</i>) in its derived characters such as large size, loss of dorsal ridges between anterior and posterior tines, enlarged central (palm) portion of antler, and horizontally orientated side branches (tines). <i>Stephanocemas palmatus</i> differs from all previously known species of <i>Stephanocemas</i> (<i>Stephanocemas aralensis</i>, <i>Stephanocemas chinghaiensis</i>, <i>Stephanocemas rucha</i>, <i>Stephanocemas thomsoni</i>) in the following derived characters: large size, expanded central (palm) part of antler, more horizontally orientated tines, and multiple (three or more) tines being plate-like.</p> <p> <i>Description</i></p> <p> The unusual arrangement of antlers in <i>Stephanocemas</i> confounded Colbert (1936), who was unable to find a suitable method to describe its peculiar patterns. Although he agreed in principle with a nomenclatural scheme by Pocock (1933), who envisioned a series of dichotomous branching patterns in the posterior beams as seen in living cervids (see also Colbert, 1940), in practice Colbert found such a system difficult to apply to <i>Stephanocemas</i>. Instead, Colbert devised a rather arbitrary system of antler nomenclature of anterior (brow tine), posterior (beam), median, and external tines. Because the branching patterns in <i>Stephanocemas</i> are apparently derived independently from those in living cervids, Colbert’s system is adopted in this description to avoid implications of homology with living cervids.</p> <p> The orientation of antlers is not always easily ascertained without associated cranial elements or large ontogenetic series. Our determination for the holotype of <i>S. palmatus</i> was based on comparisons with the best-preserved series for <i>P. elegantulus</i> and <i>S. thomsoni</i>. In general, the widest tine is the posterior one, and its opposite tine is, by definition, the anterior tine. The lateral and medial tines are generally shorter than the anterior and posterior tines. The lateral tines tend to be longer and more upwardly curved relative to their medial counterparts. Given the above observation, we regard IVPP V15722 as a left antler.</p> <p> IVPP V15722 is a large, full adult individual. A juvenile partial antler from a higher stratigraphical horizon (see ‘Geology and age’), IVPP V15724, offers some sense of ontogenetic variations. Even in a very young individual, such as IVPP V15724, the palmate nature of the antler is readily apparent (Fig. 6), and distinctly different from juveniles of <i>S. thomsoni</i> (Colbert, 1936: fig. 5). The entirely palmate IVPP V15724 has four branches and the palm portion is folded along the anteroposterior axis.</p> <p> The palm portion of IVPP V15722 is mediolaterally expanded with a relatively high width/length ratio. Toward the medial-central aspect of the palm, there is a distinct dorsal spur, which is broken off at the base. Only a rounded scar is left for this spur. Other than this spur, the dorsal surface is flat and smooth without any trace of the dorsal ridge seen in <i>Paradicrocerus</i> (see Phylogeny). On the ventral side, an indication of a burr (coronet) is marked by small bony spurs at the bases of posterior and lateral tines. The burr is oval-shaped with its long axis orientated anteroposteriorly, which obviously corresponds to an oval cross-sectioned pedicel (stem). This burr-like structure, a concave shedding scar, and rough surface surrounding the burr clearly indicate a deciduous antler.</p> <p>The most characteristic aspect of the tines in IVPP V15722 is their spread-out appearance with their nearly horizontal orientation and their flattened cross-section. In lateral view, anterior and posterior tines spread out from the palm, initially horizontally toward the periphery and then gently curving upward toward their tips (Figs 3, 4). In anterior view, the medial tines (only the posterior one is preserved) are essentially horizontal whereas the lateral tines curve upward slightly, creating an asymmetrical appearance in terms of the upward curvatures of medial and lateral tines.</p> <p>Of the preserved tines, the posterior, posterior lateral, and anterior lateral tines are bifurcated, although, judging from the condition of the anterior lateral tine, the bifurcated tines do not extend far from the base of the split. The anterior tine is relatively short and wide in dorsal view. Its cross-section is basically dorsoventrally flat with the dorsal surface slightly convex. The medial corner of the tip is broken, but judging from the remaining parts, the anterior tine may be slightly bifurcated.</p> <p>The posterior tine (beam) is the widest of all the tines and maintains its width throughout its length, and slightly widens toward the distal end. The crosssection is very flat, with a convex ventral surface and a slightly concave dorsal surface. The tine is undoubtedly bifurcated, even though its lateral tip is broken off at the base of the bifurcation (see reconstruction in Fig. 10).</p> <p>The anterior lateral tine is slightly longer than the anterior tine. This flattened tine has shallow grooves on both dorsal and ventral surfaces throughout its length. This is also the only tine that twists almost 90° – its bifurcated tips are almost vertically aligned, with the effect of a maximum width when viewed laterally. This tine also preserves the best tips (both are slightly restored with epoxy resin).</p> <p>The middle lateral tine is broken midway along the length. As in the anterior lateral tine, it is dorsoventrally compressed. Its cross-section shows shallow grooves on dorsal and ventral sides, indicating bifurcated tips.</p> <p>The posterior lateral tine is longer than the posterior tine. Its dorsoventrally compressed surfaces have shallow grooves that lead to slightly bifurcated tips, which are not as distinct as those on the anterior lateral tine. The grooves taper off toward the anterior aspect of the tine.</p> <p>The posterior medial tine is also longer than the posterior beam. Its cross-section is somewhat triangular at its base, gradually becoming flattened toward the tip, and its tip is not bifurcated. The middle and anterior medial tines are broken off at the base. Both have a rounded cross-section, indicating a single tip, as in the posterior medial tine. The base of the middle medial tine is elevated to form a prominent hump above the dorsal surface of the palm.</p> <p> <i>Measurements (mm):</i> maximum anteroposterior diameter of palm: 77.7; maximum mediolateral diameter of palm: 53.6; maximum ¥ minimum diameters at base of burr: 35.0 ¥ 20.0; length of anterior tine: 39.3; maximum ¥ minimum diameters at base of anterior tine: 19.4 ¥ 9.9; length of posterior tine: 48.3; maximum ¥ minimum diameters at base of posterior tine: 28.2 ¥ 12.8; length of posterior medial tine: 54.7; maximum ¥ minimum diameters at base of posterior medial tine: 16.6 ¥ 12.6; maximum ¥ minimum diameters at base of middle medial tine: 11.2 ¥ 10.7; maximum ¥ minimum diameters at base of anterior medial tine: 14.9 ¥ 11.7; length of anterior lateral tine: 52.7; maximum ¥ minimum diameters at base of anterior lateral tine: 17.7 ¥ 10.9; maximum ¥ minimum diameters at base of middle lateral tine: 20.3 ¥ 12.8; length of posterior lateral tine: 58.0; and maximum ¥ minimum diameters at base of posterior lateral tine: 18.3 ¥ 10.9.</p> <p>COMPARISON AND DISCUSSION</p> <p> In this study, we restricted our concept of <i>Stephanocemas</i> to those Asiatic species that fall within the advanced clade of the <i>Paradicrocerus – Stephanocemas</i> lineage (see Phylogeny). Primitive species formerly included in <i>Stephanocemas</i>, such as <i>S. elegantulus</i> (Roger, 1898) and <i>S. brevistephanos</i> Baschanov & Nurumov, 1955, are here placed in the genus <i>Paradicrocerus</i>. In such a phylogenetic framework, <i>S. palmatus</i> is easily distinguishable from species of <i>Paradicrocerus</i> in its lack of dorsal ridges between anterior and posterior tines, expanded palm portion of the antler, and laterally spread tines.</p> <p> Within the genus <i>Stephanocemas</i>, the type species, <i>S. thomsoni</i> Colbert, 1936, is still the best documented species with a large array of individual antlers representing a nearly complete ontogenetic series. Such a series demonstrates a large amount of intraspecific variation in size, number, position, and orientation of individual branches (tines). Such a range of variations is probably typical for other species of the <i>Paradicrocerus – Stephanocemas</i> lineage as well because one of its basal species, <i>P. elegantulus</i> (Roger, 1898), shows similarly large variations (see Stehlin, 1937). The large ontogenetic series of <i>S. thomsoni</i> from the Tunggur Formation also allows a certain amount of confidence that <i>S. palmatus</i> falls outside the variations of <i>S. thomsoni</i>. Despite the substantial amount of flattening in the posterior tine in <i>S. thomsoni</i>, the rest of the tines remain essentially rounded in cross-section, with the exception of the anterior median tine in the holotype (AMNH 26782). These rod-like tines with a single tip seem to be a primitive condition in contrast to the mostly platelike tines with branched tips in <i>S. palmatus</i>. Furthermore, the palm portion of the antlers in <i>S. palmatus</i> is also mediolaterally widened relative to those in <i>S. thomsoni</i>. If such a comparison is correct, then <i>S. palmatus</i> represents the most derived member of the genus and helps to envision a morphological framework for other lesser known species.</p> <p> Colbert (1936) described a second species of <i>Stephanocemas</i>, <i>S. triacuminatus</i>, from the Tunggur Formation. However, following a suggestion by Stehlin (1937), Colbert (1940) soon recognized it as a species of <i>Lagomeryx</i>, a practice that has been followed since then (e.g. Young, 1964), as well as in this study.</p> <p> Beliajeva (1949) figured and described a partial antler of <i>Stephanocemas</i> sp. near Sarybulak River from the Zaysan Basin in eastern Kazakhstan. The Zaysan form is distinguished by its relatively small palm portion, rounded tine cross-sections, and more vertically orientated tines, characters that easily differ from those in <i>S. palmatus</i>.</p> <p> Young (1964) named a new species, <i>Stephanocemas chinghaiensis</i>, based on an antler from Ledu (= Nien-Pai) County, Qinghai Province, described by Bohlin (1937) (see additional comments in ‘Problems with Bohlin’s Qinghai Materials’ below). Based on published figures (Bohlin, 1937: text figs 42–44), the Ledu antler is distinct from <i>S. palmatus</i> in its smaller size, smaller palm portion, fewer medial and lateral tines (one each), more rounded cross-sections of the tines, and slightly more upright orientation of the tines.</p> <p> Beliajeva (1974) erected a new species, <i>Stephanocemas aralensis</i>, from the middle Miocene of western Kazakhstan. The holotype (PIN 1551-67) is a nearly complete left antler collected near Bes-Tyube, north of the Aral Sea. Based on the illustrated holotype and referred materials (Beliajeva, 1974: figs 2–4), <i>S. aralensis</i> is easily distinguished from <i>S. palmatus</i> in its smaller size, much more elongated burr (i.e. shedding scar; corresponding to a more mediolaterally compressed pedicel), less mediolaterally expanded palm portion, more upwardly (vertically) orientated peripheral tines, more rounded cross-sections for most peripheral tines, fewer lateral (one to two) and medial (two to three) tines, and more proximally branched posterior tine (more so in the holotype and a referred antler, PIN 1568-42, than in PIN 1551-68). With the possible exception of the elongated burrs, most of the characters in <i>S. aralensis</i> are primitive, and are probably far removed from those of <i>S. palmatus</i>.</p> <p> More recently, Ginsburg & Ukkakimapan (1983) described a new species, <i>Stephanocemas rucha</i>, from lignitic deposits of the Li Basin in northern Thailand, which represents the southern-most occurrence of the genus. Initially thought to be late middle to early late Miocene in age mostly based on the presence of proboscideans and <i>Stephanocemas</i> (Ginsburg & Ukkakimapan, 1983; Ginsburg, 1984), the age of the Li Fauna was revised downward to early Miocene (equivalent to the MN4 of the European mammal zones), based on long-distance comparison of mammals from Europe (Mein & Ginsburg, 1997). However, more recent studies of magnetostratigraphy (Benammi <i>et al.</i>, 2002) and fossil rodents (Chaimanee <i>et al.</i>, 2007) from the nearby Mae Moh Basin, placed the Li Basin strata (considered equivalent in age to that of the Mae Moh Basin) back in the late middle Miocene age, around 13–12 Mya. Regardless of its age, the Thai species is the southernmost occurrence of the genus and is zoogeographically important. Unfortunately, the holotype of <i>S. rucha</i> is poorly preserved. Additional antler fragments were listed, but these were not figured nor described (Mein & Ginsburg, 1997). Our present comparison is thus based on the holotype (T Li 67) only. The palm portion of <i>S. rucha</i> is smaller and less laterally spread than that of <i>S. palmatus</i>. Judging from the anterior tine, the only fully preserved one on the holotype, and the bases of the two lateral tines, the radiating tines are all orientated rather vertically, i.e. forming an angle greater than 45° with the horizontal plane. In this condition, the Thai species is thus somewhat primitive among advanced species of <i>Stephanocemas</i> (see Fig. 9). If our reconstructed outline of the posterior tine (Fig. 10) is correct, an admittedly speculative reconstruction based on published figures, then <i>S. rucha</i> shares with <i>S. thomsoni</i> and <i>S. palmatus</i> a flattened posterior tine.</p> <p> Ye (1989) referred to <i>Stephanocemas</i> aff. <i>thomsoni</i> 20 partial antlers, as well as dental materials, from the lower part of the Halamagai Formation (middle Miocene), Junggar Basin, Xinjiang Province. The relatively large sample size, representing both adult and juvenile individuals, reveals a substantial amount of variations. Nonetheless, the following features in the Junggar materials are consistently different from those in <i>S. palmatus</i>: smaller size, smaller palm part of the antler, more rounded crosssections in tines, and slightly more upright tines, although the two taxa seem to share a mediolaterally expanded palm.</p> <p> Abdrachmanova (in Tleuberdina, Bayshashov & Abdrachmanova, 1993) described a new species, <i>Stephanocemas actauensis</i>, from the Aktau Mountain area in the Ily Basin of eastern Kazakhstan. The upper Aktau Mountain strata that produced <i>S. actauensis</i> were estimated to be early Miocene in age (Tleuberdina, 1994; Lucas & Bendukidze, 1997). The holotype is a small partial antler with five or six tines. Assuming the Aktau antler is an adult, it differs from <i>S. palmatus</i> by the following primitive characters: small size, five to six tines with rounded crosssections, and more upward orientation of the tines.</p> <p> Most recently, Li <i>et al.</i> (1998) referred to <i>S. thomsoni</i> four antlers, two jaws, and several teeth from the Hongliugou Formation in Tongxin, Ningxia Autonomous Region, an area that was previously known to contain some fragmentary materials of the genus (Chen, 1978). As typified by the Dingjiaergou local fauna, the middle part of the Tongxin strata is middle Miocene (Tunggurian) in age (Qiu, Wu & Qiu, 1999). Judging from a figured antler (BPV 1212) (Li <i>et al.</i>, 1998: figs 1, 2) and from the authors’ description, the Tongxin materials seem closest to <i>S. palmatus</i> from Qaidam. BPV 1212, presumably a mature individual, has reached the stage of palm expansion of the latter. Its peripheral tines seem also substanti

Figure 5. IVPP V15723 in A new species of crown-antlered deer Stephanocemas (Artiodactyla, Cervidae) from the middle Miocene of Qaidam Basin, northern Tibetan Plateau, China, and a preliminary evaluation of its phylogeny

Figure 5. IVPP V15723, referred specimen of Stephanocemas palmatus sp. nov. A, dorsal, and B, ventral views of palm portion of antler.Published as part of <i>Wang, Xiaoming, Xie, Guangpu & Dong, Wei, 2009, A new species of crown-antlered deer Stephanocemas (Artiodactyla, Cervidae) from the middle Miocene of Qaidam Basin, northern Tibetan Plateau, China, and a preliminary evaluation of its phylogeny, pp. 680-695 in Zoological Journal of the Linnean Society 156 (3)</i> on page 685, DOI: 10.1111/j.1096-3642.2008.00491.x, <a href="http://zenodo.org/record/10114804">http://zenodo.org/record/10114804</a&gt

Stephanocemas SP.

STEPHANOCEMAS SP. (FIGS 7, 8) Referred materials: IVPP V15725 (Fig. 7), a partial antler from IVPP CD9818 (N37° 14′ 12.6″ E96° 44′ 49.8″), in the north limb of the Keluke anticline of the eastern Qaidam Basin, collected by Feng Wenqing on 8 July 1998; IVPP V15726 (Fig. 8), an antler fragment from IVPP CD0406 (N36° 58′ 54.5″ E97° 23′ 29.3″), in the Bayin Mountain section of the eastern Qaidam Basin, collected by Ni Xijun on 3 September 2004. Geological age: IVPP locality CD9818 is in the upper range of the Olongbuluk Fauna (Wang et al., 2007) and has been palaeomagnetically estimated to be between 12–13 Mya (Fang et al., 2007). This locality also produced an antler fragment of Lagomeryx, another early cervid frequently associated with Stephanocemas. The Bayin Mountain exposures, to the east of the fossiliferous Naoge area, have yielded few vertebrates, and IVPP CD0406 is one of a handful that produced identifiable materials. CD0406 has been correlated with the Olongbuluk Fauna as well, based on section measurements and biostratigraphical comparisons (Wang et al., 2007). Lithologically, CD0406 and CD9818 are both in the predominantly greenish sandstones. Vertebrate fossils in the Olongbuluk Fauna in both sections (Huaitoutala and Bayin Mountain) are relatively rare, and a precise correlation of the two localities, separated by a distance of 65 km, is not possible at this moment. It is likely, however, that both CD0406 and CD9818 are stratigraphically lower than beds that produced S. palmatus in Barun Yawula, and that all Stephanocemas -producing localities in the Qaidam Basin are middle Miocene in age (see further discussion in ‘Geology and age’ under S. palmatus). Description and comparison: Both IVPP V15725 and V15726 are of similarly small sizes, but lack of additional materials from their respective localities makes it difficult to judge if they belong to juvenile individuals. Although their tips are mostly missing, V15725 has at least six tines, and those for V15726 are not clear because only the posterior half the palm is preserved. The presumed posterior tine on V15725 is short, broad-tipped, and vaguely suggestive of a branched tip. If the latter is not an artefact of weathering, it recalls the condition in S. palmatus. The pedicel is partially preserved in V15725, suggesting that this antler was not shed during life. In contrast, V15726 has a distinct burr, as seen in most materials of Stephanocemas. Both specimens display a relatively flat palm portion of the antler, and even a relatively mediolaterally expanded palm. They also possess a distinct dorsal ridge connecting the anterior and posterior tines. The latter two features are perhaps the most revealing about phylogenetic relationships. Whereas a flat and laterally expanded palm may indicate affinity with S. palmatus, the dorsal ridge, however, seems to imply a more primitive status as seen in species of Paradicrocerus and possibly in S. actauensis [the illustration in Tleuberdina et al. (1993: pl. XI, fig. 2a, b) is not clear enough to be sure of this feature]. Future discoveries may prove that IVPP V15725 and V15726 belong to a distinct species of their own if such a unique combination of primitive and derived characters is confirmed, but because of the poor state of preservations we refrain from naming a new species.Published as part of Wang, Xiaoming, Xie, Guangpu & Dong, Wei, 2009, A new species of crown-antlered deer Stephanocemas (Artiodactyla, Cervidae) from the middle Miocene of Qaidam Basin, northern Tibetan Plateau, China, and a preliminary evaluation of its phylogeny, pp. 680-695 in Zoological Journal of the Linnean Society 156 (3) on pages 688-690, DOI: 10.1111/j.1096-3642.2008.00491.x, http://zenodo.org/record/468777