Kazakhstan Encyclopedia - User contributions [en]

Macroolithus

2017-04-08T00:51:14Z

Plantdrew: taxobox cleanup

{{good article}}
{{oobox
| fossil_range = {{fossil range|99.6|65|[[Late Cretaceous]]}}
| image = TwoMacroo.jpg
| image_caption = A pair of ''Macroolithus yaotunensis'' eggs
| taxon = Macroolithus
| authority = Zhao, 1975
| type_oospecies = ''Oolithes rugustus''
| type_oospecies_authority = Young, 1965
| subdivision_ranks = [[Oospecies]]
| subdivision =
* '''''M. rugustus''''' (Young, 1965)
* '''''M. yaotunensis''''' Zhao, 1975
* '''''M. mutabilis''''' Mikhailov, 1994
* '''''M. lashuyuanensis''''' Fang ''et al.'', 2009
}}

'''''Macroolithus''''' is an [[oogenus]] (fossil-egg genus) of [[dinosaur egg]] belonging to the [[oofamily]] [[Elongatoolithidae]]. The [[Type (biology)|type oospecies]], ''M. rugustus'', was originally described under the now-defunct oogenus name ''[[Oolithes]]''. Three other oospecies are known: ''M. yaotunensis'', ''M. mutabilis'', and ''M. lashuyuanensis''. They are relatively large, elongated eggs with a two-layered eggshell. Their nests consist of large, concentric rings of paired eggs. There is evidence of blue-green pigmentation in its shell, which may have helped camouflage the nests.

''Macroolithus'' eggs have been found containing [[oviraptorid]] [[dinosaur]] [[embryos]] resembling ''[[Heyuannia]]''. Multiple other associations between oviraptorid and elongatoolithids (including other eggs containing embryos, parents brooding on nests, and a pair of shelled ''Macroolithus''-like eggs preserved within an oviraptorid's pelvis) confirm that the parent of ''Macroolithus'' was an oviraptorid.

It is found in [[Upper Cretaceous]] formations of central and eastern Asia; fossils have been found in Mongolia, Kazakhstan and China. In the Nanxiong formation in Southern China, ''Macroolithus'' fossils range up to and possibly over the [[K/T boundary|Cretaceous-Tertiary boundary]], which is traditionally assumed to mark the extinction of the non-[[bird|avian]] dinosaurs. Some paleontologists have interpreted the record of dinosaur eggs at this formation as supporting a gradual extinction event, rather than a sudden cataclysmic event. However, other paleontologists believe that these interpretations are merely based on artifacts of erosion and redeposition in the early [[Paleogene]].

== Description ==
[[File:Academy of Natural Sciences, Philadelphia - IMG 7447.JPG|thumb|right|A pair of ''Macroolithus'' eggs]]
''Macroolithus'' eggs are characterized by large size, measuring {{convert|16|to|21|cm|abbr=on}} long, and by their particularly coarse ornamentation.<ref name="Simon 2014"/><ref name="Mikhailov 1994"/> Their microstructure is not well defined in the literature,<ref name="Simon 2014"/> but generally follows the typical elongatoolithid pattern:<ref name="Mikhailov 1994"/> The eggshell is arranged into two structural layers (the mammillary layer and the continuous layer). The continuous layer forms the outer part of the eggshell; its eggshell units are fused together so that the layer appears to be a continuum. The inner layer, known as the mammillary, or cone, layer is made up of cone-shaped structures that form the base of the eggshell units.<ref name="Simon 2014"/> In ''Macroolithus'', the continuous layer is two to three times thicker than the mammillary layer.<ref name="Simon 2014"/><ref name="Mikhailov 1994"/> The eggs have great diversity of pore structure. Since gas conductance is related to the pore size and density this could imply that the eggs were laid in variable environments.<ref name="Simon 2014" />

=== Oospecies ===
Four oospecies of ''Macroolithus'' are currently recognized:
* '''''M. rugustus''''' – the [[Type (biology)|type oospecies]], these eggs are found in [[Nanxiong]] and [[Jiangxi]] in south China; it is also common in the [[Nemegt Formation]] of [[Mongolia]] and the [[Manrakskaya Svita]] in [[Kazakhstan]].<ref name="Young 1965">{{cite journal|last1=Young|first1=Chung-chien|title=Fossil eggs from Nanhsiung, Kwangtun and Kanchou, Kiangsi|journal=Vertebrata PalAsiatica|date=1965|volume=9|issue=2|pages=141–170|url=http://www.ivpp.cas.cn/cbw/gjzdwxb/xbwzxz/200912/P020091223559165777170.pdf}}</ref><ref name="Mikhailov 2000">{{cite book|last1=Mikhailov|first1=Konstantin|editor1-last=Benton|editor1-first=M.J.|editor2-last=Shishkin|editor2-first=M.A.|editor3-last=Unwin|editor3-first=D.M.|editor4-last=Kurochkin|editor4-first=E.N.|title=The Age of Dinosaurs in Russia and Mongolia|date=2000|publisher=Cambridge University Press|pages=560–572|chapter=Eggs and eggshells of dinosaurs and birds from the Cretaceous of Mongolia}}</ref> It ranges in size from {{convert|16.5|to|18|cm|abbr=on}} long by {{convert|7.5|to|8.5|cm|abbr=on}} across. Its eggshell is {{convert|0.8|to|1.7|mm|abbr=on}} thick, about 1/4 of which is the mammillary layer.<ref name="Zhao 1975">{{cite journal|date=1975|title=The microstructure of the dinosaurian eggshells of Nanxiong, Guangdong Province—on the classification of dinosaur eggs|url=http://www.ivpp.cas.cn/cbw/gjzdwxb/xbwzxz/200905/W020090813377364004471.pdf|journal=Vertebrata PalAsiatica.|volume=13|issue=2|pages=105–117|last1=Zhao|first1=Zikui}}</ref> The outer surface is covered with irregular chains of nodes (ramotuberculate ornamentation) around the outer center of the egg, transitioning to a more pitted, netlike pattern (sagenotuberculate) at the ends.<ref name="Carpenter 1999">{{cite book|last1=Carpenter|first1=K.|title=Eggs, Nests, and Baby Dinosaurs: A Look at Dinosaur Reproduction|date=1999|publisher=Indiana University Press|pages=294–295}}</ref><ref name="How to Study">{{cite book|last1=Carpenter|first1=Kenneth|title=Eggs, Nests, and Baby Dinosaurs: A Look at Dinosaur Reproduction (Life of the Past)|date=1999|publisher=Indiana University Press|location=Bloomington, Indiana|isbn=0-253-33497-7|pages=122–144|chapter=How to Study a Fossil Egg}}</ref>
* '''''M. yaotunensis''''' is native to [[Henan]] and to [[Nanxiong]].<ref name="Liang et al 2009"/><ref name="Zhao 1975"/> It has similar microstructure and ornamentation to ''M. rugustus'', but can be distinguished by being slightly larger ({{convert|17.5|to|21|cm|abbr=on}} long) and having a thicker shell ({{convert|1.4|to|1.9|mm|abbr=on}}).<ref name="Simon 2014">Simon, D. J. (2014). "[http://scholarworks.montana.edu/xmlui/bitstream/handle/1/8693/SimonD0814.pdf?sequence=1 Giant Dinosaur (theropod) Eggs of the Oogenus Macroelongatoolithus (Elongatoolithidae) from Southeastern Idaho: Taxonomic, Paleobiogeographic, and Reproductive Implications.]" (Doctoral dissertation, Montana State University, Bozeman).</ref><ref name="Mikhailov 1994">{{cite journal|last1=Mikhailov|first1=Konstantin K.E.|title=Theropod and protoceratopsian dinosaur eggs from the Cretaceous of Mongolia and Kazakhstan|journal=Paleontological Journal|date=1994|volume=28|issue=2|pages=101–120}}</ref> Also unlike ''M. rugustus'', ''M. yaotunensis'' shows a wavy boundary between its mammillary layer and its continuous layer.<ref name="Zhao 1975"/>
* '''''M. mutabilis''''' – this Mongolian oospecies is known from several eggshell fragments. The fragments range from {{convert|1.3|to|2.0|mm|abbr=on}} thick, slightly thicker than ''M. rugustus''. When complete, the egg probably measured over {{convert|17|cm|abbr=on}}. It is unique for what Mikhailov called "aberrational" features: several fragments have slitlike pore canals and lack ornamentation.<ref name="Mikhailov 1994"/>
* '''''M. lashuyuanensis''''' – this oospecies is based on an eggshell fragment found in the [[Nanxiong Basin]] of [[Guangdong]]. With a thickness of {{convert|2.3|to|2.7|mm|abbr=on}}, its eggshell is the thickest among ''Macroolithus'' oospecies. It is furthermore distinguished by having a dark, dense band near the top of its continuous layer and a gradual transition between the continuous and mammilary layers.<ref name="Fang et al 2009">{{cite journal|last1=Fang|first1=Xiaosi|last2=Li|first2=Peixian|last3=Zhang|first3=Zhijun|last4=Lin|first4=Youli|last5=Guo|first5=Shengbin|last6=Cheng|first6=Yeming|last7=Li|first7=Zhenyu|last8=Zhang|first8=Xiaojun|last9=Cheng|first9=Zhengwu|title=广东南雄白垩系及恐龙蛋到鸟蛋演化研究|journal=Acta Geoscientia Sinica|date=2009|volume=30|issue=2|pages=167–186|url=http://www.oalib.com/paper/1558382#.V-E44TVIkni|trans-title=Cretaceous Strata in Nanxiong Basin of Guangdong and the Evolution from the Dinosaur Egg to the Bird Egg|language=Chinese}}</ref>

=== Coloration ===
In 2015, Wiemann ''et al.'' announced the discovery of pigments preserved in a ''Macroolithus'' eggshell, the first evidence of egg coloration in non-avian dinosaurs.<ref name="Mancini 2015">{{cite web|last1=Mancini|first1=Mark|title=Scientists Reveal Dinosaur Egg Colors for the First Time|url=http://mentalfloss.com/article/64509/scientists-reveal-dinosaur-egg-colors-first-time|website=mental_floss|publisher=Mental Floss|accessdate=28 October 2016}}</ref> The eggshell contained the [[tetrapyrrolic]] pigments [[protoporphyrin]] and [[biliverdin]]. Due to degradation of the pigments, and the comparative stability of biliverdin to protoporphyrin, the original ratios of these chemicals are not preserved. The reddish-brown protoporphyrin is most abundant at the surface of the eggshell, whereas the blue-green biliverdin was found in deeper crystals of the eggshell. While the exact coloration is impossible to determine, this suggests that the overall color ''M. yaotunensis'' was blue or green; Wiemann ''et al.'' suggested that the most likely coloration was olive-green or blue with dark speckles.<ref name="Wiemann et al in press">{{cite journal|last1=Wiemann|first1=Jasmina|last2=Yang|first2=Tzu-Ruei|last3=Sander|first3=N. N. Sander|last4=Schneider|first4=Marion|last5=Engeser|first5=Marianne|last6=Kath-Schorr|first6=Stephanie|last7=Müller|first7=Christa E.|last8=Sander|first8=P. Martin|title=The blue-green eggs of dinosaurs: How fossil metabolites provide insights into the evolution of bird reproduction|journal=PeerJ Preprints|date=2015|volume=3|page=e1323|doi=10.7287/peerj.preprints.1080v1|url=https://peerj.com/preprints/1080/|doi-broken-date=2017-01-16}}</ref>

==Names and etymology==
The name ''Macroolithus'' is derived from the Greek roots ''macro-'' (meaning "large")<ref name="Banay 1948">{{cite journal|last1=Banay|first1=G.L.|title=An introduction to medical terminology i. Greek and Latin derivations|journal=Bulletin of the Medical Library Association|date=1948|volume=36|issue=1|page=12|url=https://www.ncbi.nlm.nih.gov/pmc/articles/PMC194697/pdf/mlab00247-0013.pdf}}</ref> and ''oolithus'' (meaning "stone egg"), the conventional suffix for oogenus names.<ref name="Mikhailov et al 1996">{{cite journal|last1=Mikhailov|first1=Konstantin E.|last2=Bray|first2=Emily S.|last3=Hirsch|first3=Karl F.|title=Parataxonomy of fossil eggs remains (Veterovata): Principles and applications|journal=Journal of Vertebrate Paleontology|date=1996|volume=16|issue=4|pages=763–769|url=https://www.ncbi.nlm.nih.gov/pmc/articles/PMC194697/pdf/mlab00247-0013.pdf}}</ref> Its Chinese name (巨形蛋 Jùxíng dàn) similarly translates to "giant egg".<ref name="Zhao 1975"/> The oospecific epiphets ''yaotunensis'' (Chinese: 窑屯 yáotún) and ''lashuyuanensis'' (Chinese: 腊树园 Làshùyuán) honor the localities where those oospecies were originally discovered (the Yaotun<ref name="Zhao 1975"/> and Lashuyuan<ref name="Fang et al 2009"/> districts of Guangdong, respectively), affixing the Latin suffix ''-ensis'' to denote the place of origin;<ref name="White 1858">{{cite book|last1=White|first1=John Tahourdin|title=Latin suffixes|date=1858|publisher=Spottiswoode and Co.|location=London|page=80|url=https://books.google.com/books?hl=en&lr=&id=I4sCAAAAQAAJ&oi=fnd&pg=PR12&dq=ensis+latin+root&ots=hSb-r84EWv&sig=LOipPbkIcu6h7m5TAxfebTqBGxI#v=onepage&q&f=false}}</ref> the name ''mutabilis'' is Latin for changeable.<ref name="Mikhailov 1994"/> When he named ''"Oolithes" rugustus'', Yang (1965) did not give the etymology of ''rugustus'', but the Chinese form (粗皮 cūpí) translates to "rough skin".<ref name="Young 1965"/>

==Paleobiology and parenting==
[[File:Nesting Nemegtomaia.jpg|thumb|right|Artist's restoration of the oviraptorid ''[[Nemegtomaia]]'' brooding on its nest.]]
Many associations between adult oviraptorosaur skeletons or embryos with elongatoolithid eggs (including ''Macroolithus'') demonstrate that ''Macroolithus'' and other elongatoolithids were laid by oviraptorosaurs.<ref name="Simon 2014"/><ref name="Wang et al 2016"/> One oviraptorosaur skeleton from the [[Upper Cretaceous]] of [[China]] was described in 2005; two shelled elongatoolithid eggs were preserved inside of its pelvis. This suggests that oviraptorosaurs had two functional [[oviduct]]s where both would produce eggs simultaneously. While Sato ''et al.'' did not refer the eggs specifically to ''Macroolithus'', they noted that the eggs closely resemble ''M. yaotunensis'', though with a thinner eggshell. The thin eggshell, however, could simply be because the shell had not finished forming when the mother died, or because of biochemical dissolution of the shell before fossilization.<ref name="Sato et al 2005">{{cite journal|last1=Sato|first1=Tamaki|last2=Cheng|first2=Yen-nien|last3=Wu|first3=Xiao-chun|last4=Zelenitsky|first4=Darla|last5=Hsiao|first5=Yu-fu|title=A Pair of Shelled Eggs Inside A Female Dinosaur|journal=Science|date=2005|volume=308|issue=5720|page=375|doi=10.1126/science.1110578}}</ref>

Multiple different genera of oviraptorids have been found on or near elongatoolithid nests indicating that oviraptorid parents would [[Egg incubation|brood]] on their eggs, most likely for extended periods of time.<ref name="Fanti et al 2012">{{cite journal|last2=Currie|first2=PJ|last3=Badamgarav|first3=D|date=2012|title=New Specimens of ''Nemegtomaia'' from the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia|url=http://journals.plos.org/plosone/article?id=10.1371%2Fjournal.pone.0031330#pone-0031330-g010|journal=PLoS ONE|volume=7|issue=2|page=e31330|doi=10.1371/journal.pone.0031330|last1=Fanti|first1=F|bibcode=2012PLoSO...731330F}}</ref> There is some evidence to suggest that oviraptorid and [[troodontid]] eggs were cared for by the father, perhaps in a [[polygamous]] system.<ref name="Varricchio et al 2008">{{cite journal|last2=Moore|first2=Jason R.|last3=Erickson|first3=Gregory M.|last4=Norell|first4=Mark A.|last5=Jackson|first5=Frankie D.|last6=Borkowski|first6=John J.|date=2008|title=Avian Paternal Care Had Dinosaur Origin|journal=Science|volume=322|issue=5909|pages=1826–1827|last1=Varricchio|first1=David J.|bibcode=2008Sci...322.1826V|doi=10.1126/science.1163245}}</ref><ref name="Wiemann et al in press"/> Given the large size of eggs relative to the parent, a mother would only lay two eggs at a time, so the eggs of a single nest may have been contributed by multiple females.<ref name="Sato et al 2005" /><ref name="Varricchio et al 2008"/>

===Nests===
[[File:OviraptoridNest.jpg|thumb|left|A fossilized nest of oviraptorosaurid eggs]]
Multiple well preserved ''Macroolithus'' nests are known, representing ''M. rugustus'' and ''M. yaotunensis''.<ref name="Young 1965"/><ref name="Zhao 1975"/> A very well preserved clutch of ''M. yaotunensis'' contains 20 eggs arrayed in two, possibly three, circular layers. When complete this nest may have contained 40 or more eggs.<ref name="Young 1965"/>

Tanaka ''et al.'' (2015) found that among modern archosaurs (the group including birds, crocodilians, and dinosaurs), the porosity of eggshells can be used to accurately predict whether the eggs are buried or laid in open nests. They concluded that ''Macroolithus'' nests were either open or only partially covered with vegetation.<ref name="Tanaka et al 2015"/> The eggs' blue-green coloration would act as a camouflage to hide the nest from predators.<ref name="Wiemann et al in press"/>

''M. rugustus'' eggs at [[Tsagan Khushu]] in Mongolia represent a coastal colonial nesting site.<ref name="Mikhailov 1994"/>

===Embryos===
A few ''Macroolithus'' eggs preserve embryonic remains of oviraptorids inside.<ref name="Cheng et al 2008"/> Two eggs containing embryos found in the [[Upper Cretaceous]] [[Nanxiong Formation]] near [[Ganzhou, Jiangxi]] were referred to ''M. yaotunensis'' in 2008. One of these embryos shows a much greater degree of bone development ([[ossification]]) than the other; it preserves ossified hind limbs and several vertebrae. The fact that the cervical vertebrae not only have ossified [[Vertebral centrum|centra]] and [[neural arch]]es, but also have ossified [[zygapophyses]], led Cheng ''et al.'' to conclude that this embryo may in fact represent a hatchling, since in modern [[chicken]]s and [[skua]]s these zygaphyses are cartilaginous until hatching. Also, the embryo's foot anatomy (specifically, the proportions of its [[Metatarsal bones|metatarsal]]s) resembles that of the oviraptorid ''[[Heyuannia huangi]]'', indicating that these eggs belong to ''H. huangi'' or a similar species. The other egg has a much less developed embryo, with only the hind limbs preserved. While this does not permit comparison on the species level, the tibia (shinbone) confirms that it is an oviraptorosaur.<ref name="Cheng et al 2008">{{cite journal|last1=Cheng|first1=Yen-nien|last2=Ji|first2=Qiang|last3=Wu|first3=Xiao-chun|last4=Shan|first4=Hsi-y|title=Oviraptorosaurian Eggs (Dinosauria) with Embryonic Skeletons Discovered for the First Time in China|journal=Acta Geologica Sinica|date=2008|volume=82|issue=6|pages=1089–1094}}</ref>

Three more eggs containing embryos from the same formation were described in 2016 by Wang ''et al.'' Though they noted that these eggs strongly resemble ''M. yaotunensis'', they declined to refer them to any ootaxon lower than Elongatoolithidae because ''Macroolithus'' is not clearly defined and is in need of revision. The embryos within these eggs are some of the most well-preserved of any oviraptorids, providing new information on oviraptorid ontogeny. The specimens show a relatively shallow head which indicates that as oviraptorids matured, their skulls grew dorsoventrally (top-to-bottom) faster than anteroposteriorly (front-to-back). This growth pattern is unusual among theropods, but is also seen in [[Derived trait|derived]] ("advanced") [[tyrannosaurid]]s. Also unusual is that, even at this early stage of development, the [[nasal bone]]s are fused. Coincidentally, tyrannosaurids also show fusion of the nasals early in development.<ref name="Wang et al 2016">{{cite journal|last1=Wang|first1=Shuo|last2=Zhang|first2=Shukang|last3=Sullivan|first3=Corwin|last4=Xu|first4=Xing|title=Elongatoolithid eggs containing oviraptorid (Theropoda, Oviraptorosauria) embryos from the Upper Cretaceous of Southern China|journal=BMC Evolutionary Biology|date=2016|volume=16|issue=67|doi=10.1186/s12862-016-0633-0}}</ref>

===Pathology and extinction===
Many ''Macroolithus'' specimens in South China have double- or multiple-layering of cones on the inner surface of the eggshell, a [[Egg paleopathology|pathological condition]] known as ovum in ovo.<ref name="Zhao 1994"/> It is especially prevalent among eggs nearest to the [[K-T boundary|Cretaceous-Tertiary (K-T) boundary]], which represents the end of the [[Mesozoic Era]]. This pathology is correlated with a higher concentration of [[trace element]]s like [[Cobalt|Co]], [[Chromium|Cr]], [[Copper|Cu]], [[Manganese|Mn]], [[Nickel|Ni]], [[Lead|Pb]], [[Strontium|Sr]], [[Vanadium|V]], and [[Zinc|Zn]].<ref name="Zhao 1994">{{cite book|last1=Zhao|first1=Zi-kui|editor1-last=Carpenter|editor1-first=Kenneth|editor2-last=Hirsch|editor2-first=Karl F.|editor3-last=Horner|editor3-first=John R.|title=Dinosaur Eggs and Babies|date=1994|publisher=Cambridge University Press|isbn=0-521-44342-3|pages=184–203|chapter=Dinosaur eggs in China: On the structure and evolution of eggshells}}</ref> Experiments on modern birds have demonstrated that exposure to high levels of these elements will cause them to be incorporated into the eggshell, but the precise mechanism behind the pathological multi-layering is unknown. These abnormalities presumably affected hatchability of the eggs and may have played a role in the extinction of the dinosaurs.<ref name="Wang et al 2016"/><ref name="Zhao 1994"/> However, the embryonic remains inside three multilayered ''Macroolithus''-like eggs from Ganzhou appear unaffected.<ref name="Wang et al 2016"/>

The K-T boundary is associated with [[iridium anomaly|abnormally high amounts of iridium]], an element which is rare in Earth's crust, but relatively common in asteroids and in Earth's core. This has been used as evidence that a meteorite impact caused the extinction of the dinosaurs (the [[Alvarez hypothesis]]). However, some paleontologists attribute the extinction event and the iridium anomaly to more gradual climatic change caused by the volcanic activity of the [[Deccan Traps]]. The Nanxiong Basin has special relevance to this discussion because it contains the K-T boundary. However, the position of boundary, as well as the duration of the extinction have been subject to debate. Zhao ''et al.'' (2002 and 2009) have postulated that there were at least two iridium-delivering events over a time period consistent with the Deccan Traps volcanism, correlated with a gradual decline in diversity and eventual disappearance of fossil eggs from the Nanxiong Basin. According to their interpretation, ''Macroolithus'' survived 250,000 years into the early [[Paleocene]].<ref name="Zhao et al 2002">{{cite journal|last1=Zhao|first1=Zikui|last2=Mao|first2=Xueming|last3=Chai|first3=Zhifang|last4=Yang|first4=Gaochuang|last5=Kong|first5=Ping|last6=Ebihara|first6=Mitsuru|last7=Zhao|first7=Zhenhua|title=A possible causal relationship between extinction of dinosaurs and K/T iridium enrichment in the Nanxiong Basin, South China: evidence from dinosaur eggshells|journal=Palaeogeography, Palaeoclimatology, Palaeoecology|date=2002|volume=178|issue=1–2|pages=1–17|doi=10.1016/S0031-0182(01)00361-3|url=http://www.sciencedirect.com/science/article/pii/S0031018201003613}}</ref><ref name="Zhao et al 2009">{{cite journal|last1=Zhao|first1=Zikui|last2=Mao|first2=Xueying|last3=Chai|first3=Zhifang|last4=Yang|first4=Gaochuang|last5=Zhang|first5=Fucheng|last6=Yan|first6=Zheng|title=Geochemical environmental changes and dinosaur extinction during the Cretaceous-Paleogene (K/T) transition in the Nanxiong Basin, South China: Evidence from dinosaur eggshells|journal=Chinese Science Bulletin|date=2009|volume=54|issue=5|page=806|doi=10.1007/s11434-008-0565-1|url=http://link.springer.com/article/10.1007/s11434-008-0565-1}}</ref><ref name="Wang et al 2010">{{cite journal|last1=Wang|first1=Qiang|last2=Wang|first2=Xiaolin|last3=Zhao|first3=Zikui|title=Recent Progress in the Study of Dinosaur Eggs in China|journal=Bulletin of the Chinese Academy of Sciences|date=2010|volume=24|issue=2|pages=95–97|url=http://english.cas.cn/bcas/2010_2/201411/P020141121535002835194.pdf}}</ref> However, Buck ''et al.'' (2004) disputed these claims, arguing that [[debris flow]]s mixed and reworked the [[Sedimentary rock|sediments]] near the boundary, causing a blurring of the iridium anomaly and the appearance of eggshell fragments on both sides.<ref name="Buck et al 2004"/>

==Classification==
{{see also|Egg fossil#Classification}}
Fossilized eggs are classified in their own, parataxonomic system parallel to [[Linnaean taxonomy]]. ''Macroolithus'' and ''Elongatoolithus'' were the first oogenera ever named in this system; they were classified in the oofamily [[Elongatoolithidae]].<ref name="Zhao 1975"/> Cladistic analysis also supports the placement of ''Macroolithus'' close to ''Elongatoolithus'', together with other elongatoolithids in a [[clade]].<ref name=VB2015>{{cite journal|last1=Varricctio|first1=David J.|last2=Barta|first2=Daniel E.|title=Revistiting Sabath's "Larger Avian Eggs" from the Gobi Cretaceous|journal=Acta Palaeontologica Polonica|date=2015|volume=60|issue=1|pages=11–25|doi=10.4202/app.00085.2014|url=https://www.app.pan.pl/article/item/app000852014.html}}</ref><ref name="Garcia et al 2006">{{cite journal|last1=Garcia|first1=Geraldine|last2=Marivaux|first2=Laurent|last3=Pélissié|first3=Thierry|last4=Vianey-Liaud|first4=Monique|title=Earliest Laurisian sauropod eggshell|journal=Acta Paleontologica Polonica|date=2006|volume=51|issue=1|pages=99–104|url=https://www.app.pan.pl/archive/published/app51/app51-099.pdf}}</ref> Four oospecies are known: ''M. rugustus'', ''M. yaotunensis'', ''M. mutabilis'', and ''M. lashuyuanensis''.<ref name="Simon 2014"/><ref name="Moreno-Azanza et al 2014">Moreno-Azanza, M., J.I. Canudo, and J.M. Gasca. (2014) "[https://www.researchgate.net/profile/Jose_Canudo/publication/263507846_Spheroolithid_eggshells_in_the_Lower_Cretaceous_of_Europe._Implications_for_eggshell_evolution_in_ornithischian_dinosaurs/links/54c7c8550cf238bb7d0b43a5.pdf Spheroolithid eggshells in the Lower Cretaceous of Europe. Implications for eggshell evolution in ornithischian dinosaurs.]" ''[[Cretaceous Research]]'' '''51''': 75–87.</ref> The oogenus's microstructures are poorly-defined<ref name="Simon 2014"/> and therefore may be in need of revision.<ref name="Wang et al 2016"/> When Mikhailov described ''M. mutabilis'', he found no microstructural difference in Zhao's original illustrations of ''M. rugustus'' and ''M. yaotunensis'', but did not [[Synonym (taxonomy)|synonymize]] the oospecies because Chinese paleontologists considered them distinct.<ref name="Mikhailov 1994"/>

==History==
{{see also|Timeline of egg fossil research}}

===Discoveries===
''Macroolithus'' eggs were first discovered in Southern China by the pioneering Chinese paleontologist [[Yang Zhongjian]]. He described the remains of several fossil eggs from that region in 1965. Working prior to the advent of modern fossil egg parataxonomy, he gave them names as species of ''[[Oolithes]]'',<ref name="Young 1965"/> a now-defunct name that was formerly used for various types of fossil eggs.<ref name="Lawver and Jackson 2015">{{cite journal|last1=Lawver|first1=Daniel R.|last2=Jackson|first2=Frankie D.|title=A Review of the Fossil Record of Turtle Reproduction: Eggs, Embryos, Nests and Copulating Pairs|journal=Bulletin of the Peabody Museum of Natural History|date=2015|volume=55|issue=2|pages=215–236|doi=10.3374/014.055.0210|url=http://scholarworks.montana.edu/xmlui/bitstream/handle/1/9308/Lawver_Jackson_BPMNH_10_2014_A1b.pdf?sequence=1}}</ref><ref name="VLM2003">{{cite journal|last1=Vianey-Liaud|first1=Monique|last2=Zelenitsky|first2=Darla|title=Historical and new perspectives on the parataxonomy of fossil eggs|journal=Paleovertebrata|date=2003|volume=32|issue=2-4|pages=189–195}}</ref> In 1975, Chinese paleontologist Zhao Zikui prototyped the modern parataxonomic system, creating a hierarchical system of oofamilies, oogenera, and oospecies. Zhao placed Yang's ''O. rugustus'' into the new oogenus, ''Macroolithus'', splitting it into two oospecies, ''M. yaotunensis'' and ''M. rugustus''. He also suggested that the American oospecies ''"Oolithes" carlylensis'' (which is now classified in a different elongatoolithid oogenus, ''[[Macroelongatoolithus]]''<ref name="Simon 2014"/>) be recombined as ''Macroolithus carlylensis''.<ref name="Zhao 1975"/>

In 1991, the Russian paleontologist Konstantin Mikhailov introduced the modern classification of fossil eggs based on Zhao's parataxonomic naming system. He classified ''"O." carlylensis'' in the oofamily [[Spheroolithidae]], but otherwise followed Zhao's 1975 classification of ''Macroolithus''.<ref name="Mikhailov 1991">{{cite journal|last1=Mikhailov|first1=Konstantin E.|title=Classification of Fossil Eggshells of Amniotic Vertebrates|journal=Acta Palaeontologica Polonica|date=1991|volume=36|issue=2|pages=193–238|url=https://www.app.pan.pl/archive/published/app36/app36-193.pdf}}</ref> In 1994 he named ''M. mutabilis'', a new oospecies of ''Macroolithus'', based on remains discovered in Mongolia.<ref name="Mikhailov 1994"/>

In 2000, a fourth oospecies, ''"M." turolensis'', was described by Spanish paleontologists Olga Amo-Sanjuán, José Ignacio Canudo, and Gloria Cuenca-Bescós based on material from Spain. However, when new material of this oospecies was uncovered in 2014, it was moved into its own oogenus, ''[[Guegoolithus]]'', which was furthermore moved to Spheroolithidae.<ref name="Moreno-Azanza et al 2014"/>

In 2005, eggs possibly attributable to ''M. yaotunensis'' were discovered paired inside the pelvis of a fossil oviraptorid skeleton.<ref name="Sato et al 2005"/> In 2008, Chinese paleontologists Cheng Yen-nien, Ji Qiang, Wu Xiao-chun and Shan Hsi-yin discovered a pair of eggs representing the first in China to contain embryonic remains of oviraptorosaurs. Both eggs were referred to ''M. yaotunensis''.<ref name="Cheng et al 2008"/> The fourth oospecies, ''M. lashuyuanensis'', was described in 2009 by the Chinese paleontologists Fang Xiao-si, Li Pei-xian, Zhang Zhi-jun, Zhang Xian-qiu, Lin You-li, Guo Sheng-bin, Cheng Ye-ming, Li Zhen-yu, Zhang Xiao-jun and Cheng Zheng-wu.<ref name="Fang et al 2009"/>

===Paleobiological interpretations===
[[File:Heyuannia profile0.jpg|thumb|Restoration of ''Heyuannia'', the putative parent of ''Macroolithus'']]
In 1994, Mikhailov suggested that ''Macroolithus'' represented the eggs of a large theropod, specifically ''[[Tarbosaurus]]'', based on the large size of the eggs and the largely overlapping distribution.<ref name="Mikhailov 1994"/> However, the discovery of an oviraptorid embryo inside an elongatoolithid egg cast doubt on this hypothesis.<ref name="Mikhailov 1997"/> With the discovery of multiple oviraptorosaur-elongatoolithid associations in the late 1990s,<ref name="Zelenitsky et al 2000">{{cite journal|last1=Zelenitsky|first1=Darla K.|last2=Carpenter|first2=Kenneth|last3=Currie|first3=Philip J.|title=Fist Record of Elongatoolithus Theropod Eggshell from North America: The Asian oogenus ''Macroelongatoolithus'' from the lower Cretaceous of Utah|journal=Journal of Vertebrate Paleontology|volume=20|issue=1|pages=130–138|doi=10.1671/0272-4634(2000)020[0130:FROETE]2.0.CO;2|url=http://www.bioone.org/doi/full/10.1671/0272-4634(2000)020%5B0130:FROETE%5D2.0.CO%3B2|year=2000}}</ref> the eggs of elongatoolithids came to be accepted as belonging to oviraptorosaurian dinosaurs.<ref name="Simon 2014"/> Oviraptorid parentage was confirmed for ''Macroolithus'' when, in 2008, oviraptorid embryos resembling ''Heyuannia'' were discovered inside a pair of ''M. yaotunensis'' eggs in Jiangxi.<ref name="Cheng et al 2008"/>

Early on, paleontologists considered ''Macroolithus'' nests to have been buried. Mou 1992 noted the high gas conductance values in ''Macroolithus'' eggs and therefore concluded that they were laid in a very humid environment, buried underground or inside a mound.<ref name="Mou 1992">{{cite journal|last1=Mou|first1=Yun|title=Nest Environments of the Late Cretaceous Dinosaur eggs from Nanxiong Basin, Guangdong Province|journal=Vertebrata PalAsiatica|date=1992|volume=30|issue=2|pages=120–134|url=http://www.ivpp.cas.cn/cbw/gjzdwxb/xbwzxz/200812/W020090813371003110009.pdf}}</ref> Deeming (2006) found a similar result. This seems to contradict evidence that oviraptorosaurids brooded bird-like on their eggs, but Deeming suggested that ''Oviraptor'' buried its eggs in a mound and then ''Oviraptor'' and its relatives sat atop a nest mound to incubate, rather than directly contacting the eggs.<ref name="Deeming 2006">{{cite journal|last1=Deeming|first1=D.C.|title=Ultrastructural and Functional Morphology of Eggshells Supports the Idea that Dinosaur Eggs were Incubated Buried in a Substrate|journal=Paleontology|date=2006|volume=49|issue=1|pages=171–185|doi=10.1111/j.1475-4983.2005.00536.x|url=http://onlinelibrary.wiley.com/doi/10.1111/j.1475-4983.2005.00536.x/abstract}}</ref> However, Tanaka ''et al.'' (2015) criticized these results for lack of statistical rigor. They found, based on comparisons to modern eggs, that ''Macroolithus'' was predicted to be laid in open or partially covered nests.<ref name="Tanaka et al 2015">{{cite journal|last1=Tanaka|first1=Kohei|last2=Zelenitsky|first2=Darla K.|last3=Therrien|first3=François|title=Eggshell Porosity Provides Insight on Evolution of Nesting in Dinosaurs|journal=PLoS ONE|date=2015|volume=10|issue=11|page=e0142829|doi=10.1371/journal.pone.0142829|url=http://journals.plos.org/plosone/article?id=10.1371/journal.pone.0142829|bibcode=2015PLoSO..1042829T}}</ref> An as-of-yet unreviewed preprint by Wiemann ''et al.'' also criticized Mou and Deeming because they had only measured eggshell porosity at the middle section of the eggs and did not take into account the fact that the pore density is much lower near the poles. This would lead to an overestimate of the total eggshell porosity and therefore an overstimate of the gas conductance value.<ref name="Wiemann et al in press"/>

The extinctions of Macroolithus and other eggs from Southern China have also had a history of different interpretations. In the 1990s, Chinese paleontologists, including the prominent egg specialist Zhao Zikui, observed a gradual reduction in dinosaur egg diversity during the final 200,000 to 300,000 years of the Cretaceous, with only ''Macroolithus'' ranging up to the boundary.<ref name="Zhao 1994"/><ref name="Zhao et al 1991"/> They postulated, contrary to the impact hypothesis, that the extinction was the result of a prolonged drought that increased the concentration of trace heavy metals, which adversely affected eggshell and embryo development of the dinosaurs causing the population to gradually decline and collapse.<ref name="Zhao et al 1991"/> Zhao ''et al.'' revised this hypothesis in 2002, postulating a gradual extinction of ''Macroolithus'' caused by the volcanism of the Deccan Traps.<ref name="Zhao et al 2002"/> In 2004, Buck ''et al.'' disputed this interpretation, arguing that the apparent gradual extinction was an illusion caused by reworking of sediments.<ref name="Buck et al 2004"/> Zhao ''et al.'' (2009) maintained that the extinction event was gradual.<ref name="Zhao et al 2009"/>

== Distribution and paleoecology ==
''Macroolithus'' is known from myriad Late Cretaceous locations in China, Mongolia, and Kazakhstan.<ref name="Simon 2014"/>

=== Henan ===
In [[Henan]], ''Macroolithus yaotunensis'' coexists with the other elongatoolithids ''Elongatoolithus andrewsi'' and ''E. elongatus'', as well as ''[[Ovaloolithus]]'' and ''[[Paraspheroolithus]]'' of the [[Hugang Formation|Hugang]], the [[Luyemiao Formation|Luyemiao]], and the [[Sigou Formation]]s. These formations were formed during the Late Cretaceous in a [[lacustrine]] or [[palustrine]] environment. Dinosaur body fossils are rarely found in the same units, but [[troodontid]]s, tyrannosaurs, and [[hadrosaur]]s are known from the same area.<ref name="Liang et al 2009">{{cite journal|last1=Liang|first1=Xinquan|last2=Wan|first2=Shunü|last3=Yang|first3=Dongsheng|last4=Zhou|first4=Shiquan|last5=Wu|first5=Shichong|title=Dinosaur eggs and dinosaur egg-bearing deposits (Upper Cretaceous) of Henan Province, China: Occurrences, palaeoenvironments, taphonomy and preservation|journal=Progress in Natural Science|date=2009|volume=19|issue=11|pages=1587–1601|doi=10.1016/j.pnsc.2009.06.012|url=http://www.sciencedirect.com/science/article/pii/S1002007109002949}}</ref>

=== Shandong ===
Some eggs tentatively assigned to ''Macroolithus'' are known from the [[Wangshi Group]] in [[Laiyang]], [[Shandong]].<ref name="Zhao 1979">{{cite journal|last1=Zhao|first1=Zikui|title=Progress in the Research of Dinosaur Eggs|journal=Mesozoic and Cenozoic Red Beds of South China|date=1979|pages=330–340|url=http://paleoglot.org/files/zhao_79.pdf|series=Selected Papers from the "Cretaceous-Tertiary Workshop," Nanxiong, Guangdong Province|publisher=Institute of Vertebrate Paleontology, Paleoanthropology & Nanjing Institute of Paleontology Science Press,}}</ref> The formations of the Wangshi group were deposited in [[alluvial fan]]s, [[braided channel]]s and shallow lakes. Common fossils include hadrosaurids and dinosaur eggs.<ref name="Wang et al 2013">{{cite journal|last1=Wang|first1=Qiang|last2=Wang|first2=Xiaolin|last3=Zhao|first3=Zikui|last4=Zhang|first4=Jialiang|last5=Jiang|first5=Shunxing|title=New turtle egg fossil from the Upper Cretaceous of the Laiyang Basin, Shandong Province, China|journal=Anais da Academia Brasileira de Ciências|date=2013|volume=85|issue=1|doi=10.1590/S0001-37652013000100008|url=http://www.scielo.br/scielo.php?pid=S0001-37652013000100103&script=sci_arttext}}</ref>

=== Southern China ===
''M. rugustus'', ''M. yaotunensis'', and ''M. lashuyuanensis'' are all known from the [[Nanxiong Basin]] in [[Guangdong]].<ref name="Zhao 1975"/><ref name="Fang et al 2009"/> This formation was deposited primary from streams, rivers, and lakes in that region.<ref name="Zhao et al 2002"/><ref name="Zhao et al 1991">{{cite journal|last1=Zhao|first1=Zikui|last2=Ye|first2=Jie|last3=Li|first3=Huamei|last4=Zhao|first4=Zhenhua|last5=Yan|first5=Zheng|title=Extinction of the dinosaurs across the Cretaceous-Tertiary Boundary in Nanxiong Basin, Guangdong Province|journal=Acta Paleontologica Sinica|date=1991|volume=29|issue=1|pages=1–20|url=http://www.ivpp.cas.cn/cbw/gjzdwxb/xbwzxz/200812/W020090813371326742778.pdf}}</ref> It spans across the [[Cretaceous-Paleogene boundary|K/T boundary]], at which point most of the dinosaur eggs disappear, but ''Macroolithus'' apparently exists on both sides of the boundary (which would imply that some species of dinosaurs survived into the early Tertiary).<ref name="Zhao et al 2002"/><ref name="Zhao et al 2009"/> However, sedimentological evidence suggests that these fossils were actually reworked by debris flows into the Tertiary rocks.<ref name="Buck et al 2004">{{cite journal|last1=Buck|first1=Brenda J.|last2=Hanson|first2=Andrew D.|last3=Hengst|first3=Richard A.|last4=Hu|first4=Shu-sheng|title="Tertiary Dinosaurs" in the Nanxiong Basin, Southern China, Are Reworked from the Cretaceous|journal=Journal of Geology|date=2004|volume=112|issue=1|pages=111–118|jstor=379695}}</ref>

The Nanxiong Basin is known for its abundance of fossil eggs, predominantly the oviraptorisaurian<ref name="Simon 2014"/> eggs ''[[Elongatoolithus]]'' and ''Macroolithus''.<ref name="Zhao et al 2002"/> Other types of eggs include other elongatoolithids, as well as [[Prismatoolithidae|prismatoolithids]], [[Megaloolithidae|megaloolithids]], and [[Ovaloolithidae|ovaloolithids]].<ref name="Zhao et al 2009"/> Footprints show that Nanxiong Basin was populated by [[Ornithopoda|ornithopods]], [[Theropoda|theropods]], and possibly [[Sauropoda|sauropods]].<ref name="Xing et al 2009">{{cite journal|last1=Xing|first1=Lida|last2=Harris|first2=Jerald D|last3=Dong|first3=Zhiming|last4=Lin|first4=Youli|last5=Chen|first5=Wei|last6=Guo|first6=Shengbin|last7=Ji|first7=Qiang|title=Ornithopod (Dinosauria: Ornithischia) tracks from the Upper Cretaceous Zhutian Formation in the Nanxiong basin, Guangdong, China and general observations on large Chinese ornithopd footprints|journal=Geological Bulletin of China|date=2009|volume=28|issue=7|pages=829–843|url=http://www.xinglida.net/pdf/Xing%20et%20al%202009-Hadrosauropodus%20nanxiongensis%20from%20Nanxiong%20Basin.pdf}}</ref>

Southern China, particularly the [[Nanxiong Formation]] in [[Jiangxi|Jiangxi Province]], possibly has the greatest oviraptorosaur diversity in the world,<ref name="Lu et al 2015">{{cite journal|last1=Lü|first1=Junchang|last2=Pu|first2=Hanyong|last3=Kobayashi|first3=Yoshitsugu|last4=Xu|first4=Li|last5=Chang|first5=Huali|last6=Shang|first6=Yuhua|last7=Liu|first7=Di|last8=Lee|first8=Yuong-Nam|last9=Kundrát|first9=Martin|last10=Shen|first10=Caizhi|title=A New Oviraptorid Dinosaur (Dinosauria: Oviraptorosauria) from the Late Cretaceous of Southern China and Its Paleobiogeographical Implications|journal=Scientific Reports|date=2015|volume=5|pages=11490|doi=10.1038/srep11490|pmid=26133245|pmc=4489096|url=http://www.nature.com/articles/srep11490|bibcode=2015NatSR...511490L}}</ref> and also includes several associations of oviraptorosaurs with ''Macroolithus'' or similar eggs.<ref name="Cheng et al 2008"/><ref name="Sato et al 2005"/><ref name="Wang et al 2016"/> [[Titanosaur]]s and [[tyrannosaurid]]s are also known from this area.<ref name="Wang et al 2016"/> Non-dinosaur fauna includes lizards and the terrestrial [[Nanhsiungchelyidae|nanhsiungchelyid]] turtles.<ref name="Tong and Mo 2010">{{cite journal|last1=Tong|first1=Haiyan|last2=Mo|first2=Jinyou|title=Jiangxichelys, a new nanhsiungchelyid turtle from the Late Cretaceous of Ganzhou, Jiangxi Province, China|journal=Geological Magazine|date=2010|volume=147|issue=6|pages=981–986|doi=10.1017/S0016756810000671}}</ref>

=== Mongolia ===
''Macroolithus rugustus'' is abundant in the [[Nemegt Formation]], which dates to the late [[Campanian]] to early [[Maastrichtian]].<ref name="Mikhailov 1995">{{cite journal|last1=Mikhailov|first1=Konstantin|title=Systematic, Faunistic and Stratigraphical Diversity of Cretaceous Eggs in Mongolia: Comparison with China|journal=Sixth Symposium on Mesozoic Terrestrial Ecosystems and Biota, Short Papers|date=1995|pages=165–168|url=https://www.researchgate.net/profile/Konstantin_Mikhailov2/publication/281406429_Systematic_faunistic_and_stratigraphic_diversity_of_Cretaceous_eggs_in_Mongolia_comparison_with_China/links/55e59e4f08aecb1a7ccc29e0.pdf}}</ref><ref name="Weishampel et al 2008"/> This formation represents depositions of a meandering river.<ref name="Weishampel et al 2008">{{cite journal|last1=Weishampel|first1=David B.|last2=Fastovsky|first2=David E.|last3=Watabe|first3=Mahito|last4=Varricchio|first4=David|last5=Jackson|first5=Frankie|last6=Tsogtbaatar|first6=Khishigjav|last7=Barsbold|first7=Rinchen|title=New Oviraptorid Embryoes from Bugin Tsav, Nemegt Formation (Upper Cretaceous), Mongolia, with Insights into Their Habitat and Crowth|journal=Journal of Vertebrate Paleontology|date=2008|volume=28|issue=4|pages=1110–1119|doi=10.1671/0272-4634-28.4.1110|url=http://www.bioone.org/doi/full/10.1671/0272-4634-28.4.1110}}</ref> Well-preserved dinosaur remains are common in the Nemegt Formation,<ref name="Clarke and Norell 2004">{{cite journal|last1=Clarke|first1=Julia A.|last2=Norell|first2=Mark A.|title=New Avialan Remains and a Review of the Known Avifauna from the Late Cretaceous Nemegt Formation of Mongolia|journal=American Museum Novitates|volume=3447|pages=1–12|doi=10.1206/0003-0082(2004)447<0001:NARAAR>2.0.CO;2|url=http://www.bioone.org/doi/full/10.1206/0003-0082(2004)447%3C0001%3ANARAAR%3E2.0.CO%3B2|year=2004}}</ref> including oviraptorosaurs,<ref name="Weishampel et al 2008"/> [[titanosaur]]s,<ref name="Wilson 2004">{{cite journal|last1=Wilson|first1=Jeffrey A.|title=Redescription of the Mongolian Sauropod ''Nemegrosaurus mongoliensis'' Nowinski (Dinosauria: Saurischia) and comments on Late Cretaceous Sauropod diversity|journal=Journal of Systematic Paleontology|date=2005|volume=3|issue=3|pages=283–318|doi=10.1017/S1477201905001628}}</ref> troodontids, tyrannosaurs, ankylosaurs,<ref name="Arbour et al 2014">{{cite journal|last1=Arbour|first1=Victoria M.|last2=Currie|first2=Philip J.|last3=Badamgarav|first3=Demchig|title=The ankylosaurid dinosaurs of the Upper Cretaceous Baruungoyot and Nemegt formations of Mongolia|journal=Zoological Journal of the Linnean Society|date=2014|volume=172|pages=631–652|doi=10.1111/zoj.12185|url=http://s3.amazonaws.com/academia.edu.documents/39611066/ArbourCurrieBadamgarav_Mongolian_ankylosaurs.pdf?AWSAccessKeyId=AKIAIWOWYYGZ2Y53UL3A&Expires=1485900444&Signature=D8hzZcUyRaUqUjCFRrAd1Ihhd8Y%3D&response-content-disposition=inline%3B%20filename%3DThe_ankylosaurid_dinosaurs_of_the_Upper.pdf}}</ref> [[pachycephalosaur]]s,<ref name="Evans et al 2011">{{cite journal|last1=Evans|first1=David C.|last2=Brown|first2=Caled Marshall|last3=Ryan|first3=Michael J.|last4=Tsogtbaatar|first4=Khishigjav|title=Cranial Ornamentation and Ontogenetic Status of Homalocephale calathocercos (Ornithischia: Pachycephalosauria) from the Nemegt Formation, Mongolia|journal=Journal of Vertebrate Paleontology|date=2011|volume=31|issue=1|pages=84–92|doi=10.1080/02724634.2011.546287|url=http://www.tandfonline.com/doi/abs/10.1080/02724634.2011.546287}}</ref> hadrosaurs,<ref name="Maryanska and Osmolska 1982">{{cite journal|last1=Maryanska|first1=Teresa|last2=Osmolska|first2=Halszka|title=Firth Lambeosaurine Dinosaur from the Nemegt Formation, Upper Cretaceous, Mongolia|journal=Acta Paleontologica Polonica|date=1982|volume=26|issue=3/4|pages=243–255|url=http://s3.amazonaws.com/academia.edu.documents/39611066/ArbourCurrieBadamgarav_Mongolian_ankylosaurs.pdf?AWSAccessKeyId=AKIAIWOWYYGZ2Y53UL3A&Expires=1485900444&Signature=D8hzZcUyRaUqUjCFRrAd1Ihhd8Y%3D&response-content-disposition=inline%3B%20filename%3DThe_ankylosaurid_dinosaurs_of_the_Upper.pdf}}</ref> [[ornithomimosaur]]s, [[alvarezsaur]]s,<ref name="Clarke and Norrell 2004"/> and [[therizinosaur]]s.<ref name="Bell et al 2012">{{cite journal|last1=Bell|first1=Phil R.|last2=Currie|first2=Philip J.|last3=Lee|first3=Yuong-Nam|title=Tyrannosaur feeding traces on ''Deinocheirus'' (Theropod:?Ornithomimosauria) remains from the Nemegt Formation (Late Cretaceous), Mongolia|journal=Cretaceous Research|date=2012|volume=37|pages=186–190|doi=10.1016/j.cretres.2012.03.018|url=https://www.researchgate.net/profile/Phil_Bell/publication/257047934_Tyrannosaur_feeding_traces_on_Deinocheirus_TheropodaOrnithomimosauria_remains_from_the_Nemegt_Formation_Late_Cretaceous_Mongolia/links/02e7e525dc05430fd6000000.pdf}}</ref> Remains of small animals are relatively rare, but several types of birds are known from Nemegt,<ref name="Clarke and Norrell 2004">{{cite journal|last1=Clarke|first1=Julia A.|last2=Norrell|first2=Mark A.|title=New Avialan Remains and a Review of the Known Avifauna from the Late Cretaceous Nemegt Formation of Mongolia|journal=American Museum Novitates|date=2004|volume=3447|pages=1–12|doi=10.1206/0003-0082(2004)447<0001:NARAAR>2.0.CO;2}}</ref> as well as several types of [[multituberculate]] [[mammals]].<ref name="Wible and Rougier 2000">{{cite journal|last1=Wible|first1=John R.|last2=Rougier|first2=Guillermo W.|title=Cranial Anatomy of ''Kryptobaatar dashzevegi'' (Mammalia, Multituberculara) and its bearing on the evolution of mammalian characters|journal=Bulletin of the American Museum of Natural History|date=2000|volume=247|page=6|url=http://digitallibrary.amnh.org/bitstream/handle/2246/1594/B247.pdf?sequence=1}}</ref> Other fossil eggs from the Nemegt Formation include ''[[Ovaloolithus]]'', ''[[Spheroolithus]]'', ''Elongatoolithus'', and ''[[Laevisoolithus]]''.<ref name="Mikhailov 1995"/>

''M. mutabilis'' is known solely at the Ikh-Shunkht locality from the [[Barun Goyot Formation]], dating from the [[Santonian]] to [[Campanian]].<ref name="Mikhailov 1995"/><ref name="Mikhailov 1994"/> ''M. rugustus'' is also known from the Barun Goyot Formation, which represents a sand-dune filled [[Aeolian processes|eolian]] environment. Compared to the Nemegt Formation, large dinosaurs are rare at Barun Goyot, where the fauna is dominated by protoceratopsids, oviraptorids, and ankylosaurids.<ref name="Longrich et al 2010">{{cite journal|last1=Longrich|first1=Nicholas R.|last2=Currie|first2=Philip J.|last3=Dong|first3=Zhi-Ming|title=A new oviraptorid (Dinosauria: Theropoda) from the Upper Cretaceous of Bayan Mandahu, Inner Mongolia Authors|journal=Paleontology|date=2010|volume=53|issue=5|pages=945–960|doi=10.1111/j.1475-4983.2010.00968.x|url=http://onlinelibrary.wiley.com/doi/10.1111/j.1475-4983.2010.00968.x/full}}</ref> Other types of fossil eggs from the Barun Goyot Formation include ''[[Protoceratopsidovum]]'', ''[[Gobioolithus]]'', ''[[Faveoloolithus]]'', ''[[Dendroolithus]]'', ''Spheroolithus'', and ''[[Subtiliolithus]]''.<ref name="Mikhailov 1995"/>

=== Kazakhstan ===
''M. rugustus'' has also been found in the [[Manrak Formation]] (also called Manrakskaya Svita) of the [[Zaisan Basin]] in the [[East Kazakhstan Region]]. This formation is near to the [[Tayzhuzgen River]], and dates to some time in the late Cretaceous, probably the Maastrictian.<ref name="Mikhailov 1994"/><ref name="Mikhailov 2000"/><ref name="Mikhailov 1997">{{cite journal|last1=Mikhailov|first1=K.E.|title=Fossil and Recent Eggshell in Amniotic Vertebrates: Fine Structure, Comparative Morphology and Classification|journal=Special Papers in Paleontology|date=1997|volume=56|page=64|url=https://www.researchgate.net/profile/Konstantin_Mikhailov2/publication/287876162_Fossil_and_recent_eggshell_in_amniotic_vertebrates_Fine_structure_comparative_morphology_and_classification/links/56a7537908ae997e22bbd4b8.pdf}}</ref><ref name="Averianov 2007">{{cite journal|last1=Averianov|first1=Alexander O.|title=Theropod dinosaurs from Late Cretacoues depositys in the northeastern Aral Sea region, Kazakhstan|journal=Cretaceous Research|date=2007|volume=28|pages=532–544|doi=10.1016/j.cretres.2006.08.008|url=http://s3.amazonaws.com/academia.edu.documents/41414265/Theropod_dinosaurs_from_the_Late_Cretace20160122-17380-1ch9jvr.pdf?AWSAccessKeyId=AKIAIWOWYYGZ2Y53UL3A&Expires=1485836116&Signature=CwUr8kvBiAgzlkXW8c15LJjlzcc%3D&response-content-disposition=inline%3B%20filename%3DTheropod_dinosaurs_from_the_Late_Cretace.pdf}}</ref>

==See also==
{{Portal|Dinosaurs|Paleontology|||}}
* [[List of dinosaur oogenera]]
* [[Paleontology in China]]
* [[Paleontology in Mongolia]]
* [[Dinosaur reproduction]]

==References==
{{Reflist|2}}

{{taxonbar}}

[[Category:Fossils of China]]
[[Category:Ootaxa described in 1975]]
[[Category:Elongatoolithids]]
[[Category:Fossils of Mongolia]]
[[Category:Fossils of Kazakhstan]]
[[Category:Cretaceous dinosaurs of Asia]]

Lesser white-toothed shrew

2017-02-11T03:19:31Z

Plantdrew: taxobox cleanup

{{Taxobox
| name = Lesser white-toothed shrew<ref name=msw3>{{MSW3 Hutterer | pages = 250}}</ref>
| image = Gartenspitzmaus.jpg
| image_caption =
| status = LC
| status_system = IUCN3.1
| status_ref = <ref name=iucn>{{IUCN2006 | assessor = Insectivore Specialist Group | year = 1996 | id = 29656 | title = Crocidura suaveolens | downloaded = 2006-05-12}} Database entry includes a brief justification of why this species is of least concern</ref>
| regnum = [[Animalia]]
| phylum = [[Chordata]]
| classis = [[Mammalia]]
| ordo = [[Eulipotyphla]]
| familia = [[Soricidae]]
| genus = ''[[Crocidura]]''
| species = '''''C. suaveolens'''''
| binomial = ''Crocidura suaveolens''
| binomial_authority = ([[Peter Simon Pallas|Pallas]], 1811)
| range_map = Lesser White-toothed Shrew area.png
| range_map_caption = Lesser white-toothed shrew range
}}

The '''lesser white-toothed shrew''' (''Crocidura suaveolens'') is a tiny [[shrew]] with a widespread distribution in [[Africa]], [[Asia]] and [[Europe]].<ref name="arkive">{{cite web|url=http://www.arkive.org/lesser-white-toothed-shrew/crocidura-suaveolens/#text=All|title=Lesser white-toothed shrew|publisher=ARKive|accessdate=21 December 2010}}</ref><ref name=Harris>{{cite web|last=Harris|first=S|title=Mammals of the British Isles|publisher=The Mammal Society|author2=Yalden, D. W }}</ref> Its preferred habitat is [[scrubland|scrub]] and gardens and it feeds on [[insect]]s, [[worm]]s, [[slug]]s, [[snail]]s, [[newt]]s and small [[rodent]]s. The closely related Asian lesser white-toothed shrew (''[[Crocidura shantungensis]]'') was once included in this species, but is now considered to be a separate species.

Like the [[common shrew]], a female lesser white-toothed shrew and her young may form a "caravan" when foraging for food or seeking a place of safety; each shrew grips the tail of the shrew in front so that the group stays together.<ref name="arkive"/>

==Distribution and habitat==
Occurs widely from France and [[Spain]], in the west, across Europe and Asia to Japan and also in North Africa.<ref name="arkive" /> There is one isolated [[United Kingdom]] population in the [[Isles of Scilly]] and another two populations off the [[France|French coast]] in the [[Channel Islands]] of [[Jersey]] and [[Sark]].<ref name=yalden>{{cite book|last1=Yalden|first1=Derek|title=The History of British Mammals|date=1999|publisher=T & A D Poyser Ltd.|location=London|isbn=0-85661-110-7}}</ref> The preferred habitat is dry ground, including scrub and gardens, and within the Isles of Scilly can be found on [[shingle beach]]es and [[Dune|sand dunes]].<ref name="arkive" />

==The Scilly shrew==
The population found on the Isles of Scilly, off the southwest coast of [[England]], was once thought to be a sub-species, ''Crocidura suaveolens cassiteridum'',<ref name="arkive"/> and is known as the Scilly shrew. Skull and tooth measurements of individuals from Scilly are found to be intermediate in size of those in the [[Channel Islands]] and the darker fur of the Scilly specimens is not considered a valid reason for the naming of a sub-species.<ref name="Harris"/> It is unusual in that it can be found on the islands' beaches.<ref>{{cite book|title=Red Data Book for Cornwall and the Isles of Scilly|year=2009|publisher=Croceago Press|location=Praze-an-Beeble|isbn=978-1-901685-01-5|pages=402–417|author=Lord D|edition=2nd|authorlink=Mammals (excluding Bats)|editor=In CISFBR}}</ref> The Scillonian name for the animal is "''teak''" or "''teke''".<ref>Robinson, H.W. (1925) ''A New British Animal Discovered in Scilly''. Scillonian 4: 123-4</ref>

Archaeological remains indicate that it was present on the islands in the [[bronze Age]], so it may have been present before the Isles of Scilly became separated from the European continent, or may have migrated from the Channel Islands or Europe onboard ships.<ref name="IOSWT">{{cite web|url=http://www.ios-wildlifetrust.org.uk/wildlife/scilly_mammals/scilly_shrew.htm|title=Scilly shrew|publisher=Isles of Scilly Wildlife Trust|accessdate=21 December 2010}}</ref> Although if shrews had survived through the [[Last Glacial Maximum|last glaciation]] or the [[Younger Dryas]], it would seem that northerly distributed species such as ''[[Sorex araneus]]'' would have been more likely to survive, rather than a southerly distributed species such as ''Crocidura suaveolens''.<ref name=yalden/>

In July 1924 W N Blair found an unknown species of shrew on [[Gugh]] and sent it to the mammal expert, Mr [[Martin Hinton|Hinton]], at the [[British Museum]]. This specimen, held at the museum, is the [[Type species|type]] for the species.<ref>Blair, W.N. (1926) ''Blair's White-toothed Shrew''. Scillonian 5:164-5.</ref> Ten years earlier H N Robinson found an unknown rodent at Old Town [[St Mary's, Isles of Scilly|St Mary's]] and sent it to Mr F W Smalley "who had the largest collection of rodents in the country".
In 2010, a Scilly shrew made headlines on [[BBC Cornwall]] when it [[Stowaway|stowed away]] on the passenger ferry [[RMV Scillonian III|RMV ''Scillonian III'']]. It was only discovered as the ship was about to arrive in Penzance. The shrew was flown back to the Isles of Scilly the next day on a [[Isles of Scilly Skybus|Skybus]] plane and then released back into its natural environment.<ref name=" shrew">[http://www.bbc.co.uk/news/10337485 Cornish ferry stowaway shrew flown home, 17 June 2010] (accessed 2011-08-16)</ref>

==References==
{{reflist}}
{{Soricomorpha|C1.}}

[[Category:Crocidura]]
[[Category:Mammals of North Africa]]
[[Category:Mammals of Europe]]
[[Category:Fauna of North Africa]]
[[Category:Mammals of Kazakhstan]]
[[Category:Mammals of Uzbekistan]]
[[Category:Mammals of Kyrgyzstan]]
[[Category:Mammals of Tajikistan]]
[[Category:Mammals of Turkmenistan]]
[[Category:Mammals of Azerbaijan]]
[[Category:Mammals of Turkey]]
[[Category:Fauna of the Palestinian territories]]
[[Category:Mammals of the Middle East]]
[[Category:Fauna of Cyprus]]
[[Category:Fauna of Northern Cyprus]]
[[Category:Environment of the Isles of Scilly]]
[[Category:Animals described in 1811]]

Eurasian water shrew

2017-02-11T03:18:19Z

Plantdrew: taxobox cleanup

{{Taxobox
| name = Eurasian water shrew<ref name=msw3>{{MSW3 Soricomorpha| id = 13700434 | page = 279}}</ref>
| image = Neomys fodiens TF 090829.jpg
| image_upright = 1.2
| status = LC
| status_system = IUCN3.1
| status_ref =<ref>{{IUCN2013.1|assessor=Hutterer, R.|assessor2=Meinig, H.|assessor3=Bertolino, S.|assessor4=Kryštufek, B.|assessor5=Amori, A.|assessor6=Sheftel, B.|assessor7=Stubbe, M.|assessor8=Samiya, R.|assessor9=Ariunbold, J.|assessor10=Buuveibaatar, V.|display-assessors=etal|year=2008 |id=29658 |title=Neomys fodiens |downloaded=2013-09-14}}</ref>
| regnum = [[Animalia]]
| phylum = [[Chordata]]
| classis = [[Mammalia]]
| ordo = [[Eulipotyphla]]
| familia = [[Soricidae]]
| genus = ''[[Neomys]]''
| species = '''''N. fodiens'''''
| binomial = ''Neomys fodiens''
| binomial_authority = ([[Thomas Pennant|Pennant]], 1771)
| range_map = Eurasian Water Shrew area.png
| range_map_caption = Eurasian water shrew range
}}

The '''Eurasian water shrew''' (''Neomys fodiens''), known in the [[United Kingdom]] as the '''water shrew''', is a relatively large [[shrew]], up to {{Convert|10|cm|in|0|abbr=on}} long, with a tail up to three-quarters as long again. It has short, dark fur, often with a few white tufts, a white belly, and a few stiff hairs around the feet and tail. It lives close to [[fresh water]], hunting aquatic prey in the water and nearby. Its fur traps bubbles of air in the water which greatly aids its buoyancy, but requires it to anchor itself to remain underwater for more than the briefest of dives.

Like many shrews, the water shrew has [[venom (poison)|venomous]] [[saliva]], making it one of the few [[venomous mammals]], although it is not able to puncture the skin of large animals such as [[human]]s. Highly [[Territory (animal)|territorial]], it lives a solitary life and is found throughout the northern part of Europe and Asia, from Britain to Korea.

==Description==
The Eurasian water shrew grows to a length of about {{convert|10|cm|in|0|abbr=on}} long with a tail length of {{convert|8|cm|in|0|abbr=on}} and weight of {{convert|15|to|19|g}}. The dense short fur on the head, back and sides is greyish-black. The underparts are dirty white and are sharply demarcated from the dorsal surface. Sometimes they are tinged with rusty brown or occasionally are entirely dark grey. There is a white spot just behind the eye and often another near the small, rounded ear which is nearly hidden in the fur. The nose is black and the snout long and tapering.<ref name=Collins>{{cite book |title=Mammals |last=Konig |first=Claus |year=1973 |publisher=Collins & Co. |isbn=978-0-00-212080-7 |pages=23–26 }}</ref> The sharp, mostly white teeth are tipped with red, typical of the shrew subfamily [[Soricinae]]. The rusty colour comes from deposits of iron which serve to harden the [[Tooth enamel|enamel]] and which are concentrated in the tips of the teeth, particularly the molars which are the teeth most subject to wear.<ref>{{cite journal|author1=Strait, S. G. |author2=Smith, S. C. | title = Elemental analysis of soricine enamel: pigmentation variation and distribution in molars of ''Blarina brevicauda'' | journal = Journal of Mammalogy | volume = 87 | issue = 4 | pages = 700–705 | year = 2006 | doi = 10.1644/05-MAMM-A-265R4.1 }}</ref> The female has five pairs of nipples. The legs are short and the hind feet are powerful, with a fringe of short, stiff hairs on the outer edge, both of which features assist while it is driving its body through the water. The tail is slender and has a keel of short white hairs on the underside.<ref name=Collins/> This shrew often utters shrill cries as it scurries about.<ref name=Collins/>

Its [[karyotype]] has [[Diploid|2n]] = 52 and [[Fundamental number|FN]] = 98.<ref name = msw3/>

==Distribution and habitat==
The Eurasian water shrew is found throughout Europe with the exception of Iceland, Ireland, The Iberian Peninsula, certain Mediterranean islands and the Balkans.<ref name=Collins/> In Asia, its range extends from western Siberia and Asia Minor to North Korea and the Pacific coast of Siberia.<ref name=ADW>{{cite web |url=http://animaldiversity.ummz.umich.edu/accounts/Neomys_fodiens/ |title=''Neomys fodiens'': Eurasian water shrew |author=Fahey, Bridget |year=1999 |work=Animal Diversity Web |publisher=University of Michigan Museum of Zoology |accessdate=30 August 2013}}</ref> It rarely strays far from water and is found in and around ditches, streams, ponds, watercress beds, fish ponds, damp meadows and rough bushy ground adjoining water.<ref name=Collins/>

==Biology==
Outside the breeding season, both male and female Eurasian water shrews maintain a [[Territory (animal)|territory]] but during the breeding season, only the females do so. At this time the males wander about visiting various female territories which indicates a [[Promiscuity|promiscuous]] [[mating system]] without pair bonding.<ref name=Cantoni>{{cite journal |author=Cantoni, Debora |year=1993 |title=Social and spatial organization of free-ranging shrews, ''Sorex coronatus'' and ''Neomys fodiens'' (Insectivora, Mammalia) |journal=Animal Behaviour |volume=45 |issue=5 |pages=975–995 |doi=10.1006/anbe.1993.1116 }}</ref> On the whole they are solitary animals that seem to mutually avoid each other and there is no social hierarchy.<ref name=ADW/>

The breeding season extends from April to September and much of the courtship takes place in the water. It either uses pre-existing burrows or digs its own. The nesting chamber is lined with moss, dry grass and leaves. Litters of four to eight or more young are born after a twenty-four-day [[gestation period]]. The young are tiny and helpless at birth. Their eyes open at fifteen to eighteen days and they are fully weaned at about seven weeks. Females can produce two or three litters a year.<ref name=Collins/> The juveniles disperse after weaning, setting up their own territories.<ref name=Cantoni/> They are sexually mature at six to eight months and their life expectancy is about three years.<ref name=ADW/>

The Eurasian water shrew is active both night and day and is thoroughly at home in the water. Its short fur holds air and the skin does not get wet when it swims. When it emerges from the water it enters one of its many burrows and any moisture adhering to the fur is absorbed by the earth walls. It mostly feeds on aquatic organisms which are caught while it is swimming. It can remain underwater for twenty seconds before it has to surface to breathe. Larger prey items can be subdued by the toxic secretions from its submaxillary glands. They feed on [[crayfish]], [[Gastropoda|water snails]], small [[fish]], aquatic [[larvae]], [[insect]]s, [[spider]]s, [[amphibian]]s, especially [[newt]]s and small [[rodent]]s are also eaten. It also feeds on land on such things as insect larvae.<ref name=ADW/>

The Eurasian water shrew has a pair of glands under its jaw which produce [[venom]], and this has been shown to be potent against the [[field vole]] (''Microtus agrestis''), and lethal at a minimum dose of fifteen milligrams per kilogram body weight.<ref>{{cite journal |author=Dufton, Mark J. |year=1992 |title=Venomous mammals |journal=Pharmacology & Therapeutics |volume=53 |issue=2 |pages=199–215 |doi=10.1016/0163-7258(92)90009-O }}</ref> The venom consists of a paralytic [[peptide]] which has been patented for use in neuromuscular therapy.<ref>{{cite web |url=https://www.google.com/patents?hl=en&lr=&vid=USPAT7485622&id=ZN-zAAAAEBAJ&oi=fnd&dq=Neomys+fodiens+venom&printsec=abstract#v=onepage&q=Neomys%20fodiens%20venom&f=false |title=Patent: Paralytic peptide for use in neuromuscular therapy |date=3 February 2009 |work=Patent 7485622 |publisher=United States Patent Office |accessdate=30 August 2013}}</ref>

==Status==
The [[International Union for Conservation of Nature]] lists the Eurasian water shrew as being of "[[Least concern]]" in its [[Red List of Threatened Species]]. This is because it has a large population distributed across a wide range and its population seems fairly stable. In some areas habitat degradation is occurring and wetlands are being drained but not to such an extent as to increase the status to "[[Vulnerable species|Vulnerable]]". Other possible threats include agricultural products and sewage which may pollute waterways and reduce the availability of food. In western Spain, a separate subspecies (''N. f. niethammeri'') has a very limited range and may be declining in numbers.<ref name=iucn>{{IUCN2008|assessor=Hutterer, R.|assessor2=Meinig, H.|assessor3=Bertolino, S.|assessor4=Kryštufek, B.|assessor5=Amori, A.|assessor6=Sheftel, B.|assessor7=Stubbe, M.|assessor8=Samiya, R.|assessor9=Ariunbold, J.|assessor10=Buuveibaatar, V.|display-assessors=etal|year=2008 |id=29658 |title=''Neomys fodiens'' |downloaded=30 August 2013}}</ref>

==References==
{{reflist}}

[[Category:Neomys]]
[[Category:Mammals of Asia]]
[[Category:Mammals of Europe]]
[[Category:Mammals of Russia]]
[[Category:Mammals of Kazakhstan]]
[[Category:Venomous mammals]]

Eurasian harvest mouse

2017-02-06T21:51:53Z

Plantdrew: taxobox cleanup

{{Taxobox
| image = Micromysminutus1.jpg
| status = LC
| status_system = IUCN3.1
| status_ref = <ref name=iucn>{{cite journal |author1=Aplin, K.|author2=Lunde, D.|author3=Batsaikhan, N.|author4=Kryštufek, B.|author5=Meinig, H.|author6=Henttonen, H. | title=''Micromys minutus''| journal = The IUCN Red List of Threatened Species| volume= 2008|publisher = [[IUCN]] | year = 2008 | page = e.T13373A3875408| url = http://www.iucnredlist.org/details/13373/0|doi=10.2305/IUCN.UK.2008.RLTS.T13373A3875408.en| accessdate = 7 August 2016}}</ref>
| regnum = [[Animal]]ia
| phylum = [[Chordata]]
| classis = [[Mammalia]]
| ordo = [[Rodentia]]
| familia = [[Muridae]]
| subfamilia = [[Murinae]]
| genus = ''[[Micromys]]''
| species = '''''M. minutus'''''
| binomial = ''Micromys minutus''
| binomial_authority = ([[Peter Simon Pallas|Pallas]], 1771)
| range_map = Distribution of Micromys minutus Map.png
| range_map_caption = Distribution of harvest mice
}}

The '''harvest mouse''' (''Micromys minutus'') is a small [[rodent]] native to [[Europe]] and [[Asia]]. It is typically found in fields of [[cereal]] crops, such as [[wheat]] and [[oat]]s, in [[reed bed]]s and in other tall ground vegetation, such as long [[grass]] and [[hedgerow]]s. It has reddish-brown fur with white underparts and a naked, highly [[prehensile tail]], which it uses for climbing. It is the smallest European rodent; an adult may weigh as little as {{Convert|4|g|oz}}. It eats chiefly [[seed]]s and [[insects]], but also nectar and fruit. Breeding nests are spherical constructions carefully woven from [[grass]] and attached to [[Plant stem|stems]] well above the ground.

==History==
Before the harvest mouse had been formally described, [[Gilbert White]] believed they were an undescribed species, and reported their nests in [[Selborne|Selborne, Hampshire]]:
<blockquote>They never enter into houses; are carried into ricks and barns with the sheaves; abound in harvest; and build their nests amidst the straws of the corn above the ground, and sometimes in [[thistle]]s. They breed as many as eight at a litter, in a little round nest composed of the blades or grass or wheat.
One of these nests I procured this autumn, most artificially platted,<ref>"artificially platted": skillfully woven.</ref> and composed of the blades of wheat; perfectly round, and about the size of a [[Cricket ball|cricket-ball]]. It was so compact and well-filled, that it would roll across the table without being discomposed, though it contained eight little mice that were naked and blind.<ref>White, ''The Natural History of Selborne'', letter xii (4 November 1767).</ref></blockquote>

Conservation efforts have taken place in Britain since 2001. Tennis balls used in play at [[The Championships, Wimbledon|Wimbledon]] have been recycled to create artificial nests for harvest mice in an attempt to help the species avoid predation and recover from near-threatened status.<ref>{{cite web|url=http://news.bbc.co.uk/1/hi/uk/1406612.stm|title='New balls, please' for mice homes|date=25 June 2001|publisher=|via=bbc.co.uk}}</ref>

==Description==
[[File:Harvest Mouse (face).jpg|220px|left|thumb|Portrait]]
The harvest mouse ranges from {{Convert|55|to|75|mm|in|abbr=on}} long, and its tail from {{Convert|50|to|75|mm|in|abbr=on}} long; it weighs from {{Convert|4|to|11|g|oz|abbr=on}},<ref>{{cite web|url=http://www.bbc.co.uk/nature/wildfacts/factfiles/266.shtml |title=Archived copy |accessdate=2004-09-11 |deadurl=yes |archiveurl=https://web.archive.org/web/20041119125341/http://www.bbc.co.uk:80/nature/wildfacts/factfiles/266.shtml |archivedate=2004-11-19 |df= }}</ref><ref>[http://www.arkive.org/harvest-mouse/micromys-minutus/#text=FactsStatus Arkive: ''Micromys minutus''.]</ref> or about half the weight of the [[house mouse]] (''Mus musculus''). Its eyes and ears are relatively large. It has a small nose, with short, stubble-like whiskers, and thick, soft fur, somewhat thicker in winter than in summer.<ref name=ADW>{{cite web |url=http://animaldiversity.ummz.umich.edu/site/accounts/information/Micromys_minutus.html |author=Ivaldi, Francesca |title=Micromys minutus |accessdate=28 May 2009 }}</ref>

The upper part of the body is brown, sometimes with a yellow or red tinge; the under-parts range from white to cream coloured. It has a prehensile tail which is usually bicoloured and furless at the tip. The mouse's rather broad feet are adapted specifically for climbing, with a somewhat opposable, large outermost toe, allowing it to grip stems with each hindfoot and its tail,<ref name=ADW/> thus freeing the mouse's forepaws for food collection. Its tail is also used for balance.

==References==
{{reflist|2}}

==External links==
*[http://www.yptenc.org.uk/docs/factsheets/animal_facts/harvest_mouse.html Young Peoples Trust for the Environment Fact Sheet]
*[http://www.bbc.co.uk/nature/wildfacts/factfiles/266.shtml BBC Wildfacts]
*[http://news.bbc.co.uk/2/hi/uk_news/1406612.stm BBC News 'New Balls Please' For Mice Homes]
{{Murinae (Melasmothrix–Mus)}}

[[Category:Old World rats and mice]]
[[Category:Mammals of Asia]]
[[Category:Rodents of Europe]]
[[Category:Mammals of Russia]]
[[Category:Mammals of Turkey]]
[[Category:Mammals of Kazakhstan]]
[[Category:Mammals of Azerbaijan]]
[[Category:Mammals of the Middle East|Mouse, Harvest]]
[[Category:Mammals of Mongolia]]
[[Category:Mammals of South Korea]]
[[Category:Mammals of North Korea]]
[[Category:Mammals of Japan]]
[[Category:Mammals of China]]

Dysoneura

2017-01-31T02:48:10Z

Plantdrew: taxobox cleanup

{{italic title}}
{{Taxobox
| fossil_range =
| regnum = [[Animalia]]
| phylum = [[Arthropoda]]
| classis = [[Insecta]]
| ordo = [[Trichoptera]]
| familia = [[Dysoneuridae]]
| genus = '''''Dysoneura'''''
}}

'''''Dysoneura''''' is the type genus of the family [[Dysoneuridae]]. It contains two species, ''[[Dysoneura trifurcata]]'' (the type species of this genus) and ''[[Dysoneura zherikhini]]''.<ref name=fossilworks>{{cite web |url=http://fossilworks.org/bridge.pl?a=taxonInfo&taxon_no=134267 |title=Fossilworks: Dysoneura |work=Fossilworks; Gateway to the Paleobiology Database |date=December 29, 2014 |accessdate=December 29, 2014}}</ref> The genus lived during the [[Jurassic]] period and is found in [[Russia]] and [[Kazakhstan]].<ref name=fossilworks/>

==References==
{{Reflist}}

[[Category:Jurassic insects]]
[[Category:Trichoptera]]
[[Category:Fossils of Kazakhstan]]

Cancriniella

2017-01-28T22:37:27Z

Plantdrew: link

{{taxobox
|image =
|image_caption =
|regnum = [[Plantae]]
|unranked_divisio = [[Angiosperms]]
|unranked_classis = [[Eudicots]]
|unranked_ordo = [[Asterids]]
|ordo = [[Asterales]]
|familia = [[Asteraceae]]
|subfamilia = [[Asteroideae]]
|tribus = [[Anthemideae]]
|genus = '''''Cancriniella'''''
|genus_authority = Tzvelev
|species = '''''C. krascheninnikovii'''''
|binomial = ''Cancriniella krascheninnikovii''
|binomial_authority = (Rubtzov) Tzvelev
|synonyms_ref=<ref>[http://www.theplantlist.org/tpl1.1/record/gcc-109790 The Plant List, ''Cancriniella krascheninnikovii'' (Rubtzov) Tzvelev ]</ref>
|synonyms =* ''[[Brachanthemum krascheninnikovii]]'' Rubtzov
*''[[Cancrinia krascheninnikovii]]'' (Rubtzov) Poljakov
}}

'''''Cancriniella''''' is a [[monotypic taxon|monotypic genus]] of flowering plants in the aster family, [[Asteraceae]].<ref>[http://www.tropicos.org/Name/40000626 Tropicos, ''Cancriniella'' Tzvelev ]</ref> It contains only one known species '''''Cancriniella krascheninnikovii''''', [[endemism|endemic]] to [[Kazakhstan]].<ref>[http://compositae.landcareresearch.co.nz/default.aspx?Page=NameDetails&TabNum=0&NameId=b6fd9aa1-a40b-47e0-92d0-af7953913560 ''Cancriniella krascheninnikovii''.] Global Compositae Checklist.</ref>

==References==
{{reflist}}

[[Category:Anthemideae]]
[[Category:Monotypic Asteraceae genera]]
[[Category:Endemic flora of Kazakhstan]]

{{Asteroideae-stub}}

Kazakhstan blind mole-rat

2017-01-25T06:21:31Z

Plantdrew: taxobox cleanup

{{Taxobox
| name = Kazakhstan blind mole-rat
| image =
| status = NT
| status_system = IUCN3.1
| status_ref = <ref name="iucn"/>
| regnum = [[Animalia]]
| phylum = [[Chordata]]
| classis = [[Mammalia]]
| ordo = [[Rodentia]]
| familia = [[Spalacidae]]
| genus = ''[[Spalax]]''
| species = '''''S. uralensis'''''
| binomial = ''Spalax uralensis''
| binomial_authority = (Tiflov and Usov, 1939)
}}

'''Kazakhstan blind mole-rat''' (''Spalax uralensis'', alternatively '''Ural blind mole-rat''') is an extant species of [[Spalax|blind mole-rat]] indigenous to [[Kazakhstan]], along the [[Ural River]] basin and the flood plains of the [[Uil River|Uil]], Temir, and [[Emba River|Emba]] rivers.<ref name="msw3"/> Whereas the species inhabits an area approximately 100,000 km2 in size, the [[International Union for Conservation of Nature]] (IUCN) describes ''S. uralensis'' as rare and its habitat prone to overgrazing.<ref name="iucn"/> In 1993, Puzachenko's [[morphometrics|morphometric]] analysis demonstrated that the species was distinct from ''S. giganteus''.<ref name="msw3"/>

==References==
{{reflist|refs=
<ref name="iucn">{{IUCN|id=136581|taxon=Spalax uralensis|assessor=Tsytsulina, K.|assessment_year=2008|version=2012.1|accessdate=12 August 2012}}</ref>
<ref name="msw3">{{cite book|last1=Wilson|first1=D. E.|last2=Reeder|first2=D. M.|year=2005|title=Mammal Species of the World|publisher=[[Johns Hopkins University Press]]|location=[[Baltimore]], [[Maryland|MD]]|isbn=978-0801882210|edition=3rd}}</ref>
}}

{{Spalacidae nav}}
{{taxonbar}}

[[Category:Animals described in 1939]]
[[Category:Mammals of Kazakhstan]]
[[Category:Spalax]]
[[Category:Spalacidae]]

Utegenia

2017-01-24T22:33:42Z

Plantdrew: taxobox cleanup

{{Use dmy dates|date=August 2013}}
{{Taxobox
| image = Ariekanerpeton sigalovi.jpg
| image_caption = Fossil
| fossil_range = [[Permian]]
| regnum = [[Animal]]ia
| phylum = [[Chordata]]
| unranked_ordo = [[Reptiliomorpha]]
| ordo = [[Seymouriamorpha]]
| familia = ''[[incertae sedis]]''
| genus = '''''Utegenia'''''
| genus_authority = Kuznetsov & Ivakhnenko, 1981
| species = '''''Utegenia shpinari'''''
| species_authority = Kuznetsov & Ivakhnenko, 1981
| synonyms =
*''Urumqia'' Zhang, Li & Wang, 1984
}}

'''''Utegenia''''' is a [[genus]] of early [[tetrapod]]. It is usually regarded as a [[Basal (phylogenetics)|basal]] [[Seymouriamorpha|seymouriamorph]],<ref>Laurin,M. A reappraisal of Utegenia, a Permo-Carboniferous seymouriamorph (Tetrapoda: Batrachosauria) from Kazakhstan. Journal of Vertebrate Paleontology, 16(3):374-383 (1996)</ref> but sometimes included in the [[Discosauriscidae]] or as a [[sister taxon]] of the latter.<ref>Klembara J.; Ruta M. The seymouriamorph tetrapod Utegenia shpinari from the ?Upper Carboniferous–Lower Permian of Kazakhstan. Part II: Postcranial anatomy and relationships. Transactions of the Royal Society of Edinburgh: Earth Sciences, Vol. 94, 1, pp. 75-93(19) (2004)</ref> Only one species, '''''Utegenia shpinari''''', found from [[Kazakhstan]], is known.<ref>Laurin, Michel. 1996. Utegenia shpinari. Version 1 January 1996. http://tolweb.org/Utegenia_shpinari/17542/1996.01.01 in The Tree of Life Web Project, http://tolweb.org/</ref> ''Urumqia'', another basal seymouriamorph, from [[Urumqi]], [[Xinjiang]] of [[China]]<ref name=Zhang1984>{{cite journal |author=Zhang Fakui, Li Yaozeng and Wang Xungang |year=1984 |title=A new occurrence of Permian seymouriamorphs in Xinjiang, China |url=http://www.ivpp.cas.cn/cbw/gjzdwxb/xbwzxz/200903/P020100316290165456321.pdf |journal=Vertebrata PalAsiatica |volume=22 |issue=4 |pages= 294–304}}</ref> is probably a [[junior synonym]] of ''Utegenia''.<ref name=Ivakhnenko87>{{cite journal |author=Ivakhnenko, M. F. |year=1987 |title=Permian parareptiles of USSR |journal=Trudy Paleontologicheskovo Instituta AN SSSR |volume=223 |pages= 1–160}}</ref>

==References==
{{reflist}}

[[Category:Seymouriamorphs]]
{{portal|Paleontology}}
[[Category:Permian tetrapods]]
[[Category:Fossils of Kazakhstan]]
{{paleo-amphibian-stub}}

Altai argali

2016-12-04T21:35:25Z

Plantdrew: taxobox cleanup

{{subspeciesbox
| name = Altai argali
| status = VU
| image=
| status_system = IUCN2.3
| status_ref = <ref>[http://www.iucnredlist.org/search/details.php/15741/all Entry in the list of endangered species]</ref>
| genus = Ovis
| species = ammon
| subspecies = ammon
| authority = ([[Carl Linnaeus|Linnaeus]], [[10th edition of Systema Naturae|1758]])
}}

'''Altai argali''' ('''''Ovis ammon ammon''''') is a traditional [[subspecies]] of [[argali]], a wild [[sheep]] that roams the highlands of the [[Altai Mountains]] in [[Central Asia]].

Altai argali are the largest of all sheep.

They have the heaviest horns. Those of mature males normally 45-50 and up to 75 pounds (35 kg).<ref>[[Rashid Jamsheed]], "Memories of a Sheep Hunter", Safari Press (1996, 2nd edition) ISBN 1-57157-021-7</ref>

==References==
{{reflist}}

{{Argali}}

[[Category:Ovis]]
[[Category:Fauna of Central Asia]]
[[Category:Megafauna of Eurasia]]
[[Category:Mammals of China]]
[[Category:Mammals of Russia]]
[[Category:Mammals of Mongolia]]
[[Category:Fauna of Kazakhstan]]
[[Category:Animals described in 1758]]

{{sheep-stub}}

Toragnostus

2016-11-20T06:17:54Z

Plantdrew: taxobox cleanup

{{italic title}}
{{Taxobox
| fossil_range = {{Geological range|506|499|late Middle Cambrian}}
| image = Toragnostus bituberculatus draw.png
| image_upright = 0.5
| image_caption = ''Toragnostus bituberculatus'' from specimen from Andrarum, Skåne, Sweden
| regnum = [[Animal]]ia
| phylum = [[Arthropod]]a
| classis = [[Trilobita]]
| ordo = [[Agnostida]]
| subordo = [[Agnostina]]
| superfamilia = [[Agnostoidea]]
| familia = [[incertae sedis]]
| genus = '''''Toragnostus'''''
| genus_authority = Robison, 1988
| subdivision_ranks = Species
| subdivision =
* ''T. bituberculatus '' (Angelin, 1851) synonyms ''Agnostus bituberculatus'', ''Phalagnostus bituberculatus'', ''Phoidagnostus bituberculatus''<ref name='Palmer'>{{cite book|last= [[Allison R. Palmer|Palmer]]|first= A.R.|year= 1967|title= Trilobites of East-Cental Alaska|journal= Geological Survey Professional Paper|issue= 559-B|url= https://books.google.com/books?id=rE7wAAAAMAAJ&pg=SL2-PA32 |accessdate= 22 May 2013}}</ref>
* ''T. placidus'' Buchholz, 1997 (tentatively assigned)
}}

'''''Toragnostus''''' is a [[genus]] of [[trilobites]] restricted to the late Middle [[Cambrian]]. Its remains have been found in the United States, Greenland, Denmark, China, Sweden, the Russian Federation, and Kazakhstan. Its headshield and tailshield are almost completely effaced and it has two thorax segments.

== Distribution ==
* ''T. bituberculatus'' occurs in the late Middle Cambrian of the United States (Hillard Peak area, Alaska)<ref name='Palmer'/> Greenland (''Lejopyge laevigata''-zone, Holm Dal Formation, West Peary-land), Denmark, China (Hunan), Sweden (''Jincella brachymetopa''-zone, [[:sv:Andrarum|Andrarum]], Skåne), the Russian Federation (''C. oriens'' and ''A. henrici''-zones, Siberian Platform, and [[Bennett Island]]), and Kazakhstan ([[Mayan stage|Mayan]], [[Tian Shan]] Range).<ref name='Treatise'>[[Harry B. Whittington|Whittington]], H. B. et al. Part O, Treatise on Invertebrate Paleontology. Revised, Volume 1 – Trilobita – Introduction, Order Agnostida, Order Redlichiida. 1997</ref>

== Description ==
Like all Agnostida, ''Toragnostus'' is diminutive, with the headshield (or [[Cephalon (arthropod head)#Trilobites|cephalon]]) and tailshield (or [[pygidium]]) of approximately the same size (or [[Pygidium#Trilobites|isopygous]]) and outline. Like all Agnostina, ''Toragnostus'' has only two thorax segments. The cephalon and pygidium are almost completely effaced, lacking a cephalic border and with wide pygidial border. The cephalon is strongly convex. The central raised area of the cephalon (or glabella) is outlined at the rear, narrow and tapering forward. It carries an elongated median node, clearly in the front half of the cephalon. The pygidial axis (or rhachis) is very broad, with the frontal lobe (M1) defined by a furrow crossing the axis. There is a weak elongated node on the rhachis.<ref name='Robison'>{{cite journal|last= Robison|first= R.A.|year= 1988|title= Stratigraphy and Palaentology of the Holm Dal Formation (late Middle Cambrian), Central North Greenland|journal= Meddelser om Gronland, Geoscience|issue= 20|url= https://books.google.com/books?id=9GihNpPF8fgC&pg=PA53 | accessdate=22 May 2013}}</ref>

== References ==
{{Reflist}}

[[Category:Agnostida]]
[[Category:Cambrian trilobites]]
[[Category:Fossils of the United States]]
[[Category:Fossils of Greenland]]
[[Category:Fossils of Denmark]]
[[Category:Fossils of China]]
[[Category:Fossils of Sweden]]
[[Category:Fossils of Russia]]
[[Category:Fossils of Kazakhstan]]

Typha pallida

2015-12-13T10:19:51Z

Plantdrew: remove Chinese name

{{italic title}}
{{taxobox
|regnum = [[Plantae]]
|unranked_divisio = [[Angiosperms]]
|unranked_classis = [[Monocots]]
|unranked_ordo = [[Commelinids]]
|ordo = [[Poales]]
|familia = [[Typhaceae]]
|genus = ''[[Typha]]''
|species = '''''T. pallida '''''
|binomial = ''Typha pallida ''
|binomial_authority =[[Evgeniia Georgievna Pobedimova|Pobed.]]
|synonyms =
|synonyms_ref=
}}
'''''Typha pallida ''''' is a plant species native to [[Kazakhstan]], [[Tajikistan]], [[Uzbekistan]] and [[China]] ([[Hebei]], [[Nei Mongol]], [[Xinjiang]]). It grows in freshwater marshes and along the banks of streams and lakes. The species is closely related to ''[[Typha minima|T. minima]].''<ref>Evgeniia Georgievna Pobedimova. 1949. Botanicheskie Materialy Gerbariya Botanicheskogo Instituta Imeni V. L. Komarova Akademii Nauk S S S R. Leningrad 11: 17, ''Typha pallida ''</ref><ref>[http://www.efloras.org/florataxon.aspx?flora_id=2&taxon_id=242353304 Flora of China v 23 p 163, ''Typha pallida'']</ref><ref>Czerepanov, S. K. 1981. Sosudistye Rasteniia SSSR 509 pages. Nauka, Leningradskoe Otd-nie, Leningrad. ''Typha minima''</ref>

==References==
{{reflist}}

[[Category:Typhaceae]]
[[Category:Freshwater plants]]
[[Category:Flora of Kazakhstan]]
[[Category:Flora of Tajikistan]]
[[Category:Flora of Uzbekistan]]
[[Category:Flora of Hebei]]
[[Category:Flora of Xinjiang]]
[[Category:Flora of Inner Mongolia]]
[[Category:Plants described in 1949]]