491:
3684:
1468:
111:
3382:
131:
392:
3394:
3717:
3388:
2861:
2866:
514:, due to its lack of cranial similarities to others. It was also the first synapsid described from the Russian area to be considered a “good pelycosaur”, as it possessed upper jaw teeth that were consistent with other known pelycosaurs (slender, recurved, enlarged incisors, single caniniform tooth). Romer and Price hypothesized that
1420:). This evolutionary stasis may be attributed to a conserved niche occupation throughout their temporal and geographical ranges. Further research on postcranial features is required in order to determine if this hypothesis is correct, as we would need to observe a similar degree of stasis throughout the entire skeleton.
490:
435:
First premaxillary tooth smaller than the second and third teeth. (There is a maximum of five teeth on the premaxilla. The first tooth is smaller than the second and third, but larger than the last two. These five teeth are closely spaced, strongly recurved, sharply pointed, and have a sharp, cutting
1447:
They possessed unbent and flattened unguals, suggesting they were diggers. It is uncertain whether their digging unguals were adapted for burrowing or solely digging for food. Many varanopids were arboreal, however the well-developed olecranon(bony prominence of the elbow) of
Mesenosaurus indicates
1415:
There is a 20 Myr gap between these two species, which exceeds the temporal range of most extinct tetrapods. Though this gap is questionable, the radioisotopic dating of speleothems, recovered for
Richards Spur, used to identify this 20 Myr gap is a reliable technique that has been used to identify
501:
The Mezen River basin is located in northern Russia and has extensive exposures of Middle
Permian sediments along the edges of affluent rivers of the Mezen River. These sediments have produced many skeletal remains of diverse amniotes, but most importantly, a partial skull of “a small synapsid of
1757:
Spindler, Frederik; Werneburg, Ralf; Schneider, Joerg W.; Luthardt, Ludwig; Annacker, Volker; Rößler, Ronny (2018-03-15). "First arboreal 'pelycosaurs' (Synapsida: Varanopidae) from the early
Permian Chemnitz Fossil Lagerstätte, SE Germany, with a review of varanopid phylogeny".
1448:
the presence of triceps and anconeus muscle, both of which would provide powerful forearm extension. This forearm extension strength combined with its somewhat small/medium body size supports the idea of a burrowing lifestyle. It is also proposed that
1431:
was a small mobile creature, capable of climbing rocks/trees. Their small size allowed them to occupy and thrive in niches as small faunivores while therapsids dominated most terrestrial environments. It was not until the late
Permian when small
1624:
MacDougall, Mark J.; Tabor, Neil J.; Woodhead, Jon; Daoust, Andrew R.; Reisz, Robert R. (June 2017). "The unique preservational environment of the Early
Permian (Cisuralian) fossiliferous cave deposits of the Richards Spur locality, Oklahoma".
1703:
Woodhead, Jon; Reisz, Robert; Fox, David; Drysdale, Russell; Hellstrom, John; Maas, Roland; Cheng, Hai; Edwards, R. Lawrence (May 2010). "Speleothem climate records from deep time? Exploring the potential with an example from the
Permian".
588:
from Russia. Some of these shared features include relative size and shape of the temporal fenestra, lateral swelling of the maxilla in the caniniform region and five premaxillary tooth positions (not reported in other mycterosaurines).
561:. Its nearly complete skull and mandible was discovered at Richards Spur locality within a series of infilled karst fissures in the Ordovician Arbuckle limestone in Oklahoma, which is one of the most plentiful sites for early
1416:
other large gaps/long temporal ranges. This gap is significant, as it indicates evolutionary stasis (persisting throughout different environments across Pangea as well as faunal turnovers throughout the
Permian-specifically
600:
share many distinct features, there are four main morphological differences between these specimens that deem a taxonomic distinction at the species level (differences insufficient for distinction above species level):
1908:
Spindler, Frederik; Werneburg, Ralf; Schneider, Jörg W. (2019-01-19). "A new mesenosaurine from the lower
Permian of Germany and the postcrania of Mesenosaurus: implications for early amniote comparative osteology".
383:
are small sized varanopid synapsids. They are characterized by mainly cranial features. Many of the postcranial features of this genus have not been analyzed fully due to a lack of fossil evidence.
466:
A very prominent, nodular ornamentation is present on the orbital margins of the prefrontal, postorbital, and jugal. (This characteristic is strikingly pronounced in well-preserved skulls of
580:), such as the “exclusion of the lacrimal from the external naris and an anteroposteriorly broad dorsal lamina of the maxilla that underlies the nasal and contacts the prefrontal”. However,
1801:
Anderson, Jason S.; Reisz, Robert R. (2004-03-25). "Pyozia mesenensis, a new, small varanopid (Synapsida, Eupelycosauria) from Russia: "pelycosaur" diversity in the Middle
Permian".
1456:. This is based on the presence of a rearward shift of center of body mass (slender trunks, elongated hindlimbs, and short forelimbs) that is necessary for facultative bipedalism.
2097:
1852:
Modesto, Christian A. Sidor, Bruce, Sean (2001-12-01). "A second varanopseid skull from the Upper Permian of South Africa: implications for Late Permian 'pelycosaur' evolution".
1444:
represents a guild of highly agile subordinate predators in their communities due to their large, slender, curved teeth, which could cause severe wounds when piercing its prey.
605:
The presence of short dorsal premaxillary processes (that do not extend to either the posterior narial margin or the posterior separation of the premaxillae by the nasals)
525:
Marginal dentition is composed of strongly recurved, laterally compressed, sharply pointed teeth front and back cutting edges restricted to the distal half of the tooth
1887:
3795:
426:
Well-developed depression on the lateral surface of the nasal that extends posteriorly from the narial border to nearly the anterior end of the prefrontal
3782:
543:
Parasphenoid plate is broad and basisphenoidal tubera are wing like and extend far laterally and posteriorly from the base of cultriform process
3884:
3879:
1668:
Benson, R.J. (2012). "Interrelationships of basal synapsids: cranial and postcranial morphological partitions suggest different topologies".
3889:
3854:
2073:
2126:
2089:
446:
3683:
3874:
522:
Dorsoventral expanded temporal fenestra occupies most of the height of temporal region, resulting in narrow subtemporal bar
537:
Tabular shrinks in size that become small, narrow elements that contacts medial margin of posterolateral wing of parietal
1581:
romeri, a small varanopseid (Synapsida: Eupelycosauria) from the Upper Permian of the Mezen River Basin, northern Russia
1011:
130:
3393:
3381:
3869:
3859:
957:
915:
2205:
1606:, Geological Society of America Special Papers, vol. 28, Geological Society of America, pp. 1–534,
540:
Absence of medial, occipital flange of posterior margin of squamosal that covers posterior margin of quadrate
3864:
2232:
2119:
1264:
1094:
1069:
891:
528:
Well-developed premaxillary subnarial shelf whose external surface is broadly rounded in transverse section
3737:
3585:
3387:
2860:
1449:
1305:
474:. However, careful examination and analysis have shown that this feature is present in other varanopids.)
3821:
3492:
3466:
1881:
1181:
728:
518:
belonged to Varanopseidae. This hypothesis was confirmed in 2001 based on the following synapomorphies:
1521:"A new varanopid synapsid from the early Permian of Oklahoma and the evolutionary stasis in this clade"
1417:
3769:
3550:
3518:
1810:
1713:
1634:
1112:
362:
as a genus. The genus name means “lizard from Mezen” while the specific epithet is given in honor of
2865:
546:
Prominent, nodular ornamentation is present on orbital margins of prefrontal, postorbital, and jugal
3849:
3608:
1481:
1473:
793:
676:
3720:
3267:
3126:
3079:
2112:
2051:
1934:
1834:
1783:
1685:
1288:
125:
442:
Postorbital cheek region of skull unusually broad and low, with nearly vertical posterior margin
3826:
2081:
417:
Dorsal process of premaxilla is long and forms anterior half of dorsal margin of external naris
110:
3808:
3800:
3207:
3180:
2043:
2008:
1971:
1926:
1869:
1826:
1775:
1739:
1650:
1584:
1550:
1035:
3813:
3237:
2976:
2836:
2479:
2035:
1998:
1961:
1918:
1861:
1818:
1767:
1729:
1721:
1677:
1642:
1607:
1540:
1532:
1222:
572:, it shares distinct cranial features with mycterosaurines (stem based group that includes
391:
3303:
3146:
3086:
2946:
2902:
2190:
1205:
877:
201:
1814:
1717:
1638:
3631:
3562:
3260:
3189:
3109:
3072:
3030:
3021:
2773:
2677:
2622:
2611:
2568:
2534:
2444:
2319:
2217:
2003:
1990:
1966:
1953:
1545:
1520:
831:
816:
659:
531:
Anterior median process of parietal extends into the supraorbital region of skull table
455:
slender, short, and rod-like, with modestly developed footplate and distally expanded
432:
Short posterior process of the maxilla fails to reach the level of the postorbital bar
3843:
3705:
3574:
3333:
3325:
3246:
3196:
3166:
3156:
3093:
2953:
2911:
2708:
2693:
2666:
2645:
2507:
2500:
2416:
1787:
456:
355:
335:
327:
68:
1938:
1838:
1689:
3710:
3700:
3664:
3524:
3481:
3455:
3317:
3253:
3058:
3044:
3037:
2960:
2932:
2575:
2514:
2458:
2451:
2409:
2402:
2395:
2344:
363:
296:
227:
2026:
Savage, Jay M.; Romer, Alfred Sherwood (1957-07-15). "Osteology of the Reptiles".
1681:
1646:
3539:
3507:
3310:
3289:
3065:
2925:
2918:
2883:
2826:
2816:
2744:
2652:
2547:
2375:
2353:
2334:
1486:
1160:
997:
770:
710:
307:
292:
188:
43:
3760:
1865:
3654:
3644:
3597:
3424:
3411:
3345:
3275:
3223:
3007:
2939:
2786:
2701:
2686:
2596:
2589:
2561:
2486:
2472:
1922:
1771:
1463:
939:
411:
323:
88:
53:
2047:
1930:
1873:
1830:
1779:
1743:
1654:
1588:
3441:
3296:
2997:
2722:
2582:
2493:
2437:
2430:
2368:
2298:
2285:
2244:
2135:
750:
634:
470:, so much so that Efremov originally considered it to be an autapomorphy of
175:
142:
93:
37:
2012:
1975:
1554:
534:
Small postfrontal medially borders narrow, posterior process of the frontal
429:
Slight lateral swelling of the maxilla at the level of the caniniform tooth
1734:
3754:
3282:
3136:
3051:
2806:
2796:
2729:
2715:
2638:
2631:
2386:
2361:
2165:
2159:
331:
162:
83:
78:
63:
58:
48:
1536:
3787:
2465:
2174:
2055:
1433:
689:
562:
510:
was discovered in 1938 by Ivan Efremov and became the first species of
319:
303:
288:
98:
73:
30:
3614:
2604:
2153:
1725:
1611:
452:
311:
152:
3731:
2039:
1822:
1954:"The flexor muscles of the forearm and hand in lizards and mammals"
2423:
489:
284:
3774:
1602:
ROMER, A. S.; PRICE, L. W. (1940), "Review of the Pelycosauria",
463:
more closely resembles those in early therapsids of the Permian.)
414:
forms a narrow, rectangular snout (from dorsal and ventral views)
1991:"A revision of the extensor muscles in the forearm in tetrapods"
3735:
3409:
2881:
2283:
2146:
2108:
342:’s stratigraphic range is the middle to late Guadalupian while
637:
modified from the analysis of Benson, after the exclusion of
423:
Palatal process of premaxilla is long with long median suture
2104:
1436:
appeared and provided competition, leading to a decline in
449:
is angled slightly anterolaterally from basal articulation.
372:
was named in honor of Ivan Efremov, who erected the genus.
420:
Expanded narial shelf that extends almost to the snout tip
576:
and all varanopseids related more closely to it than to
614:
Contact between the postorbital and supratemporal bones
3660:
3650:
3640:
3571:
3536:
3504:
3478:
3452:
3437:
3234:
3219:
3177:
3162:
3152:
3142:
3132:
3122:
3018:
3003:
2993:
2899:
2832:
2822:
2812:
2802:
2792:
2782:
2544:
2331:
2315:
234:
3744:
3629:
3596:
3570:
3561:
3535:
3503:
3477:
3451:
3422:
3344:
3233:
3206:
3176:
3107:
3017:
2974:
2898:
2771:
2743:
2676:
2621:
2543:
2532:
2385:
2343:
2330:
2296:
2243:
2216:
2189:
621:is also larger than the largest known specimen of
436:edge along the distal half of the posterior edge.)
1627:Palaeogeography, Palaeoclimatology, Palaeoecology
1519:Sigi Maho; Bryan M. Gee; Robert R. Reisz (2019).
2120:
8:
1886:: CS1 maint: multiple names: authors list (
445:Posterior edge of transverse flange of the
3732:
3567:
3430:
3419:
3406:
3212:
3115:
2986:
2982:
2895:
2891:
2878:
2768:
2553:
2540:
2340:
2308:
2304:
2293:
2280:
2222:
2195:
2186:
2143:
2127:
2113:
2105:
346:’s stratigraphic range is the Cisuralian.
109:
20:
2002:
1965:
1733:
1544:
2090:"Main Groups of Non-Mammalian Synapsida"
390:
1498:
402:The cranial features that characterize
295:. This genus includes two species: the
2100:from the original on December 3, 2023.
1879:
1903:
1901:
1899:
1897:
7:
1604:28 : Review of the Pelycosauria
1570:
1568:
1566:
1564:
1514:
1512:
1510:
1508:
1506:
1504:
1502:
611:Fewer tooth positions on the maxilla
557:The second species of this genus is
1670:Journal of Systematic Palaeontology
1280:
1256:
1197:
1173:
1166:
1085:
1061:
1051:
1027:
1003:
931:
907:
883:
873:
809:
785:
763:
756:
702:
695:
651:
644:
1803:Journal of Vertebrate Paleontology
608:More posteriorly extensive maxilla
14:
2074:"Mesenosaurus (cast of skeleton)"
439:Single, median vomerine tooth row
3716:
3715:
3682:
3392:
3386:
3380:
2864:
2859:
1466:
129:
354:Famous Russian paleontologist,
302:from the middle Permian (upper
1:
3885:Fossil taxa described in 1938
3880:Fossils of the United States
1682:10.1080/14772019.2011.631042
1647:10.1016/j.palaeo.2017.02.019
1012:Archaeovenator hamiltonensis
3890:Taxa named by Ivan Yefremov
3855:Prehistoric synapsid genera
291:. It belongs to the family
3906:
1960:. 84(Pt 1) (Pt 1): 13–29.
1866:10.1080/002411601753292971
1525:Royal Society Open Science
958:Stereophallodon ciscoensis
584:shares more features with
16:Extinct genus of synapsids
3695:
3680:
3433:
3418:
3405:
3378:
3215:
3118:
2989:
2985:
2894:
2890:
2877:
2857:
2767:
2556:
2311:
2307:
2292:
2279:
2225:
2198:
2185:
2142:
2094:Mikko's Phylogeny Archive
1997:. 73(2) (Pt 1): 211–233.
1923:10.1007/s12542-018-0439-z
1772:10.1007/s12542-018-0405-9
1575:Reisz, Robert R. (2001).
1531:(10): Article ID 191297.
1302:
1285:
1278:
1261:
1254:
1219:
1202:
1195:
1178:
1171:
1164:
1126:
1109:
1090:
1083:
1066:
1059:
1049:
1032:
1025:
1008:
1001:
954:
936:
929:
916:Varanosaurus acutirostris
912:
905:
888:
881:
871:
829:
814:
807:
790:
783:
768:
761:
754:
725:
707:
700:
693:
673:
656:
649:
502:varanopseid affinities”,
254:
249:
233:
226:
126:Scientific classification
124:
117:
108:
23:
568:In terms of classifying
459:process. (The stapes of
269:Maho, Gee et Reisz, 2019
1265:Ruthiromia elcobriensis
1095:Elliotsmithia longiceps
1070:Mycterosaurus longiceps
892:Archaeothyris florensis
574:Mycterosaurus longiceps
1450:facultative bipedalism
1306:Aerosaurus greenleorum
498:
399:
3875:Permian United States
3822:Paleobiology Database
1989:Haines, R.W. (1939).
1952:Haines, R.W. (1950).
1182:Varanops brevirostris
729:Protorothyris archeri
559:Mesenosaurus efremovi
552:Mesenosaurus efremovi
493:
394:
370:Mesenosaurus efremovi
316:Mesenosaurus efremovi
1113:Heleosaurus scholtzi
1815:2004JVPal..24..173A
1718:2010Geo....38..455W
1639:2017PPP...475....1M
1537:10.1098/rsos.191297
1482:List of pelycosaurs
1474:Paleontology portal
1130:Mesenosaurus romeri
794:Ianthodon schultzei
677:Limnoscelis paludis
504:Mesenosaurus romeri
495:Mesenosaurus romeri
485:Mesenosaurus romeri
396:Mesenosaurus romeri
300:Mesenosaurus romeri
239:Mesenosaurus romeri
119:Mesenosaurus romeri
1995:Journal of Anatomy
1958:Journal of Anatomy
1487:List of varanopids
1289:Aerosaurus wellesi
565:tetrapod fossils.
499:
400:
398:skull illustration
310:Basin of northern
3870:Fossils of Russia
3860:Permian synapsids
3837:
3836:
3809:Open Tree of Life
3738:Taxon identifiers
3729:
3728:
3691:
3690:
3678:
3677:
3674:
3673:
3625:
3624:
3401:
3400:
3376:
3375:
3372:
3371:
3368:
3367:
3364:
3363:
3208:Sphenacodontoidea
3181:Palaeohatteriidae
3103:
3102:
2970:
2969:
2873:
2872:
2855:
2854:
2851:
2850:
2847:
2846:
2763:
2762:
2739:
2738:
2528:
2527:
2524:
2523:
2275:
2274:
2271:
2270:
2267:
2266:
2263:
2262:
2084:on March 4, 2016.
1407:
1406:
1398:
1397:
1389:
1388:
1380:
1379:
1371:
1370:
1362:
1361:
1353:
1352:
1344:
1343:
1335:
1334:
1326:
1325:
1317:
1316:
1243:
1242:
1234:
1233:
1150:
1149:
1141:
1140:
1036:Pyozia mesenensis
987:
986:
978:
977:
969:
968:
860:
859:
851:
850:
842:
841:
740:
739:
276:
275:
270:
262:
222:
3897:
3830:
3829:
3817:
3816:
3804:
3803:
3791:
3790:
3778:
3777:
3765:
3764:
3763:
3733:
3719:
3718:
3686:
3662:
3652:
3642:
3573:
3568:
3538:
3506:
3480:
3454:
3439:
3431:
3420:
3407:
3396:
3390:
3384:
3238:Sphenacodontidae
3236:
3221:
3213:
3179:
3164:
3154:
3144:
3134:
3124:
3116:
3020:
3005:
2995:
2987:
2983:
2977:Haptodontiformes
2901:
2896:
2892:
2879:
2868:
2863:
2837:Protoclepsydrops
2834:
2824:
2814:
2804:
2794:
2784:
2769:
2554:
2546:
2541:
2480:Lalieudorhynchus
2341:
2333:
2317:
2309:
2305:
2294:
2281:
2223:
2196:
2187:
2180:
2179:
2144:
2129:
2122:
2115:
2106:
2101:
2085:
2080:. Archived from
2078:Fossils.valdosta
2060:
2059:
2023:
2017:
2016:
2006:
1986:
1980:
1979:
1969:
1949:
1943:
1942:
1905:
1892:
1891:
1885:
1877:
1849:
1843:
1842:
1798:
1792:
1791:
1754:
1748:
1747:
1737:
1726:10.1130/g30354.1
1700:
1694:
1693:
1665:
1659:
1658:
1621:
1615:
1614:
1612:10.1130/spe28-p1
1599:
1593:
1592:
1572:
1559:
1558:
1548:
1516:
1476:
1471:
1470:
1469:
1411:Paleoenvironment
1281:
1257:
1223:Varanodon agilis
1198:
1174:
1167:
1086:
1062:
1052:
1028:
1004:
932:
908:
884:
874:
810:
786:
764:
757:
703:
696:
652:
645:
578:Varanodon agilis
268:
260:
236:
220:
213:
200:
187:
134:
133:
113:
103:
40:
29:Temporal range:
21:
3905:
3904:
3900:
3899:
3898:
3896:
3895:
3894:
3840:
3839:
3838:
3833:
3825:
3820:
3812:
3807:
3799:
3794:
3786:
3781:
3773:
3768:
3759:
3758:
3753:
3740:
3730:
3725:
3687:
3670:
3621:
3592:
3557:
3531:
3499:
3473:
3447:
3427:
3414:
3397:
3360:
3340:
3304:Secodontosaurus
3229:
3202:
3172:
3147:Hypselohaptodus
3112:
3099:
3087:Remigiomontanus
3013:
2979:
2966:
2947:Stereophallodon
2903:Ophiacodontidae
2886:
2869:
2843:
2759:
2735:
2672:
2617:
2537:
2520:
2381:
2326:
2301:
2288:
2259:
2239:
2212:
2206:Tetrapodomorpha
2191:Tetrapodomorpha
2181:
2149:
2148:
2138:
2133:
2088:
2072:
2069:
2064:
2063:
2040:10.2307/1439431
2025:
2024:
2020:
1988:
1987:
1983:
1951:
1950:
1946:
1907:
1906:
1895:
1878:
1851:
1850:
1846:
1823:10.1671/1940-13
1800:
1799:
1795:
1756:
1755:
1751:
1702:
1701:
1697:
1667:
1666:
1662:
1623:
1622:
1618:
1601:
1600:
1596:
1574:
1573:
1562:
1518:
1517:
1500:
1495:
1472:
1467:
1465:
1462:
1426:
1413:
1408:
1399:
1390:
1381:
1372:
1363:
1354:
1345:
1336:
1327:
1318:
1244:
1235:
1206:Watongia meieri
1151:
1142:
1056:Mycterosaurinae
988:
979:
970:
878:Ophiacodontidae
861:
852:
843:
741:
631:
555:
488:
481:
389:
378:
352:
318:from the early
245:
242:
219:
211:
202:Mycterosaurinae
198:
185:
128:
104:
102:
101:
96:
91:
86:
81:
76:
71:
66:
61:
56:
51:
46:
35:
34:
27:
17:
12:
11:
5:
3903:
3901:
3893:
3892:
3887:
3882:
3877:
3872:
3867:
3865:Permian Russia
3862:
3857:
3852:
3842:
3841:
3835:
3834:
3832:
3831:
3818:
3805:
3792:
3779:
3766:
3750:
3748:
3742:
3741:
3736:
3727:
3726:
3724:
3723:
3713:
3708:
3703:
3696:
3693:
3692:
3689:
3688:
3681:
3679:
3676:
3675:
3672:
3671:
3669:
3668:
3658:
3648:
3637:
3635:
3632:Incertae sedis
3627:
3626:
3623:
3622:
3620:
3619:
3618:
3617:
3611:
3602:
3600:
3594:
3593:
3591:
3590:
3589:
3588:
3579:
3577:
3565:
3563:Eutheriodontia
3559:
3558:
3556:
3555:
3554:
3553:
3544:
3542:
3533:
3532:
3530:
3529:
3528:
3527:
3521:
3512:
3510:
3501:
3500:
3498:
3497:
3496:
3495:
3486:
3484:
3475:
3474:
3472:
3471:
3470:
3469:
3460:
3458:
3449:
3448:
3446:
3445:
3434:
3428:
3423:
3416:
3415:
3410:
3403:
3402:
3399:
3398:
3379:
3377:
3374:
3373:
3370:
3369:
3366:
3365:
3362:
3361:
3359:
3358:
3357:
3356:
3350:
3348:
3342:
3341:
3339:
3338:
3330:
3322:
3314:
3307:
3300:
3293:
3286:
3279:
3272:
3264:
3261:Ctenospondylus
3257:
3250:
3242:
3240:
3231:
3230:
3228:
3227:
3216:
3210:
3204:
3203:
3201:
3200:
3193:
3190:Palaeohatteria
3185:
3183:
3174:
3173:
3171:
3170:
3160:
3150:
3140:
3130:
3119:
3113:
3110:Sphenacodontia
3108:
3105:
3104:
3101:
3100:
3098:
3097:
3090:
3083:
3076:
3073:Melanedaphodon
3069:
3062:
3055:
3048:
3041:
3034:
3031:Bohemiclavulus
3026:
3024:
3022:Edaphosauridae
3015:
3014:
3012:
3011:
3001:
2990:
2980:
2975:
2972:
2971:
2968:
2967:
2965:
2964:
2957:
2950:
2943:
2936:
2929:
2922:
2915:
2907:
2905:
2888:
2887:
2882:
2875:
2874:
2871:
2870:
2858:
2856:
2853:
2852:
2849:
2848:
2845:
2844:
2842:
2841:
2830:
2820:
2810:
2800:
2790:
2779:
2777:
2774:Incertae sedis
2765:
2764:
2761:
2760:
2758:
2757:
2756:
2755:
2749:
2747:
2741:
2740:
2737:
2736:
2734:
2733:
2726:
2719:
2712:
2705:
2698:
2690:
2682:
2680:
2678:Varanodontinae
2674:
2673:
2671:
2670:
2663:
2656:
2649:
2642:
2635:
2627:
2625:
2623:Mesenosaurinae
2619:
2618:
2616:
2615:
2612:Thrausmosaurus
2608:
2601:
2593:
2586:
2579:
2572:
2569:Archaeovenator
2565:
2557:
2551:
2538:
2535:Eupelycosauria
2533:
2530:
2529:
2526:
2525:
2522:
2521:
2519:
2518:
2511:
2504:
2497:
2490:
2483:
2476:
2469:
2462:
2455:
2448:
2445:Cotylorhynchus
2441:
2434:
2427:
2420:
2413:
2406:
2399:
2391:
2389:
2383:
2382:
2380:
2379:
2372:
2365:
2358:
2349:
2347:
2338:
2328:
2327:
2325:
2324:
2320:Diadectomorpha
2312:
2302:
2297:
2290:
2289:
2284:
2277:
2276:
2273:
2272:
2269:
2268:
2265:
2264:
2261:
2260:
2258:
2257:
2256:
2255:
2249:
2247:
2241:
2240:
2238:
2237:
2236:
2235:
2233:Reptiliomorpha
2226:
2220:
2218:Reptiliomorpha
2214:
2213:
2211:
2210:
2209:
2208:
2199:
2193:
2183:
2182:
2178:
2177:
2168:
2162:
2156:
2147:
2140:
2139:
2134:
2132:
2131:
2124:
2117:
2109:
2103:
2102:
2086:
2068:
2067:External links
2065:
2062:
2061:
2018:
1981:
1944:
1917:(2): 303–344.
1893:
1860:(4): 249–259.
1844:
1809:(1): 173–179.
1793:
1766:(2): 315–364.
1749:
1735:1959.13/931960
1712:(5): 455–458.
1695:
1676:(4): 601–624.
1660:
1616:
1594:
1560:
1497:
1496:
1494:
1491:
1490:
1489:
1484:
1478:
1477:
1461:
1458:
1425:
1422:
1412:
1409:
1405:
1404:
1401:
1400:
1396:
1395:
1392:
1391:
1387:
1386:
1383:
1382:
1378:
1377:
1374:
1373:
1369:
1368:
1365:
1364:
1360:
1359:
1356:
1355:
1351:
1350:
1347:
1346:
1342:
1341:
1338:
1337:
1333:
1332:
1329:
1328:
1324:
1323:
1320:
1319:
1315:
1314:
1311:
1310:
1301:
1298:
1297:
1294:
1293:
1284:
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1277:
1274:
1273:
1270:
1269:
1260:
1255:
1253:
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1249:
1246:
1245:
1241:
1240:
1237:
1236:
1232:
1231:
1228:
1227:
1218:
1215:
1214:
1211:
1210:
1201:
1196:
1194:
1191:
1190:
1187:
1186:
1177:
1172:
1170:
1165:
1163:
1157:
1156:
1153:
1152:
1148:
1147:
1144:
1143:
1139:
1138:
1135:
1134:
1125:
1122:
1121:
1118:
1117:
1108:
1105:
1104:
1101:
1100:
1089:
1084:
1082:
1079:
1078:
1075:
1074:
1065:
1060:
1058:
1050:
1048:
1045:
1044:
1041:
1040:
1031:
1026:
1024:
1021:
1020:
1017:
1016:
1007:
1002:
1000:
994:
993:
990:
989:
985:
984:
981:
980:
976:
975:
972:
971:
967:
966:
963:
962:
953:
950:
949:
946:
945:
935:
930:
928:
925:
924:
921:
920:
911:
906:
904:
901:
900:
897:
896:
887:
882:
880:
872:
870:
867:
866:
863:
862:
858:
857:
854:
853:
849:
848:
845:
844:
840:
839:
836:
835:
832:Sphenacodontia
828:
825:
824:
821:
820:
817:Edaphosauridae
813:
808:
806:
803:
802:
799:
798:
789:
784:
782:
779:
778:
775:
774:
767:
762:
760:
755:
753:
747:
746:
743:
742:
738:
737:
734:
733:
724:
721:
720:
717:
716:
706:
701:
699:
694:
692:
686:
685:
682:
681:
672:
669:
668:
665:
664:
660:Tseajaia campi
655:
650:
648:
643:
630:
627:
616:
615:
612:
609:
606:
554:
549:
548:
547:
544:
541:
538:
535:
532:
529:
526:
523:
487:
482:
480:
477:
476:
475:
464:
450:
443:
440:
437:
433:
430:
427:
424:
421:
418:
415:
388:
385:
377:
374:
358:, established
351:
348:
283:is an extinct
274:
273:
272:
271:
263:
252:
251:
247:
246:
243:
231:
230:
224:
223:
209:
205:
204:
196:
192:
191:
183:
179:
178:
173:
166:
165:
160:
156:
155:
150:
146:
145:
140:
136:
135:
122:
121:
115:
114:
106:
105:
97:
92:
87:
82:
77:
72:
67:
62:
57:
52:
47:
42:
41:
28:
15:
13:
10:
9:
6:
4:
3:
2:
3902:
3891:
3888:
3886:
3883:
3881:
3878:
3876:
3873:
3871:
3868:
3866:
3863:
3861:
3858:
3856:
3853:
3851:
3848:
3847:
3845:
3828:
3823:
3819:
3815:
3810:
3806:
3802:
3797:
3793:
3789:
3784:
3780:
3776:
3771:
3767:
3762:
3756:
3752:
3751:
3749:
3747:
3743:
3739:
3734:
3722:
3714:
3712:
3709:
3707:
3706:Phthinosuchia
3704:
3702:
3698:
3697:
3694:
3685:
3667:
3666:
3659:
3657:
3656:
3649:
3647:
3646:
3639:
3638:
3636:
3634:
3633:
3628:
3616:
3612:
3610:
3606:
3605:
3604:
3603:
3601:
3599:
3595:
3587:
3586:Therocephalia
3583:
3582:
3581:
3580:
3578:
3576:
3575:Therocephalia
3569:
3566:
3564:
3560:
3552:
3548:
3547:
3546:
3545:
3543:
3541:
3534:
3526:
3522:
3520:
3516:
3515:
3514:
3513:
3511:
3509:
3502:
3494:
3490:
3489:
3488:
3487:
3485:
3483:
3476:
3468:
3464:
3463:
3462:
3461:
3459:
3457:
3450:
3444:
3443:
3436:
3435:
3432:
3429:
3426:
3421:
3417:
3413:
3408:
3404:
3395:
3391:
3389:
3383:
3354:
3353:
3352:
3351:
3349:
3347:
3343:
3336:
3335:
3334:Tetraceratops
3331:
3328:
3327:
3326:Tappenosaurus
3323:
3320:
3319:
3315:
3313:
3312:
3308:
3306:
3305:
3301:
3299:
3298:
3294:
3292:
3291:
3287:
3285:
3284:
3280:
3278:
3277:
3273:
3270:
3269:
3265:
3263:
3262:
3258:
3256:
3255:
3251:
3249:
3248:
3247:Cryptovenator
3244:
3243:
3241:
3239:
3232:
3226:
3225:
3218:
3217:
3214:
3211:
3209:
3205:
3199:
3198:
3197:Pantelosaurus
3194:
3192:
3191:
3187:
3186:
3184:
3182:
3175:
3169:
3168:
3167:Tetraceratops
3161:
3159:
3158:
3157:Kenomagnathus
3151:
3149:
3148:
3141:
3139:
3138:
3131:
3129:
3128:
3121:
3120:
3117:
3114:
3111:
3106:
3096:
3095:
3094:Xyrospondylus
3091:
3089:
3088:
3084:
3082:
3081:
3077:
3075:
3074:
3070:
3068:
3067:
3063:
3061:
3060:
3056:
3054:
3053:
3049:
3047:
3046:
3042:
3040:
3039:
3035:
3033:
3032:
3028:
3027:
3025:
3023:
3016:
3010:
3009:
3002:
3000:
2999:
2992:
2991:
2988:
2984:
2981:
2978:
2973:
2963:
2962:
2958:
2956:
2955:
2954:Stereorhachis
2951:
2949:
2948:
2944:
2942:
2941:
2937:
2935:
2934:
2930:
2928:
2927:
2923:
2921:
2920:
2916:
2914:
2913:
2912:Archaeothyris
2909:
2908:
2906:
2904:
2897:
2893:
2889:
2885:
2880:
2876:
2867:
2862:
2839:
2838:
2831:
2829:
2828:
2821:
2819:
2818:
2811:
2809:
2808:
2801:
2799:
2798:
2791:
2789:
2788:
2781:
2780:
2778:
2776:
2775:
2770:
2766:
2753:
2752:
2751:
2750:
2748:
2746:
2742:
2732:
2731:
2727:
2725:
2724:
2720:
2718:
2717:
2713:
2711:
2710:
2709:Tambacarnifex
2706:
2704:
2703:
2699:
2696:
2695:
2694:Mycterosaurus
2691:
2689:
2688:
2684:
2683:
2681:
2679:
2675:
2669:
2668:
2667:Microvaranops
2664:
2662:
2661:
2657:
2655:
2654:
2650:
2648:
2647:
2646:Elliotsmithia
2643:
2641:
2640:
2636:
2634:
2633:
2629:
2628:
2626:
2624:
2620:
2614:
2613:
2609:
2607:
2606:
2602:
2599:
2598:
2594:
2592:
2591:
2587:
2585:
2584:
2580:
2578:
2577:
2573:
2571:
2570:
2566:
2564:
2563:
2559:
2558:
2555:
2552:
2549:
2542:
2539:
2536:
2531:
2517:
2516:
2512:
2510:
2509:
2508:Ruthenosaurus
2505:
2503:
2502:
2501:Phreatophasma
2498:
2496:
2495:
2491:
2489:
2488:
2484:
2482:
2481:
2477:
2475:
2474:
2470:
2468:
2467:
2463:
2461:
2460:
2456:
2454:
2453:
2449:
2447:
2446:
2442:
2440:
2439:
2435:
2433:
2432:
2428:
2426:
2425:
2421:
2419:
2418:
2417:Callibrachion
2414:
2412:
2411:
2407:
2405:
2404:
2400:
2398:
2397:
2393:
2392:
2390:
2388:
2384:
2378:
2377:
2373:
2371:
2370:
2366:
2364:
2363:
2359:
2356:
2355:
2351:
2350:
2348:
2346:
2342:
2339:
2336:
2329:
2322:
2321:
2314:
2313:
2310:
2306:
2303:
2300:
2295:
2291:
2287:
2282:
2278:
2253:
2252:
2251:
2250:
2248:
2246:
2242:
2234:
2230:
2229:
2228:
2227:
2224:
2221:
2219:
2215:
2207:
2203:
2202:
2201:
2200:
2197:
2194:
2192:
2188:
2184:
2176:
2172:
2169:
2167:
2163:
2161:
2157:
2155:
2151:
2150:
2145:
2141:
2137:
2130:
2125:
2123:
2118:
2116:
2111:
2110:
2107:
2099:
2095:
2091:
2087:
2083:
2079:
2075:
2071:
2070:
2066:
2057:
2053:
2049:
2045:
2041:
2037:
2033:
2029:
2022:
2019:
2014:
2010:
2005:
2000:
1996:
1992:
1985:
1982:
1977:
1973:
1968:
1963:
1959:
1955:
1948:
1945:
1940:
1936:
1932:
1928:
1924:
1920:
1916:
1912:
1904:
1902:
1900:
1898:
1894:
1889:
1883:
1875:
1871:
1867:
1863:
1859:
1855:
1848:
1845:
1840:
1836:
1832:
1828:
1824:
1820:
1816:
1812:
1808:
1804:
1797:
1794:
1789:
1785:
1781:
1777:
1773:
1769:
1765:
1761:
1753:
1750:
1745:
1741:
1736:
1731:
1727:
1723:
1719:
1715:
1711:
1707:
1699:
1696:
1691:
1687:
1683:
1679:
1675:
1671:
1664:
1661:
1656:
1652:
1648:
1644:
1640:
1636:
1632:
1628:
1620:
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1613:
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1598:
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1590:
1586:
1582:
1578:
1577:The skull of
1571:
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1552:
1547:
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1394:
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1349:
1348:
1340:
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1272:
1271:
1268:
1267:
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1247:
1239:
1238:
1230:
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1209:
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1200:
1199:
1193:
1192:
1189:
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1183:
1176:
1175:
1169:
1168:
1162:
1159:
1158:
1155:
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1146:
1145:
1137:
1136:
1133:
1132:
1131:
1124:
1123:
1120:
1119:
1116:
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1114:
1107:
1106:
1103:
1102:
1099:
1097:
1096:
1088:
1087:
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1080:
1077:
1076:
1073:
1072:
1071:
1064:
1063:
1057:
1054:
1053:
1047:
1046:
1043:
1042:
1039:
1038:
1037:
1030:
1029:
1023:
1022:
1019:
1018:
1015:
1014:
1013:
1006:
1005:
999:
996:
995:
992:
991:
983:
982:
974:
973:
965:
964:
961:
960:
959:
952:
951:
948:
947:
944:
942:
941:
934:
933:
927:
926:
923:
922:
919:
918:
917:
910:
909:
903:
902:
899:
898:
895:
894:
893:
886:
885:
879:
876:
875:
869:
868:
865:
864:
856:
855:
847:
846:
838:
837:
834:
833:
827:
826:
823:
822:
819:
818:
812:
811:
805:
804:
801:
800:
797:
796:
795:
788:
787:
781:
780:
777:
776:
773:
772:
766:
765:
759:
758:
752:
749:
748:
745:
744:
736:
735:
732:
731:
730:
723:
722:
719:
718:
715:
713:
712:
705:
704:
698:
697:
691:
688:
687:
684:
683:
680:
679:
678:
671:
670:
667:
666:
663:
662:
661:
654:
653:
647:
646:
642:
640:
636:
628:
626:
624:
620:
613:
610:
607:
604:
603:
602:
599:
595:
590:
587:
583:
579:
575:
571:
566:
564:
560:
553:
550:
545:
542:
539:
536:
533:
530:
527:
524:
521:
520:
519:
517:
513:
509:
505:
496:
492:
486:
483:
478:
473:
469:
465:
462:
458:
454:
451:
448:
444:
441:
438:
434:
431:
428:
425:
422:
419:
416:
413:
409:
408:
407:
405:
397:
393:
386:
384:
382:
375:
373:
371:
367:
365:
361:
357:
349:
347:
345:
341:
337:
336:United States
333:
329:
328:Richards Spur
325:
321:
317:
313:
309:
305:
301:
298:
294:
290:
286:
282:
281:
267:
264:
261:Efremov, 1938
259:
256:
255:
253:
248:
244:Efremov, 1938
241:
240:
232:
229:
225:
221:Efremov, 1938
218:
217:
210:
207:
206:
203:
197:
194:
193:
190:
184:
181:
180:
177:
174:
171:
168:
167:
164:
161:
158:
157:
154:
151:
148:
147:
144:
141:
138:
137:
132:
127:
123:
120:
116:
112:
107:
100:
95:
90:
85:
80:
75:
70:
65:
60:
55:
50:
45:
39:
36:289–265
32:
26:
22:
19:
3746:Mesenosaurus
3745:
3711:Theriodontia
3701:Pelycosauria
3665:Niaftasuchus
3663:
3653:
3643:
3630:
3493:Dinocephalia
3482:Dinocephalia
3467:Biarmosuchia
3456:Biarmosuchia
3440:
3385:
3332:
3324:
3318:Steppesaurus
3316:
3309:
3302:
3295:
3288:
3281:
3274:
3266:
3259:
3254:Ctenorhachis
3252:
3245:
3222:
3195:
3188:
3165:
3155:
3145:
3135:
3125:
3092:
3085:
3078:
3071:
3064:
3059:Ianthasaurus
3057:
3050:
3045:Glaucosaurus
3043:
3038:Edaphosaurus
3036:
3029:
3006:
2996:
2961:Varanosaurus
2959:
2952:
2945:
2938:
2933:Echinerpeton
2931:
2924:
2917:
2910:
2835:
2825:
2815:
2805:
2795:
2785:
2772:
2728:
2721:
2714:
2707:
2700:
2692:
2685:
2665:
2660:Mesenosaurus
2659:
2658:
2651:
2644:
2637:
2630:
2610:
2603:
2595:
2588:
2581:
2576:Ascendonanus
2574:
2567:
2560:
2515:Trichasaurus
2513:
2506:
2499:
2492:
2485:
2478:
2471:
2464:
2459:Ennatosaurus
2457:
2452:Datheosaurus
2450:
2443:
2436:
2429:
2422:
2415:
2410:Arisierpeton
2408:
2403:Angelosaurus
2401:
2396:Alierasaurus
2394:
2374:
2367:
2360:
2352:
2345:Eothyrididae
2318:
2170:
2164:Superclass:
2093:
2082:the original
2077:
2031:
2027:
2021:
1994:
1984:
1957:
1947:
1914:
1910:
1882:cite journal
1857:
1853:
1847:
1806:
1802:
1796:
1763:
1759:
1752:
1709:
1705:
1698:
1673:
1669:
1663:
1630:
1626:
1619:
1603:
1597:
1580:
1579:Mesenosaurus
1576:
1528:
1524:
1454:Mesenosaurus
1453:
1452:occurred in
1446:
1442:Mesenosaurus
1441:
1438:Mesenosaurus
1437:
1429:Mesenosaurus
1428:
1427:
1424:Paleobiology
1414:
1304:
1303:
1287:
1286:
1263:
1262:
1221:
1220:
1204:
1203:
1180:
1179:
1129:
1128:
1127:
1111:
1110:
1098:(BP/1/5678)
1093:
1091:
1068:
1067:
1055:
1034:
1033:
1010:
1009:
956:
955:
938:
937:
914:
913:
890:
889:
830:
815:
792:
791:
769:
727:
726:
709:
708:
675:
674:
658:
657:
639:Basicranodon
638:
632:
622:
618:
617:
597:
593:
591:
585:
581:
577:
573:
569:
567:
558:
556:
551:
515:
512:Mesenosaurus
511:
507:
503:
500:
494:
484:
472:Mesenosaurus
471:
468:Mesenosaurus
467:
461:Mesenosaurus
460:
404:Mesenosaurus
403:
401:
395:
381:Mesenosaurus
380:
379:
369:
368:
364:Alfred Romer
360:Mesenosaurus
359:
356:Ivan Efremov
353:
343:
339:
315:
299:
297:type species
280:Mesenosaurus
279:
278:
277:
265:
257:
238:
237:
228:Type species
216:Mesenosaurus
215:
214:
169:
118:
25:Mesenosaurus
24:
18:
3551:Gorgonopsia
3540:Gorgonopsia
3525:dicynodonts
3519:Anomodontia
3508:Anomodontia
3311:Sphenacodon
3290:Macromerion
3080:Ramodendron
3066:Lupeosaurus
2926:Clepsydrops
2919:Baldwinonus
2884:Metopophora
2827:Mastersonia
2817:Knoxosaurus
2745:Metopophora
2653:Heleosaurus
2548:Varanopidae
2376:Vaughnictis
2354:Asaphestera
2335:Caseasauria
1418:Olson’s gap
1161:Varanopinae
998:Varanopidae
771:Caseasauria
711:Captorhinus
633:Below is a
619:M. efremovi
594:M. efremovi
582:M. efremovi
570:M. efremovi
376:Description
344:M. efremovi
308:Mezen River
293:Varanopidae
266:M. efremovi
195:Subfamily:
189:Varanopidae
3850:Varanopids
3844:Categories
3699:See also:
3655:Kamagorgon
3645:Dinosaurus
3609:Cynodontia
3598:Cynodontia
3425:Therapsida
3412:Therapsida
3355:see below↓
3346:Therapsida
3276:Dimetrodon
3224:Shashajaia
3008:Milosaurus
2940:Ophiacodon
2787:Dimacrodon
2754:see below↓
2702:Ruthiromia
2687:Aerosaurus
2597:Nitosaurus
2590:Dendromaia
2562:Apsisaurus
2487:Martensius
2473:Euromycter
2254:see below↓
2034:(2): 162.
1493:References
940:Ophiacodon
412:premaxilla
330:locality (
324:Artinskian
3613:includes
3523:includes
3442:Raranimus
3297:Neosaurus
2998:Ianthodon
2723:Varanodon
2583:Eoscansor
2494:Oromycter
2438:Caseopsis
2431:Caseoides
2369:Oedaleops
2299:Synapsida
2286:Synapsida
2245:Synapsida
2166:Tetrapoda
2152:Kingdom:
2136:Synapsida
2048:0045-8511
1931:0031-0220
1874:0024-1164
1831:0272-4634
1788:133846070
1780:0031-0220
1744:1943-2682
1655:0031-0182
1589:633882190
751:Synapsida
635:cladogram
629:Phylogeny
623:M. romeri
598:M. romeri
586:M. romeri
516:M. romeri
508:M. romeri
479:Discovery
447:pterygoid
350:Etymology
340:M. romeri
258:M. romeri
176:Synapsida
149:Kingdom:
143:Eukaryota
3775:10670041
3761:Q3278592
3755:Wikidata
3721:Category
3283:Eosyodon
3268:Cutleria
3137:Haptodus
3127:Cutleria
3052:Gordodon
2807:Gorgodon
2797:Driveria
2730:Watongia
2716:Varanops
2639:Cabarzia
2632:Anningia
2387:Caseidae
2362:Eothyris
2160:Chordata
2158:Phylum:
2154:Animalia
2098:Archived
2013:17105089
1976:17105089
1939:91871872
1839:59023934
1690:84706899
1633:: 1–11.
1555:31824730
1460:See also
1434:diapsids
497:skeleton
457:quadrate
410:Slender
332:Oklahoma
304:Kazanian
250:Species
182:Family:
163:Chordata
159:Phylum:
153:Animalia
139:Domain:
3814:4128063
3801:1255093
3788:4819442
3615:mammals
2466:Eocasea
2175:Amniota
2056:1439431
2004:1273351
1967:1273351
1854:Lethaia
1811:Bibcode
1714:Bibcode
1706:Geology
1635:Bibcode
1546:6837192
690:Amniota
592:Though
563:Permian
320:Permian
289:amniote
208:Genus:
31:Permian
2605:Pyozia
2054:
2046:
2028:Copeia
2011:
2001:
1974:
1964:
1937:
1929:
1872:
1837:
1829:
1786:
1778:
1742:
1688:
1653:
1587:
1553:
1543:
453:Stapes
314:, and
312:Russia
3827:37777
3796:IRMNG
2424:Casea
2171:Clade
2052:JSTOR
1935:S2CID
1835:S2CID
1784:S2CID
1686:S2CID
943:spp.
714:spp.
406:are:
387:Skull
285:genus
170:Clade
3783:GBIF
3607:see
3584:see
3549:see
3517:see
3491:see
3465:see
2231:see
2204:see
2044:ISSN
2032:1957
2009:PMID
1972:PMID
1927:ISSN
1911:PalZ
1888:link
1870:ISSN
1827:ISSN
1776:ISSN
1760:PalZ
1740:ISSN
1651:ISSN
1585:OCLC
1551:PMID
596:and
44:Preęž’
3770:EoL
2036:doi
1999:PMC
1962:PMC
1919:doi
1862:doi
1819:doi
1768:doi
1730:hdl
1722:doi
1678:doi
1643:doi
1631:475
1608:doi
1541:PMC
1533:doi
338:).
287:of
3846::
3824::
3811::
3798::
3785::
3772::
3757::
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2096:.
2092:.
2076:.
2050:.
2042:.
2030:.
2007:.
1993:.
1970:.
1956:.
1933:.
1925:.
1915:93
1913:.
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1884:}}
1880:{{
1868:.
1858:34
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1728:.
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1710:38
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1583:.
1563:^
1549:.
1539:.
1527:.
1523:.
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1440:.
641::
625:.
506:.
366:.
334:,
326:)
306:)
172::
94:Pg
38:Ma
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2114:v
2058:.
2038::
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1978:.
1941:.
1921::
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1876:.
1864::
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1821::
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