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Were mammals originally venomous?

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The extratarsal spur in extant monotremes consists of an os calcaris and a cornu calcaris. A poisonous extratarsal spur occurs only in the platypus (Ornithorhynchus); a possibly secondarily non−poisonous spur is present in echidnas (Tachyglossus and Zaglossus). Some therian mammals (e.g., bats), reptiles (Chamaeleo), and amphibians have a spur−like structure in the ankle, but this is not homologous to the extratarsal spur of monotremes. Among fossil mammals, the co−ossified os calcaris and ossified cornu calcaris have been found in the eutriconodontan Gobiconodon and in the spalacotheroid “symmetrodontan” Zhangheotherium. Here we describe the os calcaris in several multituberculate mammals from the Late Cretaceous of the Gobi Desert, Mongolia. The multituberculate os calcaris is a large, flat bone, generally similar to that in males of the extant monotreme species, but the cornu calcaris is not ossified. In Gobiconodon and Zhangheotherium the ossified cornu calcaris is fused to the os calcaris probably to provide the bony support for the keratinous spur. We hypothesize that the os calcaris in these Mesozoic mammal groups is homologous to that of monotremes. However, the extratarsal spur has not been found in non−mammalian cynodonts nor in other synapsids. A platypus−like os calcaris might be an apomorphic characteristic of basal Mesozoic mammals and is secondarily lost in crown therians; the os calcaris is confirmed to be absent in well−preserved tarsal structures of the earliest known crown therian mammals. We speculate that the os calcaris, the cornu calcaris, and its associated venom gland might have served the function of a defensive structure during the “dark ages” of mammalian history, when dinosaurs ruled the Earth. This structure is a plesiomorphic character retained in extant monotremes and cannot be used as an autapomorphy of Monotremata.
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The limb posture in early mammals is a matter of controversy. Kielan−Jaworowska and Gambaryan presented arguments for a sprawling posture in multituberculates, based mainly on three characters of the hind limbs (deep pelvis, mediolateral diameter of the tibia larger than the craniocaudal, and position of MtV, which fits the peroneal groove on the calcaneus and is not aligned with the axis of tuber calcanei). Here we present two more arguments for sprawling hind limbs in early mammals. One is the presence of an os calcaris, supporting the probably venomous spur in hind legs of docodontans, multituberculates, eutriconodontans, and “symmetrodontans”, similar to those of extant monotremes. We argue that early mammals (except for boreosphenidans) had sprawling limb posture and venomous spur; acquisition of the parasagittal stance was apparently characteristic only of boreosphenidans, in which the spur has not been found. The second argument is based on taphonomic evidence from lacustrine conditions (e.g., Early Cretaceous Jehol Biota), in which the mammalian skeletons, except for boreosphenidans (Sinodelphys and Eomaia), have been preserved compressed dorso−ventrally, suggesting sprawling stance. In similar conditions of the Eocene Messel Biota the skeletons of boreosphenidan mammals (except for bats and pangolins) are preserved lying on flanks, suggesting parasagittal stance. Sereno argued that forelimbs in multituberculates were parasagittal, based on the stated presence of a ventrally facing glenoid, a mobile shoulder joint, and an elbow joint with enhanced flexion−extension capability. However, these characters are not unequivocally indicative of parasagittalism. We demonstrate that the structure of the distal end of the multituberculate humerus is condylar, with no tendency for developing a trochlea. We reconstruct multituberculates and other early mammals with sprawling stance in resting position as plantigrade.
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