A 3D geometric morphometric analysis of digging ability in the extant and fossil cingulate humerus

Digging ability in armadillos has been shown to be closely related to the relative length of the olecranon process of the ulna. This study uses geometric morphometrics to examine the relationship between humeral shape, digging ability and size in a range of living and fossil cingulates. The extant species in the sample include representatives of 11 species of armadillo, while the fossil specimens include three species of fossil armadillos ( Peltephilus, Proeutatus and Eutatus ) and three Glyptodonts ( Propalaeohoplophorus, Glyptodon and Neosclerocalyptus ). The results show that in general, living species with good digging ability have larger sites for muscle attachment, particularly the proximal tubercles and the crests descending therefrom, and the epicondylar region at the distal end of the humerus. Some differences were found in the smallest armadillo ( Chlamyphorus truncatus ), which seems to have a different method of digging. The proportions of the olecranon process would indicate good digging ability in some glyptodonts, but humeral features do not fit with this interpretation and the differences may be related to large size. The relationship between cingulate phylogeny and humeral morphology is also examined, and it seems that while cingulates are to some extent constrained by their phylogeny, many of the humeral features are directly related to digging.     

The armadillo: a model for the neuropathy of leprosy and potentially other neurodegenerative diseases

Leprosy (also known as Hansen’s disease) is an infectious peripheral neurological disorder caused by Mycobacterium leprae that even today leaves millions of individuals worldwide with life-long disabilities. The specific mechanisms by which this bacterium induces nerve injury remain largely unknown, mainly owing to ethical and practical limitations in obtaining affected human nerve samples. In addition to humans, nine-banded armadillos (Dasypus novemcinctus) are the only other natural host of M. leprae, and they develop a systemically disseminated disease with extensive neurological involvement. M. leprae is an obligate intracellular parasite that cannot be cultivated in vitro. Because of the heavy burdens of bacilli they harbor, nine-banded armadillos have become the organism of choice for propagating large quantities of M. leprae, and they are now advancing as models of leprosy pathogenesis and nerve damage. Although armadillos are exotic laboratory animals, the recently completed whole genome sequence for this animal is enabling researchers to undertake more sophisticated molecular studies and to develop armadillo-specific reagents. These advances will facilitate the use of armadillos in piloting new therapies and diagnostic regimens, and will provide new insights into the oldest known infectious neurodegenerative disorder.     

THE MODE OF LIFE OF HYPSILOPHODON, THE SUPPOSEDLY ARBOREAL ORNITHOPOD DINOSAUR

Previous discussions of the mode of life of Hypsilophodon (Lower Cretaceous, England) are reviewed, and it is concluded that this primitive ornithopod was cursorial. There are no specific adaptations for an arboreal mode of life, and the hallux was not opposable. Metatarsal I was closely applied along its length to metatarsal II, and the first digit moved parallel to or. slightly away from the other digits. Relative to the trunk, the hind limb is long, with an elongate tibia and metatarsus, so that the proportions of the different regions fall in the range for cursorial mammals.     

Tuataras and salamanders show that walking and running mechanics are ancient features of tetrapod locomotion

The lumbering locomotor behaviours of tuataras and salamanders are the best examples of quadrupedal locomotion of early terrestrial vertebrates. We show they use the same walking (out-of-phase) and running (in-phase) patterns of external mechanical energy fluctuations of the centre-of-mass known in fast moving (cursorial) animals. Thus, walking and running centre-of-mass mechanics have been a feature of tetrapods since quadrupedal locomotion emerged over 400 million years ago. When walking, these sprawling animals save external mechanical energy with the same pendular effectiveness observed in cursorial animals. However, unlike cursorial animals (that change footfall patterns and mechanics with speed), tuataras and salamanders use only diagonal couplet gaits and indifferently change from walking to running mechanics with no significant change in total mechanical energy. Thus, the change from walking to running is not related to speed and the advantage of walking versus running is unclear. Furthermore, lumbering mechanics in primitive tetrapods is reflected in having total mechanical energy driven by potential energy (rather than kinetic energy as in cursorial animals) and relative centre-of-mass displacements an order of magnitude greater than cursorial animals. Thus, large vertical displacements associated with lumbering locomotion in primitive tetrapods may preclude their ability to increase speed.     

Dental microwear in the orthodentine of the Xenarthra (Mammalia) and its use in reconstructing the palaeodiet of extinct taxa: the case study of Nothrotheriops shastensis (Xenarthra,Tardigrada, Nothrotheriidae)

The utility of orthodentine microwear analysis as a proxy for dietary reconstruction in xenarthrans (tree sloths, armadillos) was quantitatively and statistically accessed via low‐magnification stereomicroscopy. Features such as number of scratches and pits, as well as presence of gouges, hypercoarse scratches, > four large pits, > four cross scratches, and fine, mixed or coarse scratch texture were recorded in 255 teeth from 20 extant xenarthran species. Feature patterns are consistent with scar formation through abrasional (tooth–food) and attritional (tooth–tooth) contact. Number of scratches is the most dietary diagnostic microwear variable for xenarthrans, with herbivorous sloths characterized by > ten scratches and nonherbivorous armadillos by < ten scratches. Discriminant function analysis differentiated arboreal folivores (sloths) and frugivore‐folivores (sloths) both from each other and from fossorial carnivore‐omnivores (armadillos) and insectivores (armadillos). Microwear patterns in carnivore‐omnivores and insectivores are difficult to distinguish between; armadillo microwear may reflect a fossorial lifestyle (grit consumption) rather than primary diet. Cabassous centralis is anomalous in its microwear signal relative to all other insectivores. To test the utility of orthodentine microwear analysis as an indicator of palaeodiet in extinct xenarthrans, microwear in the ground sloth Nothrotheriops shastensis was quantitatively and statistically compared to microwear in extant taxa. Microwear patterns in N. shastensis are most comparable to extant folivores based on scratch number and hierarchical cluster analysis. This strongly supports an herbivorous diet for N. shastensis that is corroborated by multiple independent lines of evidence. Thus, orthodentine microwear analysis can be used to reconstruct diet in extinct xenarthrans. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 156 , 201–222.     

Life on the Half-Shell: Consequences of a Carapace in the Evolution of Armadillos (Xenarthra: Cingulata)

Without doubt, the possession of an armored carapace represents one of the most conspicuous morphological features of all cingulates. Here, we review some of the many ways in which the carapace may have influenced the evolution of other features of extant armadillos (Xenarthra: Cingulata). Effects range from physiological impacts on respiration and thermoregulation, to mechanical and other constraints on reproduction. Additionally, in mammals, armor has been linked with relatively slow plantigrade locomotion, which in turn may have promoted the low metabolic rate and exploitation of a low quality diet typically observed in armadillos. Finally, this network of relationships may help to explain the lack of any obvious kin-selected altruism in the polyembryonic armadillos, such as the nine-banded armadillo (Dasypus novemcinctus), because of time and energy constraints associated with a short active period devoted almost exclusively to feeding. In mammals, there has been growing interest in describing an ecological “lifestyle” as the particular way in which each species makes its living, and how this lifestyle constrains the evolution of other phenotypic traits. Based on our review, it appears the carapace has been a major determinant of the lifestyle of armadillos and has played a central role in shaping the evolution of many other features of these animals.     

Earthworms near leprosy patients' homes are negative for acid-fast bacilli by fite stain,providing no link between leprous armadillos (Dasypus novemcinctus) and human leprosy

Enzootic leprosy has been recognized in armadillos in Louisiana since 1975. Contact with armadillos is being assessed as a risk factor for leprosy in three white women, lifelong residents of separate rural areas in northern Louisiana, which is a region without endemic leprosy. None has had any known exposure to human leprosy. Each was aware of armadillos (Dasypus novemcinctus) near or under her home for decades. In considering Possible environmental sources forMycobacterium leprae, we observed that all three had earthworm growth areas for fishing bait where soil was kept moist near their homes. The worms attracted armadillos. Since armadillos subsist on worms, grubs, and insects and because of the common feature of a worm farm near each home, we reasoned that earthworms might containM. leprae and be part of a cycle involving the armadillo and human beings. Worms from each home worm farm were studied. One site was sampled twice at patient 1's home, five sites were sampled once at patient 2's home, and three sites were sampled once at patient 3's home. A sample consisted of 3–4 worms, which were washed, purged, fixed live in 10% formalin, embedded in paraffin, sectioned, and stained with the Fite stain. Each was sagittally sectioned and examined by three independent observers. No acid-fast bacilli or other acid-fast structures were identified. We conclude that it is unlikely that earthworms are an environmental source or reservoir ofM. leprae.     

Effects of subsidized spiders on coastal food webs in the Baltic Sea area

The aim of this study was to examine top-down effects of cursorial spiders in subsidized coastal food webs. Top-down effects were examined by selectively removing cursorial spiders, mainly wolf spiders, from small islands (26–1834 m²) during 2004–2007. The removal success varied among islands and years, and spider densities were reduced by 30–65%. To examine treatment effects, arthropods were sampled using a vacuum sampling device at three occasions each summer. The densities of other arthropod predators, especially web spiders and carabids, were higher on islands where cursorial spiders had been removed compared to control islands. This treatment effect probably occurred through a combination of competitive release and reduced intraguild predation from cursorial spiders. No treatment effects were found on herbivore or detritivore densities and plant biomass. This lack of effect may either be because spiders indeed have fairly weak effects on herbivore and detritivore densities on Baltic shorelines or that the removal success of spiders was insufficient for observing such effects. Treatment effects may also be weak because negative effects exerted by spiders on herbivore and detritivore populations were balanced by increased predation by insect predators.     

Burrowing owls and burrowing mammals: are ecosystem engineers interchangeable as facilitators?

Terrestrial vertebrates exhibit dynamic, positive interactions that form and dissolve under different circumstances, usually with multiple species as participants. Ecosystem engineers are important facilitators of other species because they cause physical changes in the environment that alter resource availability. Although a species can be associated with more than one partner, facilitators may not be interchangeable if they differ in abundance, behavioral characteristics, or interactions with other factors in ways that condition the outcome of the association. We examined interactions between burrowing owls (Athene cunicularia) and two burrowing mammals, hairy armadillos (Chaetophractus villosus) and plains vizcachas (Lagostomus maximus), and determined whether these ecosystem engineers are interchangeable for owls. We examined reproductive success for owls nesting in these mammal burrows, constructed a logistic regression model to identify habitat characteristics associated with owl nests, and examined the engineering activities of the mammals. Data on reproduction and habitat indicate that armadillos and vizcachas are not interchangeable for owls. Thirty-five percent of the nests in vizcacha burrows produced fledglings; no fledglings were produced from nests outside vizcachas colonies, even though owls nest successfully in armadillo burrows in other parts of Argentina. Vizcachas facilitate burrowing owls by construction of burrows and by producing open understory vegetation through herbivory. In contrast, armadillos do not alter vegetation, and their burrows are suitable for nest sites only when they occur in recently burned areas or areas maintained by anthropogenic disturbance. Our habitat model also suggests that fire plays a key role in maintaining owl populations because fire is the only natural process that reduces shrubs to the level required by owls. Current management practices of eradication of vizcachas and fire suppression in shrublands could have strong negative consequences for burrowing owls.     

The phylogenetic position of the Pterosauria within the Archosauromorpha re-examined

A previous analysis of the phylogenetic position of the Pterosauria argued that pterosaurs were not closely related to dinosaurs as is generally accepted, but rather were outside the crown group Archosauria. However, that study was dismissed for the use of inappropriate methods. Here, the data set from that analysis was divided into five partitions: one with characters associated with cursorial digitigrade bipedal locomotion and the other four with characters from the skull and mandible, postcranial axial skeleton, forelimb and hindlimb, respectively. The partitions were subjected to homogeneity testing, and the Cursorial partition was found to be incongruent with other partitions and all other characters at the α = 0.01 probability level. Deletion of the Pterosauria removed all significant incongruence, demonstrating that the incongruence results from the coding of pterosaurs for the cursorial characters. The cause of the incongruence was interpreted as homoplasy in hindlimb morphology, and after re-evaluating and reformulating the characters of the Cursorial partition, the revised data set was tested for homogeneity and no significant incongruence was found. Lastly, the data set was updated with additional characters and taxa from recent analyses, tested as before, and when analysed suggested that the Pterosauria were basal archosauriforms well outside the crown group Archosauria.     

Guild mobility affects spider diversity: Links between foraging behavior and sensitivity to adjacent vegetation structure

The movement patterns of species may affect their susceptibility to modified habitat structure. It is likely that sedentary species perceive habitat features at smaller spatial extents compared to mobile species, but there is a lack of experimental research on the effects of fine-scale habitat characteristics on organisms of differing mobility. Spiders display two basic mobility levels based on foraging behavior: web-building species are restricted to specific sites whereas active hunters are mobile. We collected spiders inhabiting sagebrush shrubs with a structurally enhanced, unmodified, or removed understory, to examine (1) whether habitat structure in the immediate vicinity of shrubs affected cursorial and web spiders differently in terms of abundance and species richness and (2) which genera most contributed to changes in community composition. Shrubs without understory had reduced cursorial spider densities and species richness compared to shrubs with added and unmodified understories, whereas web spiders lacked significant responses to treatments. Community-level differences based on relative abundance of genera were detected in cursorial spiders but not in web spiders, despite a strong contribution of the web-building genus Theridion to community dissimilarities. Our results support the hypothesis that sedentary organisms may be sensitive to contiguous habitat at finer spatial scales than cursorial organisms, and highlight the risks associated with only collecting local habitat information when studying mobile species.     

Ecological differences between two sympatric species of armadillos (Xenarthra,Mammalia) in a temperate region of Argentina

Morpho-physiological specialization related to foraging can act as constraints on behaviour and ecological patterns of abundance and distribution. We tested this prediction in two species (weeping or screaming hairy armadillos Chaetophractus vellerosus Gray, 1865 and mulitas Dasypus hybridus Desmarest, 1804) that represent the two subfamilies of armadillos (Dasypodidae, Xenarthra): Euphractinae and Dasypodinae. The first subfamily possesses a well-developed masticatory apparatus that is considered primitive within the Xenarthra, while Dasypodinae show reductions in various aspects of cranial morphology. We sampled signs (burrows and foraging holes) and captured both species on 34 farms randomly selected in the north-eastern Pampas region of Argentina. We analyzed several niche dimensions. The two species showed significant differences in habitat use, seasonal patterns and diet. Weeping armadillos were generalists in their diet and active throughout the year. They were associated with calcareous-sand soil, as expected for a burrowing animal of the deserts. They tolerated a wide range of environmental conditions. We found that mulitas are myrmecophagous, and that they demonstrated a reduction in body temperature and activity during the cold season. As expected from their geographical distribution, mulitas used mainly vegetation with high cover and were associated with humic soils. Niche segregation between the two species of armadillos appeared to originate from different degrees of dietary specialization.     

Changes in hydrolytic enzyme activities in armadillos infected with Mycobacterium leprae

The activities of hydrolytic enzymes in various organs of armadillos infected with Mycobacterium leprae were compared with those in normal armadillos. Except for aspartate aminopeptidase and esterase, the levels of the other enzymes in liver, spleen and inguinal lymph nodes were significantly higher in armadillos infected with M. leprae compared with those in non-infected ones. These enzymes levels were at a maximum when the animals were sacrificed 22 to 30 months post-inoculation, a period when the bacterial load in the animals had also reached a maximum. Animals infected with M. leprae but not showing any signs of disseminated infection behaved similar to those in the non-infected group. The observed changes in enzymatic activities were not due to bacterial enzymes and so can be related to tissue damage caused by M. leprae.     

A dynamic similarity hypothesis for the gaits of quadrupedal mammals

The dynamic similarity hypothesis postulates that different mammals move in a dynamically similar fashion whenever they travel at speeds that give them equal values of a dimensionless parameter, the Froude number. Thus, given information about one species, it could be possible to predict for others relationships between size, speed and features of gait such as stride length, duty factor, the phase relationships of the feet and the patterns of force exerted on the ground.
Data for a diverse sample of mammals have been used to test the hypothesis. It is found to be tenable in many cases when comparisons are confined to quadrupedal mammals of the type described by Jenkins (1971) as "cursorial". Most mammals of mass greater than 5 kg are of this type. Although the hypothesis applies less successfully to comparisons between cursorial and non-cursorial mammals it is shown to be a reasonable approximation even for such comparisons and for comparisons between quadrupedal mammals and bipedal mammals and birds.     

The nine-banded armadillo (Dasypus novemcinctus) is naturally infected with Sarcocystis neurona

Sarcocysts were dissected from the tongue of a nine-banded armadillo (Dasypus novemcinctus). DNA was extracted and characterised by PCR amplification followed by restriction fragment length polymorphism analysis and nucleotide sequencing. A total of 1879 nucleotides were compared; the sarcocyst DNA sequence was identical to that reported for Sarcocystis neurona. DNA was extracted from the sarcocysts of five more nine-banded armadillos. A 254-nucleotide sequence was determined for each and found to be identical to S. neurona. Western blot techniques for detection of anti-S. neurona antibody were developed for use with armadillo plasma and samples from 19 wild-caught and 17 captive-raised armadillos were examined. Whereas all of the 19 wild-caught armadillos had antibodies to S. neurona, only one of 17 captive-raised armadillos did. These results suggest that the nine-banded armadillo are naturally infected with S. neurona.     

Technique,Safety, and Efficacy of Intra-Abdominal Transmitters in Nine-Banded Armadillos

ABSTRACT The future management of nine-banded armadillos (Dasypus novemcinctus) requires solid space-use and activity data, which are currently lacking and which radiotelemetry can provide. External radiotransmitters have not been successful applied with this species. To make recommendations for intra-abdominal radiotransmitter placement in nine-banded armadillos, we 1) evaluated 4 different anesthetic protocols for safety, efficiency, and cost-effectiveness; 2) evaluated a surgical technique for the intra-abdominal placement of radiotransmitters that addresses problems described in previous studies; and 3) evaluated the physiologic and behavioral effects of such a technique. We captured and surgically implanted 37 nine-banded armadillos using either butorphanol and isoflurane, ketamine alone, ketamine and xylazine, or a combination of butorphanol, ketamine, and medetomidine for anesthesia. We recovered and necropsied armadillos after the completion of the study. The objective and subjective assessment of butorphanol, ketamine, and medetomidine combination protocol, followed by reversal of the anesthesia with atipamezole, showed that it was the best overall anesthetic protocol for field use, providing both a smooth induction and fast recovery. We evaluated the fate and effects of radiotransmitters on 13 recovered animals at the end of the study and found no adverse effects. We recommend the implantation of radiotransmitters that are allowed to free-float within the abdominal cavity and specifically emphasize the need for strict aseptic technique. Wildlife managers and wildlife veterinarians aiming to implant nine-banded armadillos with radiotransmitters will benefit from using the recommended anesthetic protocol and surgical technique in future studies.     

Female reproductive tract of the lesser anteater (Tamandua tetradactyla,myrmecophagidae, Xenarthra). Anatomy and histology

The morphological and histological features of the unusual reproductive tract of the female lesser anteater, Tamandua tetradactyla (Myrmecophagidae, Xenarthra), are described for the first time. The present study aimed to establish the main similarities and differences between this species and other xenarthrans. The populations of this species are declining rapidly for a number of reasons and our study is relevant to diverse programs related to its conservation. Studies were carried out on five female genital tracts of adult specimens. Ovaries were ovoid, presenting a medulla completely surrounded by the cortex, differently from that described in other xenarthans. Like in Dasypus but different from all other armadillos studied, single oocyte follicles were observed and a simple the uterus. The uterovaginal canal connects the uterus with the urogenital sinus. The simple columnar epithelium of the uterovaginal canal ends abruptly at a septum which resembles a hymen, where the transitional epithelium of the urogenital sinus appears. This ancestral feature is shared with that of other armadillos, except Tolypeutes matacus, which has a true vagina. Characteristics of the reproductive tract and sperm morphology of other Xenarthra are comparatively discussed. These observations suggest that important reproductive features are shared between the family Myrmecophagidae and the genus Dasypus, a basal group in the phylogeny of Xenarthra. J. Morphol. 2011. © 2011 Wiley‐Liss, Inc.     

The evolution of fossoriality and the adaptive role of horns in the Mylagaulidae (Mammalia: Rodentia)

Ceratogaulus, a member of the extinct fossorial rodent clade Mylagaulidae, is the only known rodent with horns and the smallest known horned mammal. The function of the large, dorsally projecting nasal horns on this burrowing animal has been the subject of wide speculation among palaeontologists; suggested uses range from sexual combat to burrowing. Mammals have evolved adaptations for digging repeatedly; horns and other cranial appendages have also evolved numerous times. These two adaptations co-occur in mammals extremely rarely: only two fossil genera (Ceratogaulus and the xenarthran Peltephilus) and no extant mammals are both horned and fossorial. Tracing the evolution of fossoriality in aplodontoid rodents (the larger clade to which Ceratogaulus belongs) reveals that Ceratogaulus descended from ancestors who dug by head-lifting. Whereas this suggests an obvious explanation for the horns of this rodent, evidence from functional morphology, anatomy, phylogeny and geologic context indicates that the horns in Ceratogaulus were used for defence, rather than digging, and evolved to offset increased predation costs associated with spending more time foraging above ground as body size increased.     

What do we know about armadillos? An analysis of four centuries of knowledge about a group of South American mammals,with emphasis on their conservation

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Pelvic limb musculature in the emu Dromaius novaehollandiae (Aves: Struthioniformes: Dromaiidae): Adaptations to high-speed running

Emus provide an excellent opportunity for studying sustained high-speed running by a bird. Their pelvic limb musculature is described in detail and morphological features characteristic of a cursorial lifestyle are identified. Several anatomical features of the pelvic limb reflect the emus' ability for sustained running at high speeds: (1) emus have a reduced number of toes and associated muscles, (2) emus are unique among birds in having a M. gastrocnemius, the most powerful muscle in the shank, that has four muscle bellies, not the usual three, and (3) contribution to total body mass of the pelvic limb muscles of emus is similar to that of the flight muscles of flying birds, whereas the pelvic limb muscles of flying birds constitute a much smaller proportion of total body mass. Generally, the pelvic limb musculature of emus resembles that of other ratites with the notable exception of M. gastrocnemius. The presence and arrangement of four muscle bellies may increase the effectiveness of M. gastrocnemius and other muscles during cursorial locomotion by moving the limb in a cranio-caudal rather than a latero-medial plane. J. Morphol. 238:23–37, 1998. © 1998 Wiley-Liss, Inc.