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Here we examined the kinematic function of the morpho- logically unique first thoracic vertebra in giraffes. The first thoracic vertebra of the giraffe displayed similar shape to the seventh cervical vertebra in general ruminants. The flexion experiment using giraffe carcasses demonstrated that the first thoracic vertebra exhibited a higher dorsoventral mobility than other thoracic vertebrae. Despite the presence of costovertebral joints, restriction in the intervertebral movement imposed by ribs is minimized around the first thoracic vertebra by subtle changes of the articular system between the vertebra and ribs. The attachment area of musculus longus colli, mainly responsible for ventral flexion of the neck, is partly shifted posteriorly in the giraffe so that the force generated by muscles is exerted on the cervical vertebrae and on the first thoracic vertebra. These anatomical modifications allow the first thoracic vertebra to adopt the kinematic function of a cervical vertebra in giraffes. The novel movable articulation in the thorax functions as a fulcrum of neck movement and results in a large displacement of reachable space in the cranial end of the neck. The unique first thoracic vertebra in giraffes provides higher flexibility to the neck and may provide advantages for high browsing and/or male competition behaviours specific to giraffes.
1. Introduction
The mammalian vertebral column comprises successive units organized into five series that are highly conserved and easily recognized: cervical, thoracic, lumbar, sacral and caudal. Cervical vertebrae are traditionally distinguished from thoracic vertebrae by the absence of movable rib articulations. The number of cervical vertebrae has remained constant at seven for at least 200 Myr, despite variable counts in other regions of the mammalian vertebral column and in the cervical vertebrae of other vertebrate classes [1–3]. The diversity of neck length and shape in mammals has evolved under the rigorous constraint of cervical number [3]. The long neck of giraffes also follows the cervical constraint; therefore, each cervical vertebra is prominently longer than that of the common short-necked ruminants, including okapi [4,5].
A previous study raised the possibility that giraffes had gained an additional rib-bearing cervical vertebra in association with their neck elongation [6]. Skeletal comparisons between giraffe and okapi revealed morphological transformations of the vertebrae between the seventh cervical vertebra (C7) and second thoracic vertebra (T2) in giraffes. The most notable transformation is observed at C7. In okapi, C7 is obviously differentiated from other cervical vertebrae by its exhibition of an elongated neural spine and lack of the transverse foramen and ventral tubercles. Conversely, C7 in the giraffe presents the characteristics that are generally observed between C3 and C6: it contains the transverse foramen, ventral tubercles, short neural spine and vertebral body elongation [6–8]. In the case of giraffes, various morphological features generally observed in C7 are found in T1. For example, the vertebral length of T1 of the giraffe represents an intermediate value between the cervical and other thoracic vertebrae [4,6]. The facet type of vertebral articulation is transitional from the radial type to the tangential type at the joint between T1 and T2 in the giraffe, whereas the transitional point of articulation lies between C7 and T1 in okapi, which represents the typical ruminant pattern [6,7]. Additionally, the configuration of roots of the brachial plexus supports the structural similarity between T1 of the giraffe and C7 of okapi [6]. From the osteological and neurological evidences, previous studies have proposed the idea that T1 of giraffe is homologous with C7 of okapi [6,7].
Despite many morphological similarities between T1 of the giraffe and C7 of okapi, prior studies have criticized the idea of a rib-bearing cervical vertebra in giraffe by the presence of the movable rib in T1: a conclusive criterion for classifying a vertebra as a thoracic vertebra [4,5,9,10]. Additionally, the ribs articulating to T1 connects to the sternum. Therefore, T1 of the giraffe has been regarded as merely one of the thoracic vertebrae, and the functional insight of this transformation of the vertebral shape has not been mentioned. Revealing the specific function of these modified vertebrae will allow us to understand the evolutionary and ethological advantages of the morphological transformations of the vertebrae in giraffe. Given the general linkage between the vertebral shape and mobility in vertebrates [11–24], morphological similarities imply that T1 of the giraffe bears a kinematic resemblance to C7 of okapi. The C6/C7 and C7/T1 joints are highly specialized, facilitating the ventral flexion of the head–neck complex in quadrupedal mammals [25]; hence, we can presume that giraffe may have gained an additional movable joint contributing to the ventral flexion of the neck into the T1/T2 joint. Here we evaluated the ability of the ventral flexion of vertebrae around the cervicothoracic area in giraffe and described the musculoskeletal structure of the neck to reveal a functional significance of the morphological shift of the vertebral column.
2. Material and methods
2.1 Analyses of sagittal mobility of each vertebra
For evaluating the mobility of each vertebral joint, we conducted the flexion experiment using giraffe and okapi carcasses. We forcibly moved their neck from the maximum ventral flexion posture to the maximum upraised posture, and recorded the positional relationship of each vertebra in both postures by using photography and computed tomography. We calculated the angle of a vertebra relative to the posterior adjacent vertebra in the cases of the maximal dorsal and ventral flexion of the neck (figure 1b). The angle between two adjacent vertebrae was measured with reference to the line connecting the anterior edge and posterior edge of the articular process in the vertebrae (figure 1c). The motion range of each vertebra in a dorsoventral direction was evaluated by the difference between the angle in the dorsal and ventral flexion. In one giraffe and okapi, the angle was calculated from three-dimensional data obtained by computed tomographic imaging. We constructed the three-dimensional model from the computed tomography images by using the Osirix imaging software (v. 6.5.2; http://www.osirix-viewer.com/ContributionOsiriX.html), and then measured the mobility of each vertebral joint. In the other giraffes, we removed some epaxial muscles located on one side to observe the positional relationships of the articular processes, which were then photographed from a lateral view in both postures (figure 1b). In this study, we measured the motion range of the two most caudal cervical and first five thoracic vertebrae.
2.2 Description of musculoskeletal structure
We described the morphological relationships between the thoracic vertebrae and ribs in giraffe and okapi to reveal the osteological restriction on intervertebral flexion movement imposed by ribs. The costovertebral joints were observed using dried skeletons of giraffe and okapi. For inferring the effect of rib articulation against vertebral movement, we described the positions of costal foveae on thoracic vertebrae and the positional relationships between two adjacent vertebrae and a rib.
We dissected giraffe and okapi carcasses and then described the muscular system generating the force resulting in the ventral flexion of the vertebrae for inferring the active ability of the vertebral flexion of giraffe. Gross anatomical dissections of giraffe and okapi were conducted sequentially from the superficial to the deep layer. Specimens were fresh-frozen and thawed before dissection. We focused on musculus longus colli, located on the ventral surface of the vertebral body, that is the main muscle responsible for ventral flexion of the neck [26]. Previous studies about the anatomy of the giraffe have noted that musculus longus colli represents a normal structure in giraffes; however, they reported only the structure of the cervical part of the muscle without addressing the muscular structure around the cervicothoracic area displaying the particular morphological transformations of the vertebrae [27,28]. Solounias [6] mentioned differences in the attachment area of the muscle between giraffe and okapi, although the muscular structure was not reported in detail. We described the origins and insertions of the muscle in giraffe and okapi, following which the active mechanism of lowering their head and neck was discussed.
The terminology of the vertebrae and muscles was consistent with the prior works of the vertebral column of Giraffidae [6,8].
2.3 Specimens
In the flexion experiment, we used the carcasses of two giraffes and one okapi, with all being former zoo animals. All were newborn specimens of a few days old, possessing the intact axial skeleton because infant specimens allow easier conduct of the flexion experiment. The skeletal structure around the cervicothoracic area was described by observing dried skeletons of eight adult giraffes and five okapis. Data of the muscular structure were obtained from the carcasses of six giraffes and one okapi. Individual information regarding the specimens used in this study is listed in table 1.
3. Results
3.1 Estimated sagittal mobility of each vertebra
Our flexion experiment revealed that the motion range of each vertebra gradually declined through three successive vertebrae at the cervicothoracic area and that the pattern of decline was different between giraffe and okapi (figure 1a). In giraffe, the estimated ranges of motion were clearly large in C6 and C7 and decreased progressively in the caudal direction from C7 to T2. T1 of the giraffe exhibited an intermediate motion range between C7 and T2 and possessed a greater mobility than the remaining thoracic vertebrae (see the electronic supplementary material, movie S1). The motion range of each vertebra maintained a low value between T2 and T5 (figure 1a). Conversely, in okapi, the reduction of the motion range was confirmed between C6 and T1. The motion range of C7 showed an intermediate value between C6 and T1, and the ranges between T1 and T5 represented a low mobility (figure 1a). Our results showed that the decreasing point of the vertebral mobility was shifted posteriorly in giraffes, and that the intermediate mobility was observed in T1 of the giraffe and C7 of okapi. This indicates the possibility that giraffes have acquired an additional movable articulation into the T1/T2 joint and that the kinematic characteristic of T1 of the giraffe is similar to C7 of okapi in the dorsoventral movement of the neck.
3.2 Skeletal structure
All thoracic vertebrae of giraffes and okapi, with exception of T1 of the giraffe, possessed three costal foveae at a lateral surface of each vertebra: cranial costal fovea, caudal costal fovea and costal fovea of the transverse process. The cranial costal fovea was located on the lateral side of the outer border of the cranial convex extremity of the vertebral body, and the caudal costal fovea was positioned on the exterior edge of the caudal concave extremity of the vertebral body. However, the cranial and caudal costal foveae in T1 of the giraffe were not confirmed. T1 possessed an isolated costal fovea located on the central area of the lateral side of the vertebral body below the transverse process.
According to our skeletal observations, the capitulum of the first rib in giraffe made contact with the isolated fovea and did not touch the caudal part of the vertebral body of C7 (figure 2a,c). The second rib articulated with T2 at the cranial costal fovea and costal fovea of the transverse process, without contacting the vertebral body of T1 (figure 2b,d). In the third rib, the capitulum of the rib was jointed to the socket formed by the caudal costal fovea of T2 and the cranial costal fovea of T3, and the tuberculum of the rib was attached to the costal fovea of the transverse process of T3. For ribs positioned more caudal than the third rib, the costovertebral joints represent the same articulation system as that of the third rib. Namely, in giraffes, thoracic vertebrae typically possess ribs at the cranial and caudal joints of one vertebra. The T1 of giraffes has ribs only at the central area of the lateral side of the vertebral body without disturbing both joints of the vertebra.
In okapi, the costovertebral joint between the first rib and T1 displayed a structural similarity to the joint between the second rib and T2 in giraffes: the capitulum of the rib was attached at the cranial costal fovea of T1 without touching the caudal edge of the vertebral body of C7. In all ribs except the first, the capitulum of the rib was jointed to the socket formed by the caudal and cranial costal foveae of the two adjacent vertebrae, and the tuberculum of the rib articulated with the costal fovea of the transverse process. Thus, the thoracic vertebrae of okapi, including T1, generally possess the ribs on the cranial and caudal joints of the vertebral body.
3.3 Muscular structure
In giraffes, the musculus longus colli comprised partly fused fascicles covering the ventral aspect of vertebral bodies of cervical and thoracic vertebrae from atlas to T7 and was divided into cervical and thoracic portions (figure 3). The cervical part arose on the concave ventral surfaces from C2 to T2 and was composed of six incompletely separated bundles. The five cranial bundles originated from the ventral edge of the ventral tubercles between C3 and C7, were directed craniomedially and partially terminated on the ventral spine of two preceding vertebrae by tendinous fibres. The most caudal and deepest bundle arose from the hypapophysial tubercles of T1 and T2, following a parallel course on the ventral surface between C7 and T2. The thoracic part comprised two fascicles that were partly fused. The medial part of the bundle arose from the hypapophysial tubercles between T2 and T4; it inserted on the caudal edge of the dorsal tubercle of C6 as a tendon. The lateral part of the bundle originated from the convex surfaces between T2 and T7 and terminated on the dorsal tubercle of C7 as a tendon. The muscle attached to the ventrolateral surface of the vertebral body of T1 without tendinous origin and insertion.
In okapi, the muscle arose on the ventral surface of the vertebral bodies from the atlas to T6; the cervical part attached on all cervical vertebrae and the first thoracic vertebra, and the thoracic part extended from the six most cranial thoracic vertebrae to C6 (figure 3). The components of the cervical part were five bundles; the four cranial bundles were located on the ventral surface from atlas to C5 and were directed craniomedially, and the most caudal bundle was distributed on the ventral area between C6 and T1 and ran parallel to the vertebral column. No obvious tendinous insertion was confirmed in C7. The thoracic part of the muscle arose on the ventral surfaces of the vertebral bodies from T1 to T6 and terminated on the caudal edge of the ventral lamella of C6 as two tendinous fasciculi. This muscle was located on the ventrolateral space of C7 without tendinous origin and insertion. A comparison of the muscular structure of the giraffe and okapi revealed that the attachment area of musculus longus colli in the giraffe was expanded posteriorly from T1 to T2 in the cervical part and that the origin and insertion of the muscle were partly shifted posteriorly in the thoracic part (figure 3).
4. Discussion
4.1 Behavioural advantages of the additional movable vertebra in the giraffe neck
A long neck has evolved independently in various taxa and consequently plays an important role in expanding the reachable space during foraging, improving predator detection and displaying dominance in competing males [21,29]. The long neck of the giraffe is the most famous example for understanding the evolution of neck elongation. Its neck has been considered to have evolved as a result of the adaptive evolution to high browsing behaviour and/or combat behaviour between males [30–32]. The height of the head would significantly influence their fitness. In giraffes, the maximum reachable height of the head is elevated by their long necks and limbs [33]. The forelimbs of giraffes are longer than the hind limbs; therefore, their trunk slopes from the cranial to the caudal direction, although the common ruminants, including okapi, show a horizontal trunk [9,34]. The pronounced slope of the trunk in the giraffe facilitates elevating the position of the cervicothoracic transition of the axial skeleton, resulting in a rise of the entire neck. The adaptive modifications of the body plan of the giraffe pose difficulties when lowering the head to the ground surface level for drinking water [33]. As a consequence, the development of a method of lowering the head and neck to facilitate the extension of the dorsoventral reachable space of the head was necessitated.
The flexibility of the axial skeleton is generally determined by two variables: the total number of movable vertebrae and the maximum mobility of each vertebral joint [35,36]. The mammalian thoracic region is rigid to facilitate respiration during locomotion and to counteract loading forces transmitted from the limbs [37,38]. Previous studies assumed that the majority of the head and neck movement occurred in the cervical region [16,21,39,40], suggesting that the total number of movable vertebrae is at a constant seven in the mammalian neck. Nevertheless, our study proved the expansion of the motion range in C7 of the giraffe and the mobility acquisition in T1 of the giraffe (figure 1a).
The head and neck of mammals are considered collectively as a loaded beam that is supported at one end only by attachment to the cranial end of the trunk region [11]. The cervical spine is regarded as an extended third-class lever: the resistance occurs on one side of an effort force, whereas the fulcrum is located on the other side [41]. Our findings suggest that the T1/T2 joint complex in giraffes acts as a fulcrum in this system. Conversely, the C7/T1 joint complex works as a fulcrum in okapi. It is predicted that a small increment of the dorsoventral mobility in the C7/T1 and T1/T2 joints in the giraffe would result in a large displacement at the cranial end level of the cervical spine, owing to their extraordinarily long neck.
4.2 Kinematic characteristics of C7 and T1 in the giraffe
The articulation of the rib is a principal factor restricting intervertebral flexion [38]. The rib typically articulates with the vertebrae through two positions in mammals: the costal fovea of the transverse process and the socket formed by the caudal and cranial costal foveae of two adjacent vertebrae [42]. This pattern was confirmed in the rib articulations in giraffe and okapi and imposes a fixed distance between the caudal costal fovea of a vertebra and the transverse fovea of the posterior adjacent vertebra, thereby imposing a strict limitation on vertebral flexion [38]. However, in the giraffe, the capitula of the first and second ribs did not make contact with the vertebral bodies of the adjacent vertebrae; consequently, in this region, the immobilization distance resulting from the rib articulation did not affect the vertebral flexion in the C7/T1 and T1/T2 joints, suggesting that the change of articular positions of the capitula of ribs in giraffes might minimize the restriction of vertebral mobility imposed by ribs around T1.
The attachment area of musculus longus colli in giraffes demonstrates that the rotational force generated by the thoracic part of this muscle tends to be concentrated to their C6 and C7 by the strong tendons (figure 4). Although the force might act on T1, it would be lower than the power exerted in C6 and C7 because of the absence of the tendinous insertion in T1 of giraffes. Conversely, in okapi, the muscle would work more intensively on C6 than on C7, as C7 does not possess tendinous insertion (figure 4). The T1 of okapi does not possess the insertion; therefore, the muscle does not produce the rotational force on the vertebra. The vertebral movement inferred by the muscular structure is consistent with the result of our flexion experiment.
According to previous osteological studies, the C7 of giraffes possesses a long vertebral body, a round articular surface at the caudal joint between the vertebral bodies and large articular facets directed laterally on the posterior articular processes (table 2) [4–7]. These characteristics are generally observed in the vertebrae from C2 to C6 and could be regarded as the features representing high intervertebral flexibility, based on prior studies on the relationships between the vertebral shape and mobility [16,17,20,22,23]. The thoracic vertebrae from T2 to T14 display the vertebral shape representing low intervertebral mobility [16,17,20,22,23]. Namely, they present a short vertebral length, a flat articular surface at the caudal joint of the vertebral body, a long neural spine and small articular facets directed medially, just beneath the neural spine with no processes (table 2) [6,7]. Additionally, the T1 of giraffes represents the intermediate characteristics between the general cervical and thoracic vertebrae [6,7]. The vertebral body of T1 in giraffes is slightly longer than the other thoracic vertebrae and shows an intermediate length between the cervical and thoracic vertebrae [4]. T1 possesses two paired, large articular facets on the small posterior articular processes beneath the neural spine [7]. The vertebra presents a flat articular surface on the joint between T1 and T2. The long vertebral body, the large articular facets, and the posterior articular process would contribute to facilitating a higher vertebral mobility than that of general thoracic vertebrae [17,20,22,23]; however, the flat articular surface would enhance the rigidity between the joints [20]. This suggests that T1 of the giraffe potentially possesses a mobility that is intermediate of that between the general cervical and thoracic vertebrae.
Conversely, the T1 of okapi possesses the same vertebral shape as the remaining thoracic vertebrae, representing low vertebral mobility [6]. In okapi, the intermediate shape is confirmed in C7 [6], suggesting that C7 of okapi presents lower mobility than the remaining cervical vertebrae. The vertebral morphology corresponds with our results regarding the expansion and acquisition of the vertebral flexibility in the C7/T1 and T1/T2 joints in the giraffe.
Solounias [6] hypothesized that the vertebra traditionally identified as T1 might be actually regarded as 'the eighth cervical vertebra' based on specific anatomical characteristics of the general cervical vertebrae; however, the first thoracic vertebra has been overlooked as merely one of thoracic vertebrae as it possesses the movable ribs. Our study provided anatomical evidence that T1 of the giraffe played a kinematic role corresponding to the C7 of general ruminants even if the vertebra articulated with the sternum via the movable ribs. Note that, in this study, there is no evidence supporting that T1 is originally cervical vertebra. Our results indicate that the morphological shift of the vertebrae of giraffes at the cervicothoracic area has great significance in understanding evolutionary functional plasticity of the mammalian vertebral column rather than in discussing the number of cervical vertebrae.
4.3 Developmental background of the morphological shift of the vertebral column
The musculoskeletal structure of the axial skeleton consists of two elements derived from different developmental backgrounds: primaxial and abaxial elements [43–46]. The vertebrae, vertebral ribs and epaxial and hypaxial muscles develop only from somitic mesoderm and are classified as the primaxial elements. The sternum and limbs originate from lateral plate mesoderm and are identified as the abaxial elements [43]. The abaxial domain contains somitic cells that migrate into the lateral plate environment early during development and mix with lateral plate cells [43]. Curiously, in mice, the first rib is completely identified as an abaxial element [46]. The expression patterns of Hox genes, regulating the morphology of the vertebral column and rib cage [45,47–51], are independent between the primaxial and abaxial domains [43,50,52]. In the abaxial domain, the migrating somitic cells adopt the Hox expression of the lateral plate mesoderm into which they migrate [53]. Additionally, a primaxial patterning possesses a colinearity of the phenotype, whereas an abaxial patterning is not colinear [50]. Colinearity is the phenomenon that the order of the Hox genes along the chromosome corresponds with the order of the expression domains of the genes along the anterior–posterior axis of the embryo. Mutations of the Hox genes cause the homeotic transformation in primaxial elements, whereas it results in serious patterning defects in the abaxial elements [50,54–56].
In the light of the above evidence obtained from previous genetic studies, we considered the possibility that the mutations of Hox5 and/or Hox6 specific to the primaxial domain would have resulted in the transformations of the musculoskeletal structure in the neck of giraffes. Both genes are important for the specification of the identity of the vertebral column between C6 and T1 [57]. Hox6 genes function to induce ribs by regulating Myf5/Myf6 expression, suggesting that their expression pattern is responsible primarily for the regionalization of the vertebral column between the cervical and thoracic vertebrae [51]. When we consider the genetic background of the development of the vertebrae and ribs in giraffes along with the evidence obtained from mice, it could be assumed that the mutations of the Hox5 and/or Hox6 genes specific to the primaxial domain caused the morphological transformations of the vertebrae and ribs, with the exception of the first rib. From the study of mice, it has been predicted that the development of the first rib would be regulated by the Hox expression pattern in the abaxial domain [46], indicating that the phenotype of the first rib is unaffected by the mutations of the Hox genes specific to the primaxial domain. Therefore, it could now be explained that the mutations of the Hox genes specific to the primaxial domain causes the morphological transformations of the vertebrae in giraffes without displacing the position of the rib cage.
5. Conclusion
We quantitatively demonstrated the expansions of the motion range in C7 and T1 of giraffes by the flexion experiment. The muscular system generating a vertebral rotation was partly modified around the cervicothoracic region, which supported the result of the estimation of the vertebral mobility. The modifications of the attachment position of the capitula of the first and second ribs in giraffes minimize the restriction on vertebral flexion imposed by ribs, thereby enabling notable flexion of the joint between T1 and T2. The addition of movable articulation to the thoracic region results in a large displacement of the head, even if the mobility is relatively smaller than that of the cervical joints. Based on our findings, we assert the evolutionary importance of the increased mobility at the base of the neck in terms of the advanced adaptation to high browsing in giraffes. Under the rigorous cervical constraint, giraffes have achieved the functional novelties at the cervicothoracic region by subtle modifications in their existing musculoskeletal system.
Recent studies have reconsidered the regionalization of the vertebral column by focusing not on the presence of movable ribs but on the morphological characteristics of the vertebrae [58–60]. Tree sloths are well known to exhibit an abnormal number of cervical vertebrae: Bradypus (three-toed sloths) possess eight to ten cervical vertebrae, whereas Choloepus (two-toed sloths) possess five to eight cervical vertebrae [58]. However, the morphological characteristics and the ossification pattern of the vertebrae have demonstrated that the boundary between the cervical and thoracic was identified between seventh and eighth vertebrae, as is the case in other mammals [58,59,61]. These studies regarded the abnormal number of cervical vertebrae in sloths as rib-less thoracic vertebrae in the neck or rib-bearing cervical vertebrae in the thorax [59]. They proposed the idea that the constraint of the cervical vertebrae was actually conserved even in the sloths that had historically been the most famous group thought to deviate from the constraint [58]. This idea offered new insight, clearly different from the previous perception, into the regionalization of the vertebral column in mammals. Our findings additionally provide evidence that the vertebral regionalization in giraffes is slightly shifted posteriorly at the cervicothoracic transitional region without changing the traditional border, based on the presence of movable ribs. This suggests the possibility that the vertebral regionalization is additionally diversified in other mammals that maintain the number of cervical vertebrae. We advocate that subtle mutations of the Hox genes can induce the striking structural and functional novelties in the conservative existing system.
The flexibility of the neck is of major importance in inferring the neck posture and the feeding strategy in extinct long-necked species, including sauropods [29,62]. Many studies have evaluated the flexibility of the sauropod neck in the light of vertebral morphology [16,19,23,62–66], whereas they have not paid any attention to thoracic vertebrae. However, in diplodocid sauropods, it was reported that the morphological characteristics representing high intervertebral flexibility were found in three to four most anterior thoracic vertebrae [67]. This indicates the possibility that diplodocid sauropods possess movable thoracic vertebrae. The dorsoventral movement of the thoracic vertebra effectively enlarges the displacement at the level of the head. The estimated reachable area of the head will strongly depend on whether we consider the thoracic vertebrae. The evidence that the giraffe gained a movable joint in the thoracic region emphasizes the need for evaluating the flexibility of the thoracic vertebrae. Our discovery provides new insight into the evolutionary process leading to the development of the neck of the giraffe and into the reconstruction of neck function and posture in extinct long-necked animals.
Data accessibility
The datasets supporting this article have been uploaded as the electronic supplementary material. The movie of the flexion experiment in a giraffe is uploaded as the electronic supplementary material, video S1. The three-dimensional models constructed from computed tomography images are provided as the electronic supplementary material, videos S2–S5.
Authors' contributions
M.G. designed research, performed the dissection and estimation of vertebral mobility, and wrote the manuscript. H.E. contributed to the description of muscular structure and to gathering fresh carcasses of giraffe and okapi.
Competing interests
The authors declare that no competing interests exist.
M.G. was supported by JSPS fellowship for young scientists (research project no. 14J04974) and the Prince Takehiko Yamashina grant for financing the trip to the conference and investigation (Yamashina Institute for Ornithology).
We are grateful to S. Kawada, N. Kurihara (NSMT) and H. Taru (KPM) for allowing us to dissect the fresh carcasses of giraffes and okapi. Giraffe specimens were collected by the support of Key Research Project 'Studies on functional morphology and phylogeny' of NSMT. We acknowledge J. Lesur, C. Bens (MNHN), E. Hoeger and E. Westwig (AMNH) for access to skeletal specimens under their care. We thank N. Solounias (New York Institute of Technology), Y. Nakajima (The University of Tokyo) and T. K. Yamada (NSMT) for helpful discussions. We thank two anonymous reviewers for valuable comments to the draft.
- Received November 9, 2015.
- Accepted January 6, 2016.
© 2016 The Authors. Published by the Royal Society under the terms of the Creative Commons Attribution License http://creativecommons.org/licenses/by/4.0/, which permits unrestricted use, provided the original author and source are credited.


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MEMPHIS, Tenn. (AP) - Despite the loss of his three top scorers, Memphis' John Calipari was confident Thursday as he and other Conference USA coaches got together to talk about the upcoming season.
"We've got a good core of guys coming back, six guys, and we've got a good core of newcomers, so I think we're going to be fine," Calipari said.
In their preseason predictions, conference coaches picked the Tigers to be the top team in the league, as it was last season. Gone from last year's 33-4 squad are Rodney Carney, Shawne Williams and Darius Washington, who combined for 53 percent of the Tigers' scoring and led Memphis to a No. 1 seed in the NCAA playoffs. Calipari will be counting heavily on sophomores Chris Douglas-Roberts, Antonio Anderson and Robert Dozier.
"I don't know if they're all going to be scorers, but those are the three who will play probably as many minutes as they can play," Calipari said at the league's preseason media day.
The Tigers will also have senior guard Jeremy Hunt back in the lineup. Hunt was suspended from the team last season after he was charged with slapping and kicking a former girlfriend. He was allowed to avoid a criminal trial on a condition of staying out of further trouble. Calipari said Hunt will contribute badly needed experience.
"Two years ago or three years ago, he may have been the best guy we had on our team," Calipari said, "but he's been hurt throughout his career. He's just been beat up."
Hunt is healthy entering the new season. Conference coaches put Houston at No. 2 in preseason standings, and Texas transfer Dion Dowell said coach Tom Penders is preparingthe Cougars for a fast-paced attack.
"I'm glad I made the move," said Dowell, who still must sit out the fall semester. "I'm looking for a controlled, run-and-gun offense. It's a scorer's dream."
Tulane coach Dave Dickerson said his team appears to be over the trauma of being run out of New Orleans by Hurricane Katrina.
"Last year at this time, we came to this very event being displaced at Texas A&M," Dickerson said. "What a difference a year makes. Being home has served us well. Our kids love the university. They want to be there."
With the loss of leading scorer Quincy Davis, Dickerson expects to spread the scoring around.
"Our perimeter play has gotten better and our overall team speed has gotten better. It will allow us to do more things than we did last year," he said. "This year, we'll depend on a more fast-paced, perimeter attack as well as an inside game."
Mike Davis, UAB's new coach, said he's had only a few practices with the Blazers and is still "going home every night trying to figure out what to do the next day."
Any major adjustments in the Blazers' style?
"We'll have to wait and see," said Davis, who spent six seasons at Indiana. "Right now, it's too early to tell."


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Comparing Higher Ed to Wall Street
Whenever worried leaders of for-profit colleges have implied in recent months that the U.S. Education Department is gunning for the institutions, officials of the federal agency have discouraged such talk, offering evenhanded rhetoric about treating all sectors the same in their push for increased accountability.
Whenever worried leaders of for-profit colleges have implied in recent months that the U.S. Education Department is gunning for the institutions, officials of the federal agency have discouraged such talk, offering evenhanded rhetoric about treating all sectors the same in their push for increased accountability.
The words have provided little reassurance to the colleges, since they haven't always seemed to square with the aggressive approach the Obama administration is taking in rewriting federal rules governing vocational and other programs.
On Wednesday, in a speech to state regulators who oversee for-profit colleges, the chief architect of the Education Department's strategy, Robert Shireman, offered a much more critical assessment of the private sector institutions than he has in his public comments to date, according to accounts given by several people who were in the room. He compared the institutions repeatedly to the Wall Street firms whose behavior led to the financial meltdown and called them out individually, one by one, for the vast and quickly increasing sums of federal student aid money they are drawing down.
While Shireman's comments were aimed most directly at the for-profit colleges themselves, they may be most noteworthy for his indictment of accreditation, higher education's system of institutional peer review. In Shireman's narrative before the annual meeting of the National Association of State Administrators and Supervisors of Private Schools, the accrediting agencies are to the for-profit colleges what the Wall Street ratings agencies were to the misbehaving financial firms: entities charged with regulating an industry that has grown too quickly and too complex for them to control, and that have an "inherent conflict of interest" because their existence depends on financial contributions from those they regulate.
Accreditors lack the "firepower" to regulate the for-profit sector, and the states and the federal government don't necessarily have all the tools they need to do it either, Shireman said, according to the notes of several in the audience. That, he suggested, is why the Education Department must toughen its rules in the way it is now proposing.
Shireman could not be reached for comment, and an Education Department spokesman said its officials did not wish to comment on this article.
To several people in the audience, Shireman's comments represented a much more candid (and critical) appraisal of the for-profit sector than he has offered publicly since he became deputy under secretary of education almost exactly a year ago. Many supporters of the education companies feared his appointment because they believed his track record as an advocate for low-income students and a foe of student debt would result in a crackdown on the institutions, whose students are disproportionately needy and disproportionately go into heavy debt to finance their educations.
But with Wall Street analysts hanging on his every word looking for snippets that might threaten the publicly traded companies' stock prices, Shireman has often seemed to go out of his way to avoid singling the institutions out for criticism.
A typical quotation, from last summer: "Our overall goal at the Department of Education in postsecondary education is to make sure that students ... have the information they need to make good choices, and that they have good quality postsecondary education that serves both them as students and taxpayers as well," Shireman said. "...If there is not quality, we want to know about it and if we can, we want to do something about it. Whether that involves a public institution, a nonprofit, a for-profit, a two-year, a four-year, a trade program, whatever type or sector of institution, we want to do all we can to make sure that we have good quality."
Different Tone
In his comments Wednesday, Shireman laid out the context underlying the Obama administration's elevation of higher education as a central focus of its domestic policies. The economic slide created in part by the collapse of the credit markets has sent Americans streaming back to college in record numbers, and has made it more imperative than ever that more Americans get a higher education to strengthen the country's economic base for the future, Shireman said.
The administration has poured tens of billions of dollars into Pell Grants and restructured the federal student loan programs to try to ensure that Americans have access to higher education, Shireman said Wednesday. Many public institutions, facing cuts in their state funding, have had to limit or even cut their enrollments, reducing their ability to meet the increasing demand from students.
The for-profit colleges, by contrast, have stepped up, seeing their enrollments explode -- and with them, the amount of Pell Grant money that follows the students to the institutions, Shireman said. Anyone in the audience from Corinthian Colleges? Shireman asked the assembled audience Wednesday.
A hand went up. The California-based for-profit higher ed company has seen its revenue from Pell Grants grow by 38 percent in the first three quarters of this fiscal year compared to the last one, he said. Anyone from DeVry? Forty-two percent increase, Shireman said. ITT? Strayer? One by one, he ticked through a list of publicly traded companies, pointing out the increasing amounts of federal money the institutions were collecting ("It was like fourth grade, with a teacher scolding students over their grades," said one person who was in the room).
What are taxpayers and students getting in return for that investment? Shireman asked. It has historically been up to the "triad" -- the three-headed regulatory scheme involving the federal government, state governments and accrediting agencies -- to ensure access, quality and integrity in higher education, he said.
But is that regulatory system up to the job? To draw a parallel, Shireman noted that as this meeting was unfolding in St. Paul, politicians back in Washington were debating possible reforms of Wall Street, to try to fix the "flawed" regulatory process that allowed Goldman Sachs and other purveyors of subprime mortgages to engage in misbehavior that helped devastate the economy.
One major reason the process was flawed, Shireman said, was because the bond rating agencies that were supposed to be judging the riskiness of the financial instruments were supported in large part by fees from the companies that were asking the agencies to rate the financial instruments -- "a clear, inherent conflict of interest," Shireman said, according to the accounts of several in the room.
On top of that inherent conflict, the ratings agencies have been struggling to keep tabs on industries that grew quickly and adopted increasingly complex practices, Shireman said, suggesting that the ratings agencies lacked the "firepower" to regulate the financial markets.
In case anyone missed it, Shireman drove his point home, pointing out that higher education accrediting agencies are made up of (and financially supported by) their member colleges, and see it as their mission both to help the institutions "improve" and also to ensure, in what is essentially a subcontract from the federal government, that they are of sufficient quality. They are nonprofit, unlike the ratings agencies, but they are run by the institutions they regulate, in ways that the credit agencies aren't.
The peer review nature of higher education accreditation has an inherent conflict of interest similar to the ratings agencies, Shireman said. Given that, he suggested, it is crucial for state and federal agencies, as the other two parts of the triad, to step up their role in regulating higher education.
But do state regulators think they have the "firepower" to keep tabs on the big, growing and complex private market college sector? Shireman asked the state officials in the room. The response was underwhelming. "I don't think we feel we have the firepower we need," Shireman said, referring to the federal government's own powers, according to members of the audience.
The bottom line of Shireman's talk, he said, was that "federal and state governments cannot rely on accreditors to assure that consumers and taxpayers are protected to full extent that they need to be. All three legs" of the three-legged stool of higher education quality assurance need to be operating effectively, he said.
Shireman went from there into a review of the department's proposed new approaches to ensuring integrity in the financial aid programs, such as requiring most for-profit colleges and non-degree vocational programs at nonprofit colleges to show that they are preparing students for gainful employment.
Several people who heard the speech said they viewed it as a much more strident critique of for-profit colleges, and of higher education accreditation, than Shireman has delivered before. But David Dies, who heads the Wisconsin Educational Approval Board and just finished a term as president of the national group of state regulators, didn't hear it quite that way.
"I think Bob was explaining why we need state regulation and [Education] Department oversight to be part of this three-legged stool, not just accreditation, and why we all need to work together," said Dies. "He was pointing out some limitations of accreditation, but I didn't really see it" as highly critical of accreditors or for-profit colleges.
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Business License FAQs
Am I required to have a city and county business license?
State law requires that all businesses inside the City of Chattanooga have both city and county business licenses with exceptions for professionals, manufacturers, and certain disabled persons. City Business Applicatons are available. For information on obtaining a Hamilton County business license call (423) 209-6500.
What is the cost of a business license?
A business owner is assessed an initial license fee of $15.
How to renew a business license?
Business licenses are renewed with the State of Tennessee.
Where do you obtain a sales tax number?
Tennessee Department of Revenue
540 McCallie Avenue Suite 350
Chattanooga, TN 37402
(423) 634-6266
How can I get an application for business license?
For your convenience the Business Tax License Application is available for download and/or printing.
What type of permits are available?
City of Chattanooga requires permits for the following...
Going out of Business
Sidewalk Vendors (Push Carts)
Special Gathering Permits
What if my business sells beer or alcohol?
Beer - Contact Beer Board at 423-643-5156
Liquor by the Drink - Contact TN ABC board at 423-643-6434
What if I want to be on the City's wrecker list?
Contact Wrecker Board at 423-643-5156
What payment types are accepted?
Payments for NEW Business Licenses and Permits can be made by cash, personal check, money order or credit card.
City Treasurer's Office
101 East 11th Street Suite 100
Chattanooga, TN 37402 (map)
(423) 643-7262
Vacant, City Treasurer
Sharon Morris, Assistant Treasurer
For inquiries regarding property taxes
423-643-7262 or email firstname.lastname@example.com
For inquiries regarding sewer fees
Dial 311 or 423-643-6311 or email email@example.com
For inquiries regarding business license
423-643-7262 or email firstname.lastname@example.com


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|The ubiquitous African daisy.
|That's a Pelergoniumn not Geranium
I still haven't left my back yard yet and after nearly 100mm of rain in what is typically our driest month of the year everything is looking absolutely fabulous.
It certainly is an exciting time of the year with buds bursting every where.
The promise of fresh peas in their pod uummm.
I had to sneak a veggie in here somewhere. Radish that'll be ready in a week or two.
Cheers and bye for now,
The best time to plant a tree was 20 years ago. The second best time is now. Chinese proverb
Stewart says, The best time to plant a tree was yesterday. The second best time is today.


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As Jeffrey Kluger writes: "We're suckers for scale. Things that last for a long time impress us more than things that don't, things that scare us by their sheer size strike us more than things we dwarf. We grow hushed," he writes in his book Simplexity "at say, a star and we shrug at a guppy. And why not?" he asks. "A guppy is cheap, fungible, eminently disposable, a barely conscious clump of proteins that coalesce, swim about for a few months, and then expire entirely unremarked upon." He then suggests that "a star roars and burns across the epochs, birthing planets, consuming moons, sending showers of energy to the limits of its galaxy." Yet, he points out, "the guppy is where the magic lies." A star is just a furnace whereas a guppy is a symphony of systems. So it is with the world of finance.
And joining us today on the Financial Impact Factor Radio we have Roger Wohlner, Certified Financial adviser at Asset Strategy Consultants based in Arlington Heights, Ill., where he provides advice to individual clients, retirement plan sponsors, foundations, and endowments. He recently cofounded Retirement Fiduciary Advisors to provide direct investment and retirement planning advice to 401(k) plan participants. Roger also blogs at Chicago Financial Planner and columnist for USNews and World Report. He has been kind enough to join us today to help us sort through the complexity, the scale we are so impressed with.
That scale, that complex mechanism that Roger has agreed to speak about is the target date fund. If you use one, you should listen to this explanation. If you invest, you should tune in as well.
Listen to Financial Impact Factor Radio with your hosts:
Paul Petillo of Target2025.com/BlueCollarDollar.com and Dave Kittredge and Dave Ng of FinancialFootprint.com
The show is broadcast daily, online at 6amPST/9amEST.


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Coming soon on our very own '3 Beat Blue' is the full blown release of the massive Natural Born Grooves – Candy On The Dancefloor. With new mixes by Warren Clarke and the Wideboys adding to the existing mix package by Cahill and Alex K.
Then there will be the long awaited second single by Cahill called Sexshooter. With mixes by Cahill, Soul Avengerz, Dave Ramone, Agent X, Crazy Cousins and Outsource.
Promo cds and vinyl available soon… this can not be missed out on!!!


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Moscow: Maria Sharapova`s return from injury after an almost three-month break was cut short on Monday after the world number three suffered a wrist injury at the Wuhan Open in China.
The Russian, who missed the entire U.S. hardcourt season with a leg injury, was into a deciding set against Czech player Barbora Strycova when she called for the trainer.
Despite playing on for another couple of games she quit while leading 7-6(1) 6-7(4) 2-1.
The former world number one had not played a match since losing to Serena Williams in the Wimbledon semi-finals in July.


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Menthol-cigarette smokers may be at higher risk of having a stroke than those who smoke other types of cigarettes, a new study suggests. Women and non-black smokers are at highest risk, HealthDay reports.
Researchers found those who smoked menthol cigarettes had more than twice the risk of suffering a stroke than regular cigarette smokers. Women and non-black smokers had more than three times the risk, the article notes.
The way in which menthol cigarettes might increase stroke risk is not known, the researchers said.
Lead researcher Dr. Nicholas Vozoris of St. Michael's Hospital, in Toronto, said the findings do not indicate that any type of cigarette is safer than others. "There is no 'good' cigarette type," he said. "Smoking any kind of cigarette is bad for one's health, and serves to increase one's risk for a variety of cancers, heart diseases and lung diseases. However, this study shows that smoking mentholated cigarettes may place one at even higher risk for stroke than smoking regular, non-mentholated cigarettes."
The study appears in the Archives of Internal Medicine.
In 2011, a Food and Drug Administration advisory panel said a ban on menthol cigarettes could benefit public health, because the mint flavor seemed to make smoking more appealing. The panel did not conclude menthol cigarettes were more harmful than regular cigarettes.
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HomeFootballFirst TeamNewsStill in the hunt! (1-2)
Jordi Clos
26/02/2012 | 00:00
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Messi, celebrant l'1-2 contra l'Atlètic de Madrid. FOTO: MIGUEL RUIZ-FCB.
The footballing Gods were certainly fair to Barça tonight and a cheeky strike from Messi towards the end of the game gave them a thoroughly deserved win that keeps them in the chase for the title. After a dominant first half against an ultra defensive Atletico Madrid topped off by a Dani Alves strike, Barça were stunned after the break by Simeone's team, who got an early goal, but Guardiola's men kept their heads, withstood the pressure and hit back with Messi's winner.
Great start
Barça made perhaps their brightest start to a game all season and after Xavi shot just wide early on and Messi had a great goal disallowed for an earlier handball, they dominated their hosts and kept them penned back in their own half with some careful patient football.
Alves goal
Xavi then brought a great save from Courtois from a 20th minute free kick and the team were boasting an astonishing 83% of possession, with Atletico trying to claw their way back into the match with a more physical approach . It was the creative team who got the rewards though, as on 36 minutes Messi played a clever ball out left to Cesc and the Catalan's pinpoint pass set up Alves to score.
Equaliser changes the storryline
Although the goal spurred Atletico to come out of their shell and Valdes made a good save to cut out a dangerous looking attack, Barça immediately took control again and finished the half in the lead and clearly the better team. Then, just after the break, as Simeone pushed his lines forward for a more attacking approach, Atletico got the equaliser through Falcao after Busquets had failed to clear a corner. Now it was the home team who imposed their trademark more intense and vertical style of play on the game.
Cheeky Messi catches hosts cold
Guardiola reacted on the hour mark by switching to a 3-4-3 formation - a high risk strategy that nearly backfired when Falcao forced a great save from Valdes. Alexis then had a couple of good opportunities – one of which he put away, but was ruled out for handball - before Messi took the home team by surprise with a quick free kick before the wall was set to give Barça the lead with about ten minutes left to play.
The team defended bravely in the closing minutes with Valdes again standing strong. It was a great win for the team, who celebrated the win together in the centre circle.
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Cesc proving to be an artist of assists
Messi and Busquets to miss Sporting clash
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Anna Edney and Meg Tirrell
Wednesday, 13 November 2013 06:56 PM
While the U.S. Food and Drug Administration advisory panel decided that potential safety risks don't preclude approval of Lemtrada, its members voted 14-2 Wednesday that the drug didn't help improve a patient's disability. The agency isn't required to accept the recommendations of its advisory panels.
Lemtrada, approved in Europe earlier this year, was a key part of Paris-based Sanofi's $20 billion acquisition of Genzyme Corp. in 2011. The drug if cleared in the U.S. would enter a crowded field in which relapsing MS patients now have 10 treatment options with varying degrees of efficacy, including Biogen Idec Inc.'s Tecfidera and Teva Pharmaceutical Industries Ltd.'s Copaxone.
"If the study is biased, then everything that flows from the study can't be trusted," Robert Clancy, a panel member and professor of neurology and pediatrics at the University of Pennsylvania School of Medicine, said at Wednesday's meeting. The panel voted 11-6, with one abstention, that the two large studies Sanofi submitted on the drug's behalf weren't adequate and well-controlled.
Contingent Value
The FDA is expected to decide whether to approve Lemtrada by Dec. 27. Genzyme stockholders had received rights to additional fees of as much as $14 a share, in the form of contingent value rights, or CVRs, at the end of 2020 if Sanofi meets certain goals, most of them tied to the approval and sale of Lemtrada.
Sanofi declined less than 1 percent to 77.78 euros earlier Wednesday at the close in Paris. The tradeable CVR was halted during the vote.
Lemtrada may generate sales of $672 million in 2017, according to the average of eight analysts' estimates compiled by Bloomberg. Bayer AG plans to co-promote the drug and will receive payments based on sales.
Multiple sclerosis is a debilitating disease that attacks the central nervous system. Relapses, or flare ups, are episodes of worsening neurological function, according to the National MS Society.
Several patients who said they had severe MS testified Wednesday in favor of Lemtrada's approval, imploring panel members to leave the weighing of risk versus benefit to those who have the disease and their physicians.
Patient's Plea
"It's critical that people with significant disease who stand to lose function and become disabled have the opportunity to access this drug," said David Goldblatt, a neuroradiologist from Austin, Texas, who said he started taking Lemtrada 10 years ago.
"I understand your issues with safety," Goldblatt told the panel. "I think that nowadays patients are taking more responsibility and should be allowed to make the decision that they may decide that the risk/benefit ratio is adequate for them, that they would like to take this medication."
The panel agreed, with many members saying they voted that the drug's safety issues didn't preclude its approval because patients should be allowed to make that decision with their doctor. They also voted unanimously, though, that the drug should not be a first option for patients, if approved.
"For the foreseeable future, I would rank it as a third-line drug," said Justin Zivin, a panel member and professor emeritus at the University of California at San Diego.
Staff Report
FDA staff determined in a Nov. 8 report that Sanofi's annual infusion has "serious and potentially fatal safety issues" including risk of cancer and autoimmune and thyroid diseases. FDA staff also questioned Sanofi's claims the drug is effective. The company's decision not to keep secret which patients in clinical research were taking the medicine or an older treatment and the subjective nature of determining whether the drug was working may have skewed results, agency staff said. That was a main focus of the discussions Wednesday.
Sanofi said it couldn't keep the treatments confidential because of the differences in the drugs, including annual dosing compared with three times a week.
Lemtrada won European Union approval in September and the active ingredient alemtuzumab was cleared by the FDA in 2001 to treat a certain form of leukemia, though the drug is no longer for sale.
Other treatments for relapsing MS include another Sanofi drug Aubagio, Biogen's Tecfidera and Tysabri and Teva's Copaxone. Lemtrada is given through two courses of infusions given a year apart.
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Palace Hotel wins honours at 'Emirates Culinaire'
Dubai, March 2, 2008
The Palace Hotel, managed by Sofitel Luxury Hotels, made a strong debut at the Emirates Salon Culinaire which was hosted by Gulf Food's - one of the world's leading food, drink, foodservice and hospitality equipment exhibitions.
The food and beverage team of The Palace took part in the competition for the first time since its opening and made an impact by winning an impressive eight medals.
The Palace winners included Danushka Karunarathne who won two gold medals in the categories of 'Open Showpiece' and 'Pastry Showpiece' and also walked away with three bronze medals in 'Chocolate Carving Showpiece', 'Fruit and/or Vegetable Carving Showpiece' and the 'Live Vegetable Carving' compeition.
Vrushali Jadhav won two silver medals in the 'Three Tier Wedding cake' and 'Five Plated Dessert' categories. Hiran Jayamanne was rewarded a silver medal for his 'Five Plated Dessert'.
The Emirate Salon Culinare was a four-day culinary event that includes seminars, competitions and demonstrations that took place from February 24 to 28. – TradeArabia News Service


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After I wrote my last post about the first two days of school, I had a comment by one of my sweet long-term readers who asked if every day went as smoothly as those two days. Those two days did go smoothly, but certainly it is not always smooth. Sometimes it is super rough and awful. Or one child is having a hard time and it is impacting the flow of all the other children and our day. That is life homeschooling multiple children.
Part of life in homeschooling is also just life. This week involved going to the barn, our family attending (and me leading) a breastfeeding support group session, numerous calls and emails and such that needed to be returned after said meeting, two visits by friends to our home on separate days, a run to the allergist and grocery store, a visiting aunt who is here through the weekend to teach machine sewing and work on a mini-quilting project with my seventh grader (which is normally more eighth grade in a Waldorf School, but this particular aunt lives far away so I am happy to take her up on it now!), (our fourth grader also doing a mini project to help brush up on measurement skills and look at textiles and then will have a turn machine sewing in eighth grade for her very own), a husband who traveled out of state the majority of the week, and the pet care of two hamsters, fish, frogs, and a large dog plus meals and housekeeping. That is all life and part of homeschooling as well. Especially as your children grow older, they may have more activities or passions they are investigating and have distinctly different needs than the children in grades one through four. Life may expand outside the home, but being within the home is still the basis of homeschooling and the more you are home, the more smoothly things will run, in my experience.
So, how does one manage life and homeschooling? Continue reading


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Cary, NC, April 17, 2009 – cdv Software Entertainment USA and Ascaron Entertainment are pleased to announce that the action-RPG, Sacred 2: Fallen Angel, has gone gold for the Xbox 360 video game and entertainment system from Microsoft, and will be released on May 12, 2009. For more information about Sacred 2: Fallen Angel, please visit https://www.sacred2.com or https://www.cdvusa.com.
Sacred 2: Fallen Angel is a hack-and-slash action-RPG with a rich story that takes place in a gigantic seamless world. This world contains massive dungeons, treacherous opponents and hundreds of challenging quests. Intelligent enemies, which adapt in number and difficulty based on player progress, challenge gamers in heroic single- and multiplayer battles for up four players on next-generation consoles.
About cdv Software Entertainment USA
cdv Software Entertainment USA, a wholly owned subsidiary of cdv Software Entertainment AG, is a leading publisher of third-party video games and interactive entertainment with offices in New Jersey and headquarters in Cary, North Carolina. cdv Software Entertainment USA has published critically acclaimed and commercially successful titles including Divine Divinity, Blitzkrieg, American Conquest, Codename: Panzers, City Life, DarkStar One, War Front: Turning Point and the acclaimed Sherlock Holmes: The Awakened. The company will continue to expand its offerings with upcoming titles including the highly anticipated Sacred 2: Fallen Angel. For more information, please visit https://www.cdvusa.com.
About Ascaron Entertainment, GmbH
Ascaron Entertainment GmbH is one of the pioneers in the development of computer games. It is also one of the leading developers of computer games in Germany and has been prominent in the sales charts for more than a decade with titles such as Patrician, ANSTOSS, Vermeer, Port Royale, DarkStar One and the original SACRED. Ascaron Entertainment is headquartered in Gütersloh, Germany, and maintains an office in Potsdam, a development studio in Aachen, Germany, as well as an international department in Birmingham (UK). Currently, Ascaron Entertainment markets its products in more than 34 countries


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About Spectra
The Greater China Leader
SPECTRA Technologies, founded in 1993, is the Greater China Leader in design and development of electronic transaction solutions. Our state-of-the-art technologies include the world's first integrated cordless transaction solution, self-service electronic transaction solution, security magnetic card reader and detachable IC card and SAM card module. With the combination of innovation and pragmatism, SPECTRA Technologies' pioneering hardware and software won top honours at various award ceremonies.
Our Dedication and Commitment
With the rapid spread of smart cards, new concepts and applications emerge. We at SPECTRA Technologies take pride in our broad vision. Our people have the expertise and creativity necessary to anticipate market needs, and we strive to develop technologies with the future in mind.
We continue to place special emphasis on innovative and pragmatic solution development not just to keep you abreast with time, but also to ensure that your investment is protected. Using our hardware, software, comprehensive ranges of professional and customer services, you can count on smooth and effective operation with secured and advanced solutions.
Management Team
SPECTRA Technologies' management team is one of the localized experts, which centred in Asia's market. We at SPECTRA Technologies share the same goal of contributing to the public with quality living through bringing electronic efficiency and convenience to people. We have customers and partners around the world in more than 65 countries. Our management team includes:
Yeung Tat Wing – CEO & General Manager
Alfred Hung – Sales & Marketing Director, Asia Pacific
Timothy Wong – Director
Sam Or – General Manager of China Office


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Carlson Opens First Park Inn By Radisson Hotel In New Mexico
January 12, 2012 // Franchising.com // ALBUQUERQUE, N.M. – Carlson, a global hospitality and travel company, today announced the further expansion of the Park Inn by Radisson brand in North America with the opening of the Park Inn by Radisson Hotel & Water Park, Albuquerque, N.M.
"The opening of the first Park Inn by Radisson in New Mexico represents the continued momentum of our Ambition 2015 growth strategy and is aligned with plans to grow Park Inn by Radisson as a winning midscale brand in North America," said Thorsten Kirschke, president, Carlson Hotels, Americas.
The brand has grown to more than 121 hotels in operation globally with 61 hotels currently in development.
"We look forward to offering a bright, bold and fresh, uncomplicated, friendly and fun environment that provides both business and leisure travelers with the affordable hotel experience they desire," said Michael Gallegos, president and CEO, Pacific Pearl Hotel Management, LLC, owning company of the Park Inn by Radisson Albuquerque, N.M.
The 290-room hotel features a large on-site water park with water slides, a wave pool, lazy river, waterfalls and more. The water park can accommodate just over 700 people. The hotel also offers an indoor pool, a fitness center, a business center and free Wi-Fi access. Breakfast, lunch and dinner are available in Brio Grille Restaurant, the hotel's restaurant serving American and New Mexican cuisine. The hotel can accommodate events with 26,000 square feet of flexible meeting space.
The Park Inn by Radisson Albuquerque is located seven miles from Albuquerque International Sunport and offers complimentary shuttle service. For reservations and more information, visit http://www.parkinn.com/hotel-albuquerque
About Carlson
Carlson is a global hospitality and travel company headquartered in Minneapolis, Minn. Carlson encompasses 1,070 hotels, including, Radisson®, Country Inns & Suites By CarlsonSM, Park Inn by Radisson and Park Plaza®; more than 900 T.G.I. Friday's® restaurants; and a majority stake in Carlson Wagonlit Travel®, the global leader in business travel management. Carlson operates in more than 150 countries and its brands employ more than 170,000 people. http://www.carlson.com
About Pacific Pearl Hotel Management, LLC
Pacific Pearl Hotel Management, LLC., one of the nation's largest independent hotel companies, is based in San Diego, California and Albuquerque, New Mexico. With a portfolio value of over $2 billion, APMC currently owns and operates 30 lodging properties encompassing over 6,500 rooms and employs over 2,000 personnel throughout the United States. In addition to a successful network of independent hotels, boutique properties and resorts, APMC's hotel flags have included Radisson, Le Meridien, Sheraton, Hilton, Wyndham, Doubletree, Hilton Garden Inn and Holiday Inns. For more information on APMC, please visit our website at www.apmc.net.
Robert Davis
General Manager
+1 (505) 888-3311
Ben Gardeen
Carlson Hotels
+1 (763) 212-8129


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May 16, 18, 09:10 AM #1
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Small Claims AdviceOk so back in December I hired a guy to do my floors. To make a long story short the guy messed my floors up, agreed to pay me back for the materials I bought and the deposit I paid him up front. He then gave a different excuse each time we were supposed to meet up so I ended up taking him to Small Claims Court.
So I took him to Small Claims about a month and a half ago, he didn't show, so I won the case totaling around $1,500. Now I thought the battle was finally over and I'd be able to stop worrying about all of this since I had done everything on my own without any legal help. Little did I know that actually collecting the money would be a whole other process in its own.
So after winning the case I called the Circuit Clerk and asked how I would go about receiving the payment from the case. They essentially told me I'd have to research it on my own and that they couldn't give me any legal advice. So after googling what to do I found out that there was a 7 or 10 day grace period for an appeal.
I gave it about 2 weeks and called the County Sheriff to find out if they'd be able to help me. They gave me the phone number for the Circuit Clerk. I called them knowing that they wouldn't be of any help but figured I would at least try. Again no help from them. I called the Sheriff's office again and they gave me a phone number that may be able to help out. I can't remember who it was, I think maybe a law firm, but they essentially gave me the phone number for the Circuit Clerk and hung up on me before I could even get a word out.
So I googled some more and figured out that maybe a Wage Garnishment was my best route. So now I have the Wage Garnishment form and have no idea how I'm supposed to fill it out.
Not sure if I should just try my best on it and hope I can figure it out, or if I should contact a lawyer and hire them to do it.
Can anyone guide me in the right direction on what to do?Advertisement
Last edited by brown4; May 16, 18 at 09:35 AM.
May 16, 18, 09:28 AM #2Small claims is a bit of a farce. Hard to collect and almost not worth the hassle. It ends up being more out of principle. You might be able to place a lien on his home or business as an option, but once again, I'm sure the paperwork is a nightmare.
I'd also be posting this guys name so he does not scam anyone else??
Good luck!
May 16, 18, 09:32 AM #3
Is the guy self employed? If so, since he wouldn't receive a regular paycheck, it will be impossible to garnish his wages.
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Not to be a downer, but as TAC said, winning a debt case in small claims is the easy part. Actually getting paid is the hard part. You're likely one of many that's waiting to get paid by the guy, and if he had the money to satisfy his debts, you wouldn't have ended up in small claims court to begin with.
Last edited by rjs4470; May 16, 18 at 09:51 AM.
May 16, 18, 09:59 AM #4
That's what I'm afraid of. I've spent so much time and effort into this that I really don't want to give up based on the principle of it. If a wage garnishment is not the route to take, is there another way to go?
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May 16, 18, 10:06 AM #5Yeah that's rough. A judgment is just a piece of paper until you take the effort to enforce it.
Have you looked into an Non-wage garnishment? If you have any notion of where he might do his banking, you could file a non-wage garnishment and seize assets from his bank account(s). These are relatively inexpensive to serve on banking institutions so even if they come back with a notice that the debtor holds no funds there, you won't have lost much.
Given the size of the judgment, a writ of execution on personal property or a judgment lien may not be worth the trouble.
May 16, 18, 10:19 AM #6
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May 16, 18, 10:22 AM #7
File a lien on any property he has. You may never get funds if he doesn't sell or die, but if when he does you will. Its worth it and its not to bad of a process. You could also file a non wage garnishment (think bank garnishment here). It takes some time to process, but you will either get funds from it or a notice of no funds in account from the asset.
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May 16, 18, 10:34 AM #8
May 16, 18, 10:40 AM #9I'm with TAC, this clown needs to be named and drug through the mud.
May 16, 18, 10:52 AM #10
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The reasoning for not using my friend with his own company is a whole other story that essentially put me in this mess.


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Young men who want to become an Eagle Scout must perform a service-leadership project that totals 80 to 150 hours. A service project of that size is no small task, and Ben Freed, a 17-year-old Boy Scout from Wisconsin, wanted to come up with a great idea for his Eagle Scout Service Project that would have a lot of meaning.
After two project ideas that didn't pan out, Ben became interested in doing something for a local cemetery that would help preserve the records and make them available for others to research.
Ben's mother, Deborah, suggested he do a project for the White Creek Cemetery in Wisconsin. This cemetery has special significance for Ben and his family, because at least 1/3 of the burials in the cemetery are somehow related to them--including Ben's grandfather.
Ben's father did some online research and learned that they could create their own online digital cemetery map on Names In Stone, and that those records would be linked with FamilySearch, the world's largest genealogical database. That project sounded ideal.
Ben's Eagle Scout Service Project
The White Creek Cemetery is located in Adams, Wisconsin. It is an older cemetery, with tombstones dating from the late 1850s, some with barely legible inscriptions. The cemetery is still in use, with newer monuments along the north and west edges of the cemetery. In 2002, an acre of land was donated to the White Creek Cemetery Association, which will allow the cemetery to be enlarged and a new fence added.
The Cemetery is a wonderful American memorial, with many veterans buried there, including veterans from the Spanish-American War, the Civil War, World War I, World War II, and the Gulf War.
Ben was excited about mapping the White Creek Cemetery for his Eagle Project because he felt it was an enduring project that could be built upon, would be useful into the future, and could be used by many people.
Ben divided his project into two parts:
- Data Collection
- Data Entry
Data Collection:
The first step was accomplished at the White Creek Cemetery on April 28, 2012.
Ben divided a group of 33 volunteers into two-man teams, who took photos of each headstone, or photos of a rubbing of a headstone if the inscription was illegible. They marked each photo with a label that identified where in the cemetery the headstone is located. The teams also used a data collection form to record information including: name of deceased, birth date, death date, and all other information found on the headstone.
After the data collection was complete, Ben recruited 12 people to work on computers and enter the data on an online map he created on the Names in Stone website. During one data entry session, scout leaders were able to provide enough laptops to have four groups enter data simultaneously.
Ben learned quickly how important it was to standardize their methods--and he put himself in charge of quality control. That's one bit of advice he would give to any other Scout who wants to map a cemetery: include "Quality Control" as one of the steps of the project. "I would recommend to anyone who wishes to complete a similar project to use a standard format for all data both in recording and in data upload--as well as standard equipment on the collection end."
When Ben was finished, he had created an online digital cemetery with 524 graves! Users from anywhere in the world are able to click on any grave on Ben's map and see a photo of the headstone and all the data linked with that grave.
How do Ben and his family feel about this project now that it's complete?
Ben says, "Doing this project felt like…ROOTS. I was connecting back to my past. When I was entering names in the computer, it hit me—how these people lived and how they were connected to ME."
Ben's mother, Deborah, helped on the project by entering headstone information on the computer. She says, "This project was near and dear to our hearts, because my husband's family is buried there. It was a profound spiritual experience for me. It was almost like I felt them say to me, 'Thank you. Now I can be found, and I am not forgotten.'"
Ben gives special thanks to all the volunteers who donated their time and efforts to make his cemetery mapping project a big success!
Click here to view the White Creek Cemetery map on Names In Stone.
Would you like to learn more about creating a digital map of a cemetery? Click here for detailed instructions. Get more information about Names In Stone and Eagle Scout Service Projects here.