Evolution and development of the synarcual in early vertebrates

The synarcual is a structure incorporating the anterior vertebrae of the axial skeleton and occurs in vertebrate taxa such as the fossil group Placodermi and the Chondrichthyes (Holocephali, Batoidea). Although the synarcual varies morphologically in these groups, it represents the first indication, phylogenetically, of a differentiation of the vertebral column into separate regions. Among the placoderms, the synarcual of Cowralepis mclachlani Ritchie, 2005 (Arthrodira) shows substantial changes during ontogeny to produce an elongate, spool-shaped structure with a well-developed dorsal keel. Because the placoderm synarcual is covered in perichondral bone, the ontogenetic history of this Cowralepis specimen is preserved as it developed anteroposteriorly, dorsally and ventrally. As well, in the placoderm Materpiscis attenboroughi Long et al., 2008 (Ptyctodontida), incomplete fusion at the posterior synarcual margin indicates that both neural and haemal arch vertebral elements are added to the synarcual. A survey of placoderm synarcuals shows that taxa such as Materpiscis and Cowralepis are particularly informative because perichondral ossification occurs prior to synarcual fusion such that individual vertebral elements can be identified. In other placoderm synarcuals (e.g. Nefudina qalibahensis Lelièvre et al., 1995; Rhenanida), cartilaginous vertebral elements fuse prior to perichondral ossification so that individual elements are more difficult to recognize. This ontogenetic development in placoderms can be compared to synarcual development in Recent chondrichthyans; the incorporation of neural and haemal elements is more similar to the holocephalans, but differs from the batoid chondrichthyans.     

Ontogenetic variation in the stratum granulosum of the epidermis of Chaetophractus vellerosus (Xenarthra,Dasypodidae) in relation to the development of cornified scales

The epidermis of mammals is characterized by having a stratum granulosum that produces an orthokeratotic stratum corneum, different from the typical reptilian parakeratotic stratum. Nonetheless, some mammals show distinct degrees of parakeratosis in epidermal regions with few or no pilose follicles (e.g., areas subjacent to cornified scales). With respect to the epidermis and the development of cornified scales in the Dasypodidae, previous studies have supported the presence of a continuous stratum granulosum without any variations during ontogeny. This condition, in which the cornified scales develop without a loss of the stratum granulosum, was interpreted as primitive for eutherians. The present contribution expands the knowledge on the epidermis of Chaetophractus vellerosus in distinct ontogenetic stages in order to determine whether the cornified scales show the same developmental pattern as in other eutherians. The presence of a stratum granulosum in C. vellerosus neonates and its reduction in more advanced ontogenetic stages, in direct relationship with cornified scale development, supports the hypothesis that the partial parakeratosis in the xenarthran integument is secondary, as in other eutherians, and can be interpreted as a derived character state.     

A new elasmothere (Perissodactyla, Rhinocerotidae) from the late Miocene of the Linxia Basin in Gansu, China

Ningxiatherium euryrhinus sp. nov. is a relatively large elasmothere rhino from the Linxia Basin in northwestern China found in the early Bahean, which corresponds to the early Vallesian (MN 9), ca. 11.1 Ma. It is much larger than the extant Ceratotherium simum and has a single horn. Ningxiatherium is similar to the late Miocene Parelasmotherium from Gansu and Shanxi, but differs in having partially ossified nasal septum, terminal nasal horn boss, shallower nasal notch above the P3/P4 boundary, and much shorter premolars. N. euryrhinus sp. nov. differs from N. longirhinus from the late Miocene of Zhongning, Ningxia in northwestern China by its larger size, relatively wider nasals, shallower nasal notch, and sub-quadrangular occlusal surface of M3. In the Linxia Basin, the other early late Miocene (Bahean or Vallesian) elasmotheres are known from complete skulls or isolated teeth. They include two genera and three species: Parelasmotherium linxiaense, P. simplum, and Iranotherium morgani. They lack a nasal septum, but Ningxiatherium has one. The age of N. euryrhinus is early late Miocene based on direct association with biochronologic indicator taxa, such as Dinocrocuta gigantea, Hipparion dongxiangense, and Chilotherium primigenius. N. euryrhinus is more primitive than N. longirhinus sp. nov. in having a sub-quadrangular M3, a shallower nasal notch, and the presence of DP1 in adults.     

Males Resemble Females: Re-Evaluating Sexual Dimorphism in Protoceratops andrewsi (Neoceratopsia,Protoceratopsidae)

BackgroundProtoceratops andrewsi (Neoceratopsia, Protoceratopsidae) is a well-known dinosaur from the Upper Cretaceous of Mongolia. Some previous workers hypothesized sexual dimorphism in the cranial shape of this taxon, using qualitative and quantitative observations. In particular, width and height of the frill as well as the development of a nasal horn have been hypothesized as potentially sexually dimorphic.Conclusions/SignificanceSexual dimorphism within Protoceratops andrewsi is not strongly supported by our results, as previously proposed by several authors. Anatomical traits such as height and width of the frill, and skull size thus may not be sexually dimorphic. Based on PCA for a data set focusing on the rostrum and associated ANOVA results, nasal horn height is the only feature with potential dimorphism. As a whole, most purported dimorphic variation is probably primarily the result of ontogenetic cranial shape changes as well as intraspecific cranial variation independent of sex.     

A nearly complete skeleton of an early juvenile diplodocid (Dinosauria: Sauropoda) from the Lower Morrison Formation (Late Jurassic) of north central Wyoming and its implications for early ontogeny and pneumaticity in sauropods

A nearly complete skeleton of a juvenile sauropod from the Lower Morrison Formation (Late Jurassic, Kimmeridgian) of the Howe Ranch in Bighorn County, Wyoming is described. The specimen consists of articulated mid-cervical to mid-caudal vertebrae and most appendicular bones, but cranial and mandibular elements are missing. The shoulder height is approximately 67 cm, and the total body length is estimated to be less than 200 cm. Besides the body size, the following morphological features indicate that this specimen is an early juvenile; (1) unfused centra and neural arches in presacral, sacral and first to ninth caudal vertebrae, (2) unfused coracoid and scapula, (3) open coracoid foramen, and (4) relatively smooth articular surfaces on the limb, wrist, and ankle bones. A large scapula, short neck and tail and elongate forelimb bones relative to overall body size demonstrate relative growth. A thin-section of the mid-shaft of a femur shows a lack of annual growth lines, indicating an early juvenile individual possibly younger than a few years old. Pneumatic structures in the vertebral column of the specimen SMA 0009 show that pneumatisation of the postcranial skeleton had already started in this individual, giving new insights in the early ontogenetic development of vertebral pneumaticity in sauropods.

The specimen exhibits a number of diplodocid features (e.g., very elongate slender scapular blade with a gradually dorsoventrally expanded distal end, a total of nine dorsal vertebrae, presence of the posterior centroparapophyseal lamina in the posterior dorsal vertebrae). Although a few diplodocid taxa, Diplodocus, cf. Apatosaurus, and cf. Barosaurus, are known from several fossil sites near the Howe Ranch, identification of this specimen, even at a generic level, is difficult due to a large degree of ontogenetic variation.     

Major cranial changes during Triceratops ontogeny

This is the first cranial ontogenetic assessment of Triceratops, the well-known Late Cretaceous dinosaur distinguished by three horns and a massive parietal-squamosal frill. Our analysis is based on a growth series of 10 skulls, ranging from a 38 cm long baby skull to about 2 m long adult skulls. Four growth stages correspond to a suite of ontogenetic characters expressed in the postorbital horns, frill, nasal, epinasal horn and epoccipitals. Postorbital horns are straight stubs in early ontogeny, curve posteriorly in juveniles, straighten in subadults and recurve anteriorly in adults. The posterior margin of the baby frill is deeply scalloped. In early juveniles, the frill margin becomes ornamented by 17-19 delta-shaped epoccipitals. Epoccipitals are dorsoventrally compressed in subadults, strongly compressed and elongated in adults and ultimately merge onto the posterior frill margin in older adults. Ontogenetic trends within and between growth stages include: posterior frill margin transitions from scalloped to wavy and smooth; progressive exclusion of the supraoccipital from the foramen magnum; internal hollowing at the base of the postorbital horns; closure of the midline nasal suture; fusion of the epinasal onto the nasals; and epinasal expansion into a morphologically variable nasal horn. We hypothesize that the changes in horn orientation and epoccipital shape function to allow visual identity of juveniles, and signal their attainment of sexual maturity.     

Hot receptors in the brain

Background

The complex neuronal circuitry of the dorsal horn of the spinal cord is as yet poorly understood. However, defining the circuits underlying the transmission of information from primary afferents to higher levels is critical to our understanding of sensory processing. In this study, we have examined phosphodiesterase 1C (Pde1c) BAC transgenic mice in which a green fluorescent protein (GFP) reporter gene reflects Pde1c expression in sensory neuron subpopulations in the dorsal root ganglia and spinal cord.

Results

Using double labeling immunofluorescence, we demonstrate GFP expression in specific subpopulations of primary sensory neurons and a distinct neuronal expression pattern within the spinal cord dorsal horn. In the dorsal root ganglia, their distribution is restricted to those subpopulations of primary sensory neurons that give rise to unmyelinated C fibers (neurofilament 200 negative). A small proportion of both non-peptidergic (IB4-binding) and peptidergic (CGRP immunoreactive) subclasses expressed GFP. However, GFP expression was more common in the non-peptidergic than the peptidergic subclass. GFP was also expressed in a subpopulation of the primary sensory neurons immunoreactive for the vanilloid receptor TRPV1 and the ATP-gated ion channel P2X₃. In the spinal cord dorsal horn, GFP positive neurons were largely restricted to lamina I and to a lesser extent lamina II, but surprisingly did not coexpress markers for key neuronal populations present in the superficial dorsal horn.

Conclusion

The expression of GFP in subclasses of nociceptors and also in dorsal horn regions densely innervated by nociceptors suggests that Pde1c marks a unique subpopulation of nociceptive sensory neurons.     

Three-Dimensional Distribution of Sensory Stimulation-Evoked Neuronal Activity of Spinal Dorsal Horn Neurons Analyzed by In Vivo Calcium Imaging

The spinal dorsal horn comprises heterogeneous populations of interneurons and projection neurons, which form neuronal circuits crucial for processing of primary sensory information. Although electrophysiological analyses have uncovered sensory stimulation-evoked neuronal activity of various spinal dorsal horn neurons, monitoring these activities from large ensembles of neurons is needed to obtain a comprehensive view of the spinal dorsal horn circuitry. In the present study, we established in vivo calcium imaging of multiple spinal dorsal horn neurons by using a two-photon microscope and extracted three-dimensional neuronal activity maps of these neurons in response to cutaneous sensory stimulation. For calcium imaging, a fluorescence resonance energy transfer (FRET)-based calcium indicator protein, Yellow Cameleon, which is insensitive to motion artifacts of living animals was introduced into spinal dorsal horn neurons by in utero electroporation. In vivo calcium imaging following pinch, brush, and heat stimulation suggests that laminar distribution of sensory stimulation-evoked neuronal activity in the spinal dorsal horn largely corresponds to that of primary afferent inputs. In addition, cutaneous pinch stimulation elicited activities of neurons in the spinal cord at least until 2 spinal segments away from the central projection field of primary sensory neurons responsible for the stimulated skin point. These results provide a clue to understand neuronal processing of sensory information in the spinal dorsal horn.     

中国第一个完整的锯齿龙头骨

中国已经命名了7种锯齿龙,但是没有一个有完整的头骨。本文描述一个产自鄂尔多斯盆地孙家沟组的新材料,包括一个近于完整头骨及关连的下颌。根据下列特征将其归入埃尔金龙科(Elginiidae):头骨的膜质骨突起形成长而尖的角,面颊上装饰有显著的锥型角,上颞骨角从头骨一角向侧后方伸出,副基蝶骨细长且腹面中间有凹沟,案骨加大并在中间相互接触,使后顶骨不参与形成头骨边缘。该标本与矮小三川龙(Sanchuansaurus pygmaeus)有下列共同特征:鼻孔后的上颌骨前背支有小的突起,上颌齿齿冠扇形,9—11个齿尖;但是有以下特征区别于后者:眶下神经两个外开口距离更近且更接近上颌骨腹缘,齿冠不重叠,只有14个牙槽。与柳林黄河龙(Huanghesaurus)共有以下特征:下颌腹缘直、光滑,隅骨突位于后缘,下颌齿有中脊,17个齿尖;不过其下颌前部厚度比后部大。新标本表明三川龙和黄河龙关系接近,两者都应可能属于埃尔金龙科。根据新标本建立一个新种并暂时归入石千峰龙属,即完整石千峰龙(Shitienfenia completus);但是目前不能排除这个标本属于二叠石千峰龙的可能性。     

Anatomy and taxonomic status of the chasmosaurine ceratopsid Nedoceratops hatcheri from the upper Cretaceous Lance Formation of Wyoming, U.S.A

Background

The validity of Nedoceratops hatcheri, a chasmosaurine ceratopsid dinosaur known from a single skull recovered in the Lance Formation of eastern Wyoming, U.S.A., has been debated for over a century. Some have argued that the taxon is an aberrant Triceratops, and most recently it was proposed that N. hatcheri represents an intermediate ontogenetic stage between “young adult” and “old adult” forms of a single taxon previously split into Triceratops and Torosaurus.

Methodology/Principal Findings

The holotype skull of Nedoceratops hatcheri was reexamined in order to map reconstructed areas and compare the specimen with other ceratopsids. Although squamosal fenestrae are almost certainly not of taxonomic significance, some other features are unique to N. hatcheri. These include a nasal lacking a recognizable horn, nearly vertical postorbital horncores, and relatively small parietal fenestrae. Thus, N. hatcheri is tentatively considered valid, and closely related to Triceratops spp. The holotype of N. hatcheri probably represents an “old adult,” based upon bone surface texture and the shape of the horns and epiossifications on the frill. In this study, Torosaurus is maintained as a genus distinct from Triceratops and Nedoceratops. Synonymy of the three genera as ontogenetic stages of a single taxon would require cranial changes otherwise unknown in ceratopsids, including additions of ossifications to the frill and repeated alternation of bone surface texture between juvenile and adult morphotypes.

Conclusions/Significance

Triceratops, Torosaurus, and likely Nedoceratops, are all distinct taxa, indicating that species richness for chasmosaurine ceratopsids in the Lance Formation just prior to the Cretaceous-Paleocene extinction was roughly equivalent to that earlier in the Cretaceous.     

黑犀(奇蹄目,犀科)化石在中国的首次发现

黑犀(Diceros属)的惟一现生代表D.bicornis生活在非洲。该属在新近纪时期曾广泛分布于希腊、土耳其和伊朗等地区,但从未在东亚地区发现过。新种甘肃黑犀(Diceros gan- suensis sp.nov.)是该属在中国和东亚的首次发现。化石采自甘肃临夏盆地晚中新世柳树组中部。新种以尺寸较小、头型短、枕顶高耸、枕面窄而高、枕嵴无中沟、副枕突短小、下颌上升支距m3较近、前臼齿较小、DP1无后脊、P2原脊孤立、P2和P3后脊细窄而区别于东地中海地区的Diceros neumayri。D.neumayri的分类位置一直是一个争论的焦点,曾在黑犀(Diceros属)和白犀(Ceratotherium属)之间反复变更。研究显示,甘肃黑犀和D.neumayri的一系列共同的原始特征表明它们与更进步的白犀有明显的区别,应该归入黑犀属。     

Palatal thickening and facial form in Paranthropus: Examination of alternative developmental models

Paranthropus is distinctive among hominoids in its possession of a greatly thickened hard palate. Although traditionally considered a structural adaptation to counter high-magnitude masticatory stress, alternative developmental models are equally viable. Three models of palatal thickening were evaluated in this study. A mechanical model interprets palatal thickening as a compensatory response to increased instability of the midpalatal suture effected by an anterior placement of the masseteric muscle mass. This model predicts that palatal thickness is correlated with the length of the palate posterior to the masseteric tubercle. Two non-mechanical models consider the thickness of the hard palate to be structurally related to and therefore correlated with either 1) the degree to which the premaxilla overlaps the hard palate in the subnasal region or 2) the height of the posterior facial skeleton. The correlation of craniofacial variables was assessed intraspecifically in ontogenetic series of great ape and human crania. Tests of correlation were performed for each comparison using both residuals calculated from reduced major axis regression of the variable of interest against a measure of cranial size and shape ratios. A significant correlation of palatal thickness with posterior facial height in Pan suggests that the unusually thick hard palate of Paranthropus is directly related to the increased posterior facial height characteristic of this taxon. Further evaluation suggests that extreme palatal thickening in these specimens occurred by virtue of their possession of a nasal septum morphology in which the vomer extends onto the superior nasal surface of the premaxilla. Such a morphology would have constrained the palatal nasal lamina to maintain the approximate level of the premaxillary nasal lamina throughout the growth process thereby promoting palatal thickening. Am J Phys Anthropol 103:375–392, 1997. © 1997 Wiley-Liss, Inc.     

A new Late Miocene ovibovine-like bovid (Bovidae,Mammalia) from the Kassandra Peninsula (Chalkidiki,Northern Greece) and implications to the phylogeography of the group

A new Late Miocene bovid, Urmiatherium kassandriensis sp. nov., from Northern Greece is described. The material comes from the Fourka locality in the Kassandra Peninsula (Chalkidiki), and the included fauna is estimated to be of Vallesian age. The two preserved crania represent a medium-sized taxon with short, conical horn cores, a flat cranial roof (consisting of the posterior part of the frontals, parietal and occipital), thick and porous frontals and pneumatized short parietals, an extremely thick basioccipital with voluminous posterior tuberosities and accessory articular facets for the atlas. The specialized atlanto-occipital joint recalls Pleistocene and extant ovibovines, but the braincase structure as a whole and the horn core features closely match Late Miocene ovibovine-like taxa, especially Plesiaddax and even more Urmiatherium. Nevertheless, the Kassandra bovid differs from representatives of both genera in the simpler horn core morphology and external brain anatomy. Urmiatherium is known to appear first in China and Iran at about 7.8 Ma, whereas its westernmost appearance on Samos Island (Greece) is dated much later. The presence of Urmiatherium kassandriensis sp. nov. in N. Greece suggests a farther west and earlier (Vallesian at least) first appearance of the genus. This would justify a basic geographic and phylogenetic split of Urmiatherium into two main Turolian lineages: a central-eastern Asian one leading to the sister species U. polaki and U. intermedium and a western one leading to U. rugosifrons.     

Morphological Variations within the Ontogeny of Deinonychus antirrhopus (Theropoda,Dromaeosauridae)

This research resulted from the determination that MCZ 8791 is a specimen of Deinonychus antirrhopus between one and two years of age and that the morphological variations within particular growth stages of this taxon have yet to be described. The primary goal of the research is to identify ontogenetic variations in this taxon. Histological analyses determined that the Deinonychus specimens AMNH 3015 and MOR 1178 were adults. Comparisons are made between MCZ 8791 and these adult specimens. The holotype, YPM 5205, and the other associated specimens of this taxon within the YPM collection are similar in size and morphology to AMNH 3015. Further comparisons were made with the three partial specimens OMNH 50268, MCZ 4371, and MOR 1182. Although these specimens represent only a partial ontogenetic series, a number of morphological variations can be described. One secondary goal of this research is to compare the known pattern of variable, informative, ontogenetic characters in MCZ 8791 to a similar pattern of morphological characters in the sub-adult dromaeosaurid specimen Bambiraptor feinbergorum, AMNH FR: 30556. If the characters that have been determined to represent variable juvenile morphology in the ontogeny of Deinonychus are exhibited in Bambiraptor, this study will begin the process of determining whether a similar, conservative, ontogenetic pattern exists throughout the rest of Dromaeosauridae. If defensible, it may reduce the number of sympatric taxa within this clade. The other secondary goal relates to the forelimb function. The approximate body size, forelimb length, wrist development, and the presence of a more prominent olecranon on the ulna of MCZ 8791 support the hypothesis that juveniles of this taxon possessed some form of flight capability.     

Three Echinostome Species from Wild Birds in the Republic of Korea

Three echinostome species, i.e., Patagifer bilobus, Petasiger neocomense, and Saakotrema metatestis, are newly recorded in the trematode fauna of the Republic of Korea. They were recovered from 3 species of migratory birds (Platalea minor, Podiceps cristatus, and Egretta garzetta), which were donated by the Wildlife Center of Chungbuk (WCC) and the Conservation Genome Resource Bank for Korean Wildlife (CGRB). Only 1 P. bilobus specimen was recovered from the intestine of a black-faced spoonbill (P. minor), and characterized by the bilobed head crown with a deep dorsal incision and 54 collar spines. Twenty P. neocomense were recovered from the intestine of a great crested grebe (P. cristatus), and they had a well-developed head crown with 19 spines and 2 testes obliquely located at the posterior middle of the body. Total 70 S. metatestis were collected from the bursa of Fabricius of 1 little egret (E. garzetta). It is characterized by stout tegumental spines covered in the entire leaf-shaped body, posterior extension of the uterus, presence of the uroproct and a well-developed head crown with 12 pairs of collar spines on each side. By the present study, these 3 echinostome species are newly added to the trematode fauna in Korea.     

Shape Variation in the Dermatocranium of the Greater Short-Horned Lizard <Emphasis Type="Italic">Phrynosoma hernandesi</Emphasis> (Reptilia: Squamata: Phrynosomatidae)

Dermatocranial shape and horn morphology display great disparity among the species of Phrynosoma. Ontogenetic change in dermatocranial shape in a series of 79 specimens of the short-horned Phrynosoma hernandesi (54F: 25M) was examined using geometric morphometric techniques. A multivariate ANCOVA of Procrustes residuals with sex as a factor and ln(centroid size) as the covariate indicated sexual shape dimorphism. Separate multivariate regressions of Procrustes residuals on ln(centroid size) for each sex indicated that allometry accounts for ~52–54% of the total sample shape variance. Comparisons of ontogenetic shape change between sexes indicate that sexual shape dimorphism is minimal and of uncertain biological significance. Groupings of multivariate regression coefficients by magnitude and sign suggest that allometric integration of the dermatocranium is not uniform over the dermatocranium. Principal component analysis of the landmark configurations corrected for sex and allometry yields a first principal component which describes shape variance concentrated in the posterolateral and posterior regions of the dermatocranium, and again is indicative of non-uniform shape variation over the dermatocranium. Our findings for P. hernandesi indicate that the adult shape of the dermatocranium may contribute to a passive defence against predation. We hypothesize that the complexity in dermatocranial shape demonstrated here for P. hernandesi indicates parcellation of shape variance, which may contribute to explanations of the pronounced dermatocranial disparity exhibited by the species of Phrynosoma.     

Time-related changes in the labeling pattern of motor and sensory neurons innervating the gastrocnemius muscle,as revealed by the retrograde transport of the cholera toxin B subunit

Summary Morphological changes in the motor and sensory neurons in the lumbar spinal cord and the dorsal root ganglia were investigated at different survival times following the injection of the B subunit of cholera toxin (CTB) into the medial gastrocnemius muscle. Unconjugated CTB, visualized immunohistochemically, was found to be retrogradely transported through ventral and dorsal roots to motor neurons in the anterior horn, each lamina in the posterior horn, and ganglion cells in the dorsal root ganglia at L₃–L₆. The largest numbers of labeled motor neurons and ganglion cells were observed 72 h after the injection of CTB. Thereafter, labeled ganglion cells were significantly decreased in number, whereas the amount of labeled motor neurons showed a slight reduction. Motor neurons had extensive dendritic trees filled with CTB, reaching lamina VII and even the pia mater of the lateral funiculus. Labeling was also seen in the posterior horn, but the central and medial parts of laminae II and III had the most extensively labeled varicose fibers, the origin of which was the dorsal root ganglion cells. The results indicate that CTB is taken up by nerve terminals and can serve as a sensitive retrogradely transported marker for identifying neurons that innervate a specific muscle.     

Morphology,ontogeny and molecular phylogeny of a new urostylid ciliate,Bakuella (Pseudobakuella) guangdongica n. sp. (Ciliophora,Hypotrichia) from southern China

The morphology, morphogenesis, and molecular phylogeny of Bakuella (Pseudobakuella) guangdongica n. sp., isolated from southern China, were investigated. The new species is characterized by a body length of 150–225 μm in vivo; 35–42 adoral membranelles; 3–5 buccal, two frontoterminal, 7–12 transverse and two pretransverse ventral cirri; midventral complex comprised of 10–20 pairs and two rows extending to transverse cirri; posterior part of marginal rows slightly overlapping; colorless cortical granules about 1 μm across, arranged in small groups; soil habitat. Its main ontogenetic features are: (1) in the proter, the parental adoral zone of membranelles is completely renewed by new structures; (2) in the opisthe, the oral primordium originates apokinetally, some old midventral cirri join the formation of frontoventral-transverse cirral anlagen; (3) the anlagen for marginal rows and dorsal kineties develop intrakinetally; and (4) the numerous macronuclear nodules fuse into a single mass before dividing. Phylogenetic analyses based on the SSU rDNA sequence suggest the non-monophyly of the genus Bakuella.