The comparative osteology and phylogenetic relationships of African and South American lungfishes (Sarcopterygii: Dipnoi)

Lepidosirenidae is a clade of freshwater lungfishes that include the extant South American Lepidosiren paradoxa Fitzinger, 1837 and African species of the genus Protopterus. These genera have been geographically separated since the break‐up of Gondwana in the Early Cretaceous, but they display similar biology and morphology. Species were distinguished by a combination of features such as head‐to‐body ratios, the number of pairs of ribs, and the presence of external gills, but no discrete skeletal characters were identified, and no comparative studies including all extant species have been published. I used computed tomography (CT), X‐ray photography, and specimens from museum collections to describe the skeletal morphology of all species of lepidosirenid in a comparative context. I digitally disarticulated the bones in each specimen to compile a comparative atlas of the cranial and pectoral elements of all extant lungfishes, which has the potential to increase the correct identifications of specimens in museum collections. The morphology of the frontoparietal, parasphenoid, supraorbital, and suboperculum differ between species. I used those characters, along with molecular sequence data from the ribosomal RNA gene 16S, to run combined morphological and molecular phylogenetic analyses. Lepidosirenidae is monophyletic in all analyses, but the interrelationships of the species of Protopterus vary with the different sources of character data. © 2015 The Linnean Society of London     

Lymphatic vessels and cancer

Survival and development of tumors depends on nutritional and respiratory biological events and exchanges ensured by blood and lymph. Tumor proliferation is associated with an increase in the vascular networks either near the tumor or intra-tumorally. Tumor tissues are able to increase their provisionment according to their needs while directing and optimizing the development of peri-tumoral vessels. The production of growth factors stimulating neo-formation of lymphatic vessels by cancer cells constitutes one of the adaptations responsible for metastatic propagation. During tumor development the lymphatic system is considered in many cases of cancer as the primary means of metastasis dissemination. The study of the lymphatic system setting and ways to block it are important points to consider in oncology.     

A new assemblage of Cenozoic lungfishes (Dipnoi: Lepidosirenidae) from the late Oligocene Nsungwe Formation,Rukwa Rift Basin,southwestern Tanzania

Lungfish (Dipnoi) date back to the Devonian, and some fossil taxa as well as extant African lungfishes are known for their ability to aestivate, tolerating low-oxygen environments associated with seasonal drying. Extant lungfishes are separated into two families: Lepidosirenidae (Protopterus in Africa and Lepidosiren in South America) and Neoceratodontidae (Neocerotadus in Australia). African lungfishes were more geographically and phylogenetically diverse on the continent in the past than they are today, with only 5% of extinct taxa recorded from the sub-Saharan fossil record. Given the sparse record of Lepidosirenidae fossils from continental Africa, any new materials are important for understanding diversification of the clade. Here we describe new lungfish fossils cautiously referable to Protopterus annectens and Protopterus aethiopicus, which are strongly supported sister taxa based on the molecular phylogeny. Specimens were collected from the late Oligocene Nsungwe Formation in the Rukwa Rift Basin (RRB) of southwestern Tanzania. The late Oligocene Nsungwe Formation represents a sequence of continental rift-fill deposits of the Songwe sub-basin of the RRB and is subdivided into the lower Utengule and upper Songwe members. Recovery of such material from the Paleogene of Africa below the equator addresses a sizable gap in the lungfish fossil record. It also expands the Nsungwe Formation fauna that includes invertebrates, alestid fishes, ptychadenid anurans, snakes, and several clades of mammals, deepening paleoecological insights into the late Oligocene record of the continental African interior. At present, P. aethiopicus and P. dolloi have an extensive modern eastern African distribution associated with the rift lakes and a region where extant members of P. annectens are not presently known. Fossil specimens described herein document presence of the clade during Paleogene volcanic activity in the western branch of the Eastern African Rift System.     

The interrelationships of Devonian lungfishes (Sarcopterygii: Dipnoi) as inferred from neurocranial evidence and new data from the genus Soederberghia Lehman, 1959

New data are presented on the neurocranial complex (endocranium plus intimately associated bones of the palate) of the Late Devonian (Famennian) ‘rhynchodipterid’ lungfish Soederberghia groenlandica from the Aina Dal Formation (Celsius Bjerg Group) of East Greenland. Only the otic and occipital regions of the braincase are ossified in Soederberghia. The neurocranium of this genus shares a series of derived features with ‘Griphognathus’whitei, including a cranial centrum, fenestrate lateral cristae, and gutters on the ventral surface of the parasphenoid interpreted as accommodating the lateral dorsal aortae. The interrelationships of early lungfishes have been the subject of considerable disagreement. New data from Soederberghia are coupled with a data set focused on the character‐rich neurocranial complex in order to examine the systematic utility of this underexploited morphological system. Different methods of phylogenetic inference (maximum parsimony, Bayesian) return broadly consistent results. The Early to Middle Devonian forms Dipnorhynchus, Stomiahykus and Uranolophus are placed among the most basal of lungfishes. ‘Holodontids’ plus ‘rhynchodipterids’ (comprising Griphognathus and Soederberghia) occupy an apical position, and are separated from the earliest lungfishes by a paraphyletic assemblage of taxa generally identified as ‘chirodipterids’ and ‘dipterids.’ This finds broad agreement with the results of previous cladistic studies focused on non‐neurocranial data sets, but diverges from functional‐adaptive scenarios that posit three lineages of early lungfishes based on aspects of the dentition. As currently defined, both ‘chirodipterids’ and the genus Chirodipterus are heterogeneous assemblages; this analysis fails to find support for the monophyly of either. While this study indicates that Griphognathus is probably paraphyletic, it nevertheless supports a close relationship between the nominal species of this genus and Soederberghia. © 2007 The Linnean Society of London, Zoological Journal of the Linnean Society, 2007, 151 , 115–171.     

Lymphatic vessels in bone support regeneration after injury

1. Download : Download high-res image (216KB)
2. Download : Download full-size image

     

Lymphatic vessels in the capsule of the lymph node

By means of the injection method the lymphatic vessels, running to the lymph nodes of various localization, have been studied. Their architectonics in the lymph node capsule is revealed. In the capsule the afferent vessels make peculiar broom-like formations. They are named terminal arborizations of afferent lymphatic vessels (TAALV). Two types of such arborizations are described: palm-like, peculiar for the somatic type of the lymph nodes, and tree-like, specific for visceral nodes. The TAALV diameter is 15-20 mcm. They come across the nodal capsule, penetrating it obliquely with numerous holes. In the TAALV wall myocytes are revealed. Together with the capsule muscular elements they might play a role of the most important factors in the mechanism of lymph circulation.     

Das geruchsorgan der rezenten Actinistia und Dipnoi (Pisces)

The olfactory organs of all recent genera from the Actinistia (Latimeria) and Dipnoi (Neoceratodus, Lepidosiren, Protopterus) were studied morphologically, and compared with each other. Whereas the olfactory organ of Latimeria resembles that of the Brachiopterygii, the olfactory organ of the Dipnoi is similar to that, of the Elaemohra.nnhii and ArtinnnfarAraii

---

---

Nit dankenswerter Unterstützung durch die Deutsche Forschungsgemeinschaft.     

Regionalization of axial skeleton in the lungfish Neoceratodus forsteri (Dipnoi)

Differentiation of the axial skeleton into distinct regions, once thought to be characteristic of the Tetrapoda, also occurs in the actinopterygian Danio rerio. In these taxa, the boundary between the cervical-thoracic regions correlates with Hoxc6 expression and morphological features such as position of the pectoral fin and associated nerves, and the absence of ribs. In the lungfish Neoceratodus, a member of the extant sister taxon to the Tetrapoda, the first vertebral element to chondrify is situated well posterior to the skull, developing from somites 6 and 7 (6/7) and associated with an enlarged cranial rib and nerves innervating the pectoral fin. Two vertebral elements develop later and more anteriorly, associated with somites 4/5 and 5/6. These three elements become incorporated into the occipital region of the skull during Neoceratodus ontogeny, until the cranial rib itself articulates to the rear of the skull. These features of early development indicate a regionalization of the Neoceratodus vertebral column: the cranial rib marks the boundary between the cervical and thoracic regions, the two more anterior vertebrae lacking ribs represent the cervical region, while somites 1-4 (cranial half), lacking any vertebral development, represent the occipital region. However, the cervical region of the vertebral column is effectively lost during ontogeny of Neoceratodus. A recognizable cervical region in the tetrapod vertebral column, as in zebrafish, suggests that cervical vertebrae are not incorporated into the skull but maintained as distinct elements of the column, representing an important shift in relative developmental timing and the influence of heterochrony in this region during the fish-tetrapod transition.     

Lymphatic vessels in the developing diaphragm of the rat.

Diaphragms of fetal, neonatal and young albino rats have been observed both under light and electron microscopes to examine the presence and distribution of lymphatic vessels and their morphological features. In fetal diaphragms of between 18 and 22 days of gestation, no normal lymphatic vessels can be seen; only after birth, specifically in neonatal and 2-day-old rats, small lymphatic vessels appear; they are in close proximity to the blood vessels in the inner areas of the muscle. As the rats get older, lymphatic vessels are also observed in the subserosa where an abundant connective tissue is present. The fine structure of diaphragmatic lymphatic vessels is different at different ages. In neonatal rats of up to 2 days, the endothelial wall is very thin and often holed. The relationships between contiguous endothelial cells are characterized by simple end-to-end or overlapping structures. The basement membrane is virtually absent. Within the first week of life, the endothelial wall becomes more complex; along the wall, complex interdigitations between two contiguous endothelial cells often touch. A discontinuous basement membrane and collagen and elastic fibers surround the vessels. In the older rats (from 14 to 25 to 140 days), next to the complex interdigitations which characterize the junction between two contiguous endothelial cells, cellular flaps interdigitate forming a channel which opens out either to the exterior or the interior of the vessel. Dense bundles of elastic and collagen fibers are closely apposed to the endothelial wall.     

Some remarks on the axial skeleton in Protopterus (Pisces, Dipnoi)

Based on a comparative morphological study of the axial skeleton in a broader range of "primitive bony fishes" a reevaluation of some characteristic traits of the vertebral column in the African lungfish has been done. The axial skeleton of living dipnoans has to be regarded as reduced from a more complex phylogenetic state and may be neotenic in some respect. The formation of perichordal bony autocentra could be confirmed, at least in one specimen. The well-known invasion of mesenchymal cells into the fibrous sheet of the notochord is interpreted as a rudimentary chordacentric "vertebralization". Also included is an analysis of the peculiar urostyle of lungfishes, its regeneration capacity and its bearing upon the principles of vertebra-formation.     

Prismatic dentine in the Australian lungfish, Neoceratodus forsteri (Osteichthyes: Dipnoi)

The Australian lungfish, Neoceratodus forsteri, has a dentition consisting of enamel, mantle dentine and bone, enclosing circumdenteonal, core and interdenteonal dentines. Branching processes from cells that produce interdenteonal dentine leave the cell surface at different angles, with collagen fibrils aligned parallel to the long axis of each process. In the interdenteonal dentine, crystals of calcium hydroxyapatite form within fibrils of collagen, and grow within a matrix of non-collagenous protein. Crystals are aligned parallel to the cell process, as are the original collagen fibrils. Because the processes are angled to the cell surface, the crystals within the core or interdenteonal dentine are arranged in bundles set at angles to each other. Apatite crystals in circumdenteonal dentine are finer and denser than those of the interdenteonal dentine, and form outside the fibrils of collagen. In mature circumdenteonal dentine the crystals of circumdenteonal dentine form a dense tangled mass, linked to interdenteonal dentine by isolated crystals. The functional lungfish tooth plate contains prisms of large apatite crystals in the interdenteonal dentine and masses of fine tangled crystals around each denteon. This confers mechanical strength on a structure with little enamel that is subjected to heavy wear.     

Scale structure in the Australian lungfish,Neoceratodus forsteri (Osteichthyes: Dipnoi)

Scales of the Australian lungfish, Neoceratodus forsteri, are secreted within the dermis by a capsule of scleroblasts, and enclosed in a pouch made of collagen fibers, in contact with the epidermis over the posterior third of the scale. Each scale grows from a focus, which represents the first formed part of the scale. On the internal surface of the scale is elasmodin, made of collagen fiber bundles arranged in layers. Elasmodin, unmineralized in N. forsteri, contains cells in the living animal, and the number of layers increases as the scales grow. Squamulin, on the thin external part of the scale, is also laid down in layers, and based on a matrix of fine collagen fibrils, mineralized with a poorly crystalline biogenic calcium hydroxylapatite. Squamulin is divided into separate sections called squamulae, and contains long tubules with cells applied to the wall of the tubule. The anterior and lateral surfaces of the squamulin are ornamented with pediculae, and the posterior surface has longitudinal ridges, from which collagen fibers extend to anchor the scale within the pouch. Elasmodin and squamulin are linked by unmineralized collagen fibrils. The layers, formed at irregular intervals, are connected around the margin of the scale, effectively converting the whole scale into a flat structure resembling a pearl, with the first formed tissues deeply embedded inside the scale, and the youngest on the outer surface. Incremental lines in the hard tissue, and the number of layers in the elasmodin, do not reflect the chronological age of the fish. J. Morphol. 276:1137–1145, 2015. © 2015 Wiley Periodicals, Inc.     

The First Virtual Cranial Endocast of a Lungfish (Sarcopterygii: Dipnoi)

Lungfish, or dipnoans, have a history spanning over 400 million years and are the closest living sister taxon to the tetrapods. Most Devonian lungfish had heavily ossified endoskeletons, whereas most Mesozoic and Cenozoic lungfish had largely cartilaginous endoskeletons and are usually known only from isolated tooth plates or disarticulated bone fragments. There is thus a substantial temporal and evolutionary gap in our understanding of lungfish endoskeletal morphology, between the diverse and highly variable Devonian forms on the one hand and the three extant genera on the other. Here we present a virtual cranial endocast of Rhinodipterus kimberleyensis, from the Late Devonian Gogo Formation of Australia, one of the most derived fossil dipnoans with a well-ossified braincase. This endocast, generated from a Computed Microtomography (µCT) scan of the skull, is the first virtual endocast of any lungfish published, and only the third fossil dipnoan endocast to be illustrated in its entirety. Key features include long olfactory canals, a telencephalic cavity with a moderate degree of ventral expansion, large suparaotic cavities, and moderately enlarged utricular recesses. It has numerous similarities to the endocasts of Chirodipterus wildungensis and Griphognathus whitei, and to a lesser degree to ''Chirodipterus'' australis and Dipnorhynchus sussmilchi. Among extant lungfish, it consistently resembles Neoceratodus more closely than Lepidosiren and Protopterus. Several trends in the evolution of the brains and labyrinth regions in dipnoans, such as the expansions of the utricular recess and telencephalic regions over time, are identified and discussed.