Morphological cranial diversity contributes to phylogeny in soft-furred sengis (Afrotheria, Macroscelidea)

Despite the well-supported Macroscelidea phylogeny proposed at the end of the 1960s, several systematic arrangements have been suggested in the last 20 years, raising doubts about the phylogeny of the Macroscelidinae; sengi inter-specific relationships are still debated to this day. The main issue of concern involves the supposed Elephantulus diphyly. To solve this persisting debate about sengi phylogeny, we examined the cranium ventral surface of 13 species using geometric morphometric techniques and neighbour-joining algorithms. This study supported the idea that the ventral side of the sengi cranium has the potential to provide important signals for reconstructing the Macroscelidea phylogeny. The phylogenetic signals seemed to differentiate between two major clades in the sengi radiation. In the first clade, the two monospecific genera (Petrodromus and Macroscelides), the two African Horn species (Elephantulus revoilii and E. rufescens), and the only North African species (E. rozeti) were clustered together. The second clade includes the remnant south-central African Elephantulus species. Our results were in agreement with both mitochondrial and nuclear data, confirmed that there is no Elephantulus monophyly and highlighted the close relationship between Petrodromus and E. rozeti. It appears that all the soft-furred sengi species are organised in two evolutionary lines: an old monophyletic clade, comprising only Elephantulus species, and a new polyphyletic clade, including P. tetradactylus, M. proboscideus, and E. rozeti. This requires a taxonomic and nomenclatural rearrangement within Macroscelidinae, where the phylogenetic position of the remnant 4 (of 12) Elephantulus species has yet to be fully defined.     

Reassessment of the cranial characters of Glossotherium and Paramylodon (Mammalia: Xenarthra: Mylodontidae)

Nomenclatural confusion has existed within the Mylodontinae for several genera, and has resulted in the supposition that Paramylodon of North America is synonymous with Glossotherium of South America. A taxonomic revision of crania for Glossotherium and Paramylodon upholds their separation as distinct genera and provides a list of diagnostic characters that have been lacking. Assessment was made using principal components analysis for suites of cranial and mandibular measurements, evaluation of ratios and measurement distribution, and by examining qualitative characters. Results show the greatest characterization for the skull comes from differences relating to cranial length versus width, whereas the mandible is predominantly distinguished by qualitative characters of the predental spout. Examination of the Pliocene species Glossotherium chapadmalense from South America shows a combination of characters indicative of each genus, but exhibits more with Glossotherium and is tentatively retained under that genus. The mix of characters indicates that G. chapadmalense is the likely ancestor to Paramylodon, although when and where the transition took place is still unclear. During the evolutionary transition, Paramylodon crania emphasized an increase in length of the palate, whereas those of Glossotherium emphasized an increase in cranial width. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 885–903.     

Mandible/basihyal relationships in red howler monkeys (Alouatta seniculus): A craniometrical approach

The morphological relationships between the cranium and basihyal of red howler monkeys (Alouatta seniculus) were analyzed based on measurements of 36 cranial and 3 basihyal dimensions and observations of the female specimen in spirits. In this study, 115 crania from 111 red howler monkeys and 4 mantled howler monkeys (Alouatta palliata) were used. The analyses from the standpoints of the correlation coefficients, principal component analysis, discriminant function analysis, age changes and sex differences were performed and the following results were obtained: (1) The cranial measurements closely related to the basihyal measurements were mandibular length, occipital breadth, facial length, ramus height, cranial base length, bigonial breadth and pr-i length. (2) Age changes for the mandibular measurements in males of red howler monkeys were remarkable, and, in particular, the development of the gonion toward posterior and lateral directions were characteristic. (3) The largest sex differences were found in the mandibular measurements of red howler monkeys among the Anthropoidea of seven genera compared. (4) The existence of a “cline” in the cranial measurements of the red howler monkey was recognized. (5) The inter-species differences in the crania between the red howler monkey and mantled howler monkey are obvious, metrically and non-metrically. Based on the results mentioned above, the morphological relationships between the mandible and basihyal in the red howler monkey are discussed.     

Primate cranial diversity

Many studies in primate and human evolution focus on aspects of cranial morphology to address issues of systematics, phylogeny, and functional anatomy. However, broad analyses of cranial diversity within Primates as an Order are notably absent. In this study, we present a 3D geometric morphometric analysis of primate cranial morphology, providing a multivariate comparison of the major patterns of cranial shape change during primate evolution and quantitative assessments of cranial diversity among different clades. We digitized a set of 18 landmarks designed to capture overall cranial shape on male and female crania representing 66 genera of living primates. The landmark data were aligned using a Generalized Procrustes Analysis and then subjected to a principal components analysis to identify the major axes of cranial variation. Cranial diversity among clades was compared using multivariate measurements of variance. The first principal component axis reflects differences in cranial flexion, orbit size and orientation, and relative neurocranial volume. In general, it separates strepsirrhines from anthropoids. The second axis reflects differences in relative cranial height and snout length and primarily describes differences among anthropoids. Eulemur, Mandrillus, Pongo, and Homo are among the extremes in cranial shape. Anthropoids, catarrhines, and haplorhines show a higher variance than prosimians or strepsirrhines. Hominoids show the highest variance in cranial shape among extant primate clades, and much of this diversity is driven by the unique cranium of Homo sapiens. Am J Phys Anthropol 142:565–578, 2010. © 2010 Wiley‐Liss, Inc.     

Cranial osteology of Picini (Aves: Piciformes: Picidae)

The Picini is composed of 26 species of woodpeckers from the old world that are grouped into seven genera. Given the absence of detailed studies on the cranial osteology of the Picini, the purpose of this study is to describe the cranial osteology of 14 species of Picini from six genera and to compare these species with each other, with other species of woodpeckers and with other bird groups. The results of the analysis of the bone structures of the skull of the Picini indicate that there are seven outstanding characteristic of bones involved in the functional mechanistics of the jaw apparatus. In the ossa cranii, these include the frontal overhang, the postorbital process, the extension of the parietal versus the frontal region and the temporal fossa. In the ossa faciei, these include the rostral expansion of the pterygoid, the ventral palatine fossa and the orbital process of the quadrate bone. For all of the structures associated with the cranial osteology of the investigated species, there is a clear distinction between the species of the genus Picus and the other species studied. Blythipicus rubiginosus, Chrysocolaptes lucidus, Reinwardtipicus validus, Gecinulus viridis and the species of the genus Dinopium exhibit additional particularities.     

Molecular systematics of the Cactaceae

Bayesian, maximum‐likelihood, and maximum‐parsimony phylogenies, constructed using nucleotide sequences from the plastid gene region trnK‐matK, are employed to investigate relationships within the Cactaceae. These phylogenies sample 666 plants representing 532 of the 1438 species recognized in the family. All four subfamilies, all nine tribes, and 69% of currently recognized genera of Cactaceae are sampled. We found strong support for three of the four currently recognized subfamilies, although relationships between subfamilies were not well defined. Major clades recovered within the largest subfamilies, Opuntioideae and Cactoideae, are reviewed; only three of the nine currently accepted tribes delimited within these subfamilies, the Cacteae, Rhipsalideae, and Opuntieae, are monophyletic, although the Opuntieae were recovered in only the Bayesian and maximum‐likelihood analyses, not in the maximum‐parsimony analysis, and more data are needed to reveal the status of the Cylindropuntieae, which may yet be monophyletic. Of the 42 genera with more than one exemplar in our study, only 17 were monophyletic; 14 of these genera were from subfamily Cactoideae and three from subfamily Opuntioideae. We present a synopsis of the status of the currently recognized genera.
© The Willi Hennig Society 2011.     

Molecular phylogeny of Pamphagidae (Acridoidea,Orthoptera) from China based on mitochondrial cytochrome oxidase II sequences

Abstract Phylogenetic relationships of Pamphagidae were examined using cytochrome oxidase subunit II (COII) mtDNA sequences (684 bp). Twenty‐seven species of Acridoidea from 20 genera were sequenced to obtain mtDNA data, along with four species from the GenBank nucleotide database. The purpose of this study was analyzing the phylogenetic relationships among subfamilies within Pamphagidae and interpreting the phylogenetic position of this family within the Acridoidea superfamily. Phylogenetic trees were reconstructed using neighbor‐joining (NJ), maximum parsimony (MP) and Bayesian inference (BI) methods. The 684 bp analyzed fragment included 126 parsimony informative sites. Sequences diverged 1.0%–11.1% between genera within subfamilies, and 8.8%–12.3% between subfamilies. Amino acid sequence diverged 0–6.1% between genera within subfamilies, and 0.4%–7.5% between subfamilies. Our phylogenetic trees revealed the monophyly of Pamphagidae and three distinct major groups within this family. Moreover, several well supported and stable clades were found in Pamphagidae. The global clustering results were similar to that obtained through classical morphological classification: Prionotropisinae, Thrinchinae and Pamphaginae were monophyletic groups. However, the current genus Filchnerella (Prionotropisinae) was not a monophyletic group and the genus Asiotmethis (Prionotropisinae) was a sister group of the genus Thrinchus (Thrinchinae). Further molecular and morphological studies are required to clarify the phylogenetic relationships of the genera Filchnerella and Asiotmethis.     

Evolutionary integration of the frog cranium

Evolutionary integration (covariation) of traits has long fascinated biologists because of its potential to elucidate factors that have shaped morphological evolution. Studies of tetrapod crania have identified patterns of evolutionary integration that reflect functional or developmental interactions among traits, but no studies to date have sampled widely across the species-rich lissamphibian order Anura (frogs). Frogs exhibit a vast range of cranial morphologies, life history strategies, and ecologies. Here, using high-density morphometrics we capture cranial morphology for 172 anuran species, sampling every extant family. We quantify the pattern of evolutionary modularity in the frog skull and compare patterns in taxa with different life history modes. Evolutionary changes across the anuran cranium are highly modular, with a well-integrated “suspensorium” involved in feeding. This pattern is strikingly similar to that identified for caecilian and salamander crania, suggesting replication of patterns of evolutionary integration across Lissamphibia. Surprisingly, possession of a feeding larval stage has no notable influence on cranial integration across frogs. However, late-ossifying bones exhibit higher integration than early-ossifying bones. Finally, anuran cranial modules show diverse morphological disparities, supporting the hypothesis that modular variation allows mosaic evolution of the cranium, but we find no consistent relationship between degree of within-module integration and disparity.     

Symbiotic Relationships of Legumes and Nodule Bacteria on Barro Colorado Island,Panama: A Review

New data on 129 bacterial isolates were analyzed together with prior samples to characterize community-level patterns of legume–rhizobial symbiosis on Barro Colorado Island (BCI), Panama. Nodules have been sampled from 24 BCI legume species in 18 genera, representing about one quarter of the legume species and one half of the genera on the island. Most BCI legumes associated exclusively with nodule symbionts in the genus Bradyrhizobium, which comprised 86.3% of all isolates (315 of 365). Most of the remaining isolates (44 of 365) belonged to the β-proteobacterial genus Burkholderia; these were restricted to two genera in the legume subfamily Mimosoideae. Multilocus sequence analysis indicated that BCI Bradyrhizobium strains were differentiated into at least eight lineages with deoxyribonucleic acid divergence of the same magnitude as found among currently recognized species in this bacterial genus. Two of these lineages were widely distributed across BCI legumes. One lineage was utilized by 15 host species of diverse life form (herbs, lianas, and trees) in 12 genera spanning two legume subfamilies. A second common lineage closely related to the taxon B. elkanii was associated with at least five legume genera in four separate tribes. Thus, BCI legume species from diverse clades within the family frequently share interaction with a few common lineages of nodule symbionts. However, certain host species were associated with unique symbiont lineages that have not been found on other coexisting BCI legumes. More comprehensive sampling of host taxa will be needed to characterize the overall diversity of nodule bacteria and the patterns of symbiont sharing among legumes in this community.     

A new taxonomy of the family Teloschistaceae

The lichen family Teloschistaceae is one of the larger families of lichenized fungi. Currently it includes one very large heterogenous genus, Caloplaca, with some 1000 or more species with a vast variation in morphology, anatomy and chemistry. The rest of the family is split into 10–15 smaller genera, each with 20 or fewer species. There is no modern classification of the family based on molecular data. Here we attempt a first phylogenetic evaluation of a large part of the family, including a total of 337 species. Of these, 162 were used in a combined phylogenetic analysis of the ribosomal RNA sequence markers nrITS, nrLSU and mrSSU, using Bayesian inference. We also analysed all species using nrITS data, split into four different analyses. As a result we propose a new classification of the family, where a total of 39 genera are recognized, of which 31 are newly described or resurrected. The new genera are: Athallia, Austroplaca, Bryoplaca, Calogaya, Cerothallia, Flavoplaca, Gondwania, Haloplaca, Orientophila, Pachypeltis, Parvoplaca, Rufoplaca, Shackletonia, Scutaria, Sirenophila, Solitaria, Squamulea, Stellarangia, Teloschistopsis, Usnochroma, Variospora, Villophora and Wetmoreana. Resurrected genera are Blastenia, Dufourea, Follmannia, Gyalolechia, Leproplaca, Polycauliona, Pyrenodesmia and Xanthocarpia. The species Orientophila subscopularis is described as new. A third subfamily, Teloschistoideae, is proposed to accommodate the genus Teloschistes and related genera, parallel to the two previously recognized subfamilies Xanthorioideae and Caloplacoideae. We also show the large plasticity in both morphological and anatomical characters between closely related species within genera, indicating the low value of these as evolutionary markers. The secondary chemistry is a better marker in some parts of the family. We recognize a large number of geographically delimited clades with clear centres of evolution, but often showing large variation in morphology and anatomy.     

A reappraisal of the family Comasteridae A. H. Clark, 1908 (Echinodermata: Grinoidea), with the description of a new subfamily and a new genus

The family Comasteridae is divided into four subfamilies on the basis of syzygy positions in the division series and proximal arm. Use of this character results in major differences in perceived relationships between genera compared with that obtained when traditional subfamilial characters are used. The primary character previously used to determine comasterid subfamilies, cirrus ornamentation, is rejected as unimportant in recognizing taxa any higher than species. Seventeen genera, including a new genus from the tropical western Atlantic, and 99 species are now recognized in the Comasteridae. Four of these genera and five species cannot be placed within the newly diagnosed subfamilies. Characters of generic importance are considered to be pinnule comb form, mouth position and, to a lesser extent, the characters traditionally used to distinguish genera.     

Redefining the damselfly families: a comprehensive molecular phylogeny of Zygoptera (Odonata)

An extensive molecular phylogenetic reconstruction of the suborder Zygoptera of the Odonata is presented, based on mitochondrial (16S, COI) and nuclear (28S) data of 59% of the 310 genera recognized and all (suspected) families except the monotypic Hemiphlebiidae. A partial reclassification is proposed, incorporating morphological characters. Many traditional families are recovered as monophyletic, but reorganization of the superfamily Coenagrionoidea into three families is proposed: Isostictidae, Platycnemididae and Coenagrionidae. Archboldargia Lieftinck, Hylaeargia Lieftinck, Palaiargia Förster, Papuargia Lieftinck and Onychargia Selys are transferred from Coenagrionidae to Platycnemididae, and Leptocnemis Selys, Oreocnemis Pinhey and Thaumatagrion Lieftinck from Platycnemididae to Coenagrionidae. Each geographically well‐defined clade of Platycnemididae is recognized as a subfamily, and thus Disparoneurinae (i.e. Old World ‘Protoneuridae’) is incorporated, Calicnemiinae is restricted, and Allocnemidinae (type genus: Allocnemis Selys) subfam.n ., Idiocnemidinae (type genus: Idiocnemis Selys) subfam.n . and Onychargiinae (type genus: Onychargia Selys) subfam.n . and Coperini trib.n . (type genus: Copera Kirby) are described. Half of Coenagrionidae belongs to a well‐supported clade incorporating Coenagrion Kirby and the potential subfamilies Agriocnemidinae, Ischnurinae and Pseudagrioninae. The remainder is less well defined, but includes the Pseudostigmatidae and New World Protoneuridae that, with Argiinae and Teinobasinae, may prove valid subfamilies with further evidence. Ninety‐two per cent of the genera formerly included in the polyphyletic Amphipterygidae and Megapodagrionidae were studied. Pentaphlebiidae, Rimanellidae and Devadattidae fam.n . (type genus: Devadatta Kirby) are separated from Amphipterygidae, and Argiolestidae, Heteragrionidae, Hypolestidae, Philogeniidae, Philosinidae and Thaumatoneuridae from Megapodagrionidae. Eight further groups formerly placed in the latter are identified, but are retained as incertae sedis; the validity of Lestoideidae, Philogangidae and Pseudolestidae is confirmed. For some families (e.g. Calopterygidae, Chlorocyphidae) a further subdivision is possible; Protostictinae subfam.n . (type genus: Protosticta Selys) is introduced in Platystictidae. Numerous new combinations are proposed in the Supporting Information. Many long‐established families lack strong morphological apomorphies. In particular, venation is incongruent with molecular results, stressing the need to review fossil Odonata taxonomy: once defined by the reduction of the anal vein, Protoneuridae dissolves completely into six clades from five families.     

Nucleolar organiser region (NOR) location in karyotypes of Australian ground frogs (family Myobatrachidae)

Nucleolar organiser regions (NORs) were examined in over 90% of the species of Australian ground frogs (familiy Myobatrachidae), including representatives from all twenty currently recognised genera and the three subfamilies. Throughout the family, location of the NOR within the karyotype showed considerable variation yet karyotype morphology showed uniformity. The precise mechanism(s) whereby variation in NOR location evolved while karyotype morphology was unchanged remains uncertain. Comparison of the two major subfamilies showed that the Limnodynastinae had a greater diversity of NOR location than the Myobatrachinae. The limnodynastine genus,Heleioporus, was the only one to show multiple NOR sites in several species. NOR location was particularly stable within most polytypic genera. Differences in NOR location within the remaining polytypic genera (Heleioporus, Limnodynastes, Neobatrachus, Philoria andRanidella) pointed to taxonomic discriminations that were generally consistent with recent proposals based on other criteria.     

Evolution and systematics of the Late Paleozoic brachiopod family Linoproductidae with descriptions of species from the Lower Permian of the Northern Timan

The analysis of the morphological features of the genera of the family Linoproductidae in the Late Paleozoic substantiates its three subfamilies as three evolutionary trends beginning with the initial subfamily Coopericinae Lazarev, 2004, which is known from the beginning of the Early Carboniferous and two its derivatives: subfamily Linipalinae subfam. nov., which appeared in the Podolskian Time (Upper Moscovian Age), and subfamily Linoproductinae Stehli, 1954, which appeared in the Kasimovian Age. The problems and prospects of the further detailing of the system of these subfamilies are discussed. Three new species of the genus Sublinoproductus are described from the Lower Permian of Northern Timan.     

Systematic classification of Asian colobines

In order to study the differentiation of Asian colobines, fourteen variables were analysed in one way, on 123 skulls, includingRhinopithecus, Presbytis, Presbytiscus, Pygathrix, andNasalis with both cluster and differentiated functions tests. Information on paleoenvironment changes in China and South-East Asia since late Tertiary have been used to examine the influences of migratory habits and the distribution range in Asian colobines. The cladogram among different Asian colobines genera was made from the results of various analysis. Some new points or revisions were suggested: 1. Following the second migratory way, ancient species of Asian colobines perhaps passed through Xizang along the northern bank of Tethis sea and Heng-Duan Shan regions, across Yunnan into Vietnam, since the ancient continent between Yunnan and Xizang was already located in on eastern bank of Tethis sea. Thus, during the evolution, Asian colobines must have had two original centres, i.e. “Sundaland” and Heng-Duan Shan Chinese regions; 2. Pygatrix possesses a lot of cranial features more similar toPresbytiscus than toRhinopithecus. The small difference from the modification combinesPygatrix with other two genera as shown by Groves (1970), but it is better to putPygatrix andPresbytiscus together as one genus; 3.Nasalis (2n=48) may be the most primitive genus within Asian colobines. Some features shared withRhinopithecus, for example body size, terrestrial activities, limb proportion etc. ...seem to be considered as a common inheritance of symlesiomorphus characters; 4.Rhinopithecus, with reference to cranioface and cranium or to its origin, is a special genus of Asian colobine. It may represent the highest level of evolutionary position within various genera (Peng et al., 1985).     

Effects of different kinds of cranial deformation on the incidence of wormian bones

Researchers have debated whether the presence and frequency of wormian bones (sutural bones, supernumerary bones, and ossicles) are attributable to genetic factors, environmental factors, or both. This research examines the effects of many different kinds of cranial deformation on the incidence of wormian bones. A sample of 127 deformed and undeformed crania from New World archaeological sites was examined. An undeformed cranial sample (n=35) was compared to the following cranially deformed groups: 1) occipital, 2) lambdoid, 3) annular, 4) fronto-vertico-occipital, 5) parallelo-fronto-occipital, and 6) sagittal synostosis. Three levels of degree of cultural cranial deformation were qualitatively determined. Type and number of wormian bones along each major suture were recorded for each cranium. Group means were analyzed using Kruskal-Wallis one-way ANOVA statistical tests to test the null hypothesis that cranial deformation does not have an effect on wormian bone incidence. Results indicate that all forms of cranial deformation affect the frequency of some types of wormian bones. In particular, all cranially deformed groups exhibited significantly greater frequencies of lambdoid ossicles. Apical, parieto-mastoid, and occipito-mastoid wormian bones also appeared with greater frequency in some groups of culturally deformed crania. Further, varying degrees of cultural deformation all had more lambdoid wormian bones than the undeformed group. These results suggest that wormian bone development in posteriorly placed sutures may be affected more by environmental forces than are their anteriorly placed counterparts.     

Evaluation of a New Method of Fossil Retrodeformation by Algorithmic Symmetrization: Crania of Papionins (Primates,Cercopithecidae) as a Test Case

Diagenetic distortion can be a major obstacle to collecting quantitative shape data on paleontological specimens, especially for three-dimensional geometric morphometric analysis. Here we utilize the recently -published algorithmic symmetrization method of fossil reconstruction and compare it to the more traditional reflection & averaging approach. In order to have an objective test of this method, five casts of a female cranium of Papio hamadryas kindae were manually deformed while the plaster hardened. These were subsequently “retrodeformed” using both algorithmic symmetrization and reflection & averaging and then compared to the original, undeformed specimen. We found that in all cases, algorithmic retrodeformation improved the shape of the deformed cranium and in four out of five cases, the algorithmically symmetrized crania were more similar in shape to the original crania than the reflected & averaged reconstructions. In three out of five cases, the difference between the algorithmically symmetrized crania and the original cranium could be contained within the magnitude of variation among individuals in a single subspecies of Papio. Instances of asymmetric distortion, such as breakage on one side, or bending in the axis of symmetry, were well handled, whereas symmetrical distortion remained uncorrected. This technique was further tested on a naturally deformed and fossilized cranium of Paradolichopithecus arvernensis. Results, based on a principal components analysis and Procrustes distances, showed that the algorithmically symmetrized Paradolichopithecus cranium was more similar to other, less-deformed crania from the same species than was the original. These results illustrate the efficacy of this method of retrodeformation by algorithmic symmetrization for the correction of asymmetrical distortion in fossils. Symmetrical distortion remains a problem for all currently developed methods of retrodeformation.