On the systematic position ofChaetomys subspinosus (Rodentia: Caviomorpha) based on evidence from the incisor enamel microstructure

We have studied the incisor enamel microstructure ofChaetomys subspinosus and other possibly closely related caviomorph rodents.Chaetomys subspinosus lacks the important synapomorphy of the Octodontoidea, reactangular plate-like interprismatic matrix (IPM) in the portio interna (PI) of the incisor Schmelzmuster. Therefore its transfer from the Erethizontidae to the Echimyidae, as proposed by Patterson and Wood (Bull. Mus. Comp. Zool. 149, 371–543, 1982) based on retention of dP4, is contested. The parallel to acute angular IPM in the PI ofChaetomys and the Erethizontidae is a symplesiomorphy and does not indicate close relationship. Contrary to previous claims, a posterior carotid foramen is also retained inChaetomys. Chaetomys is characterized by an unusual thin enamel which is considered primitive after outgroup comparison. Therefore, it is proposed to leaveChaetomys in the monospecific erethizontid subfamily Chaetomyinae, until additional information on the species is available.     

Allometric disparity in rodent evolution

In this study, allometric trajectories for 51 rodent species, comprising equal representatives from each of the major clades (Ctenohystrica, Muroidea, Sciuridae), are compared in a multivariate morphospace (=allometric space) to quantify magnitudes of disparity in cranial growth. Variability in allometric trajectory patterns was compared to measures of adult disparity in each clade, and dietary habit among the examined species, which together encapsulated an ecomorphological breadth. Results indicate that the evolution of allometric trajectories in rodents is characterized by different features in sciurids compared with muroids and Ctenohystrica. Sciuridae was found to have a reduced magnitude of inter‐trajectory change and growth patterns with less variation in allometric coefficient values among members. In contrast, a greater magnitude of difference between trajectories and an increased variation in allometric coefficient values was evident for both Ctenohystrica and muroids. Ctenohystrica and muroids achieved considerably higher adult disparities than sciurids, suggesting that conservatism in allometric trajectory modification may constrain morphological diversity in rodents. The results provide support for a role of ecology (dietary habit) in the evolution of allometric trajectories in rodents.     

Enamel microstructure ofTribosphenomys (Mammalia,glires): Character analysis and systematic implications

Enamel distribution on the upper and lower incisors ofTribosphenomys minutus (from Late Paleocene-Early Eocene of Inner Mongolia of China) is typically rodent-like, i.e., primarily confined to the anterior surface throughout these transversely compressed, evergrowing teeth. AlthoughTribosphenomys incisor enamel is differentiated into two layers, it does not possess Hunter-Schreger bands (HSB). The incisor and molar enamels are radial in type, a condition regarded as either an autapomorph or a primtive retention forTribosphenomys. Character polarities concerning enamel thickness, enamel layer number, HSB, enamel types, and functional and phylogenetic implications of the enamel structures are discussed. Overall, enamel microstructural evolution at high taxonomic levels within Glires displays considerably more homoplasy than generally appreciated. A phylogenetic definition of Rodentia is proposed.Tribosphenomys is the sister-group of a taxon here named Rodentia, and thus is not itself a member of the order, from a systematic viewpoint.     

Distribution and variation in enamel structure in the oral teeth of sarcopterygians: Its significance for the evolution of a protoprismatic enamel

The property of tooth enamel to resist alteration during fossilization, is used to analyse the unique arrangements of biological crystallites amongst genera of Paleozoic sarcopterygians, with both polarized light and s.e.m. Previous concepts of crystallite organization in reptiles and mammal‐like reptiles are evaluated. Two of the Devonian sarcopterygians, are shown to exhibit a protoprismatic pattern, identical with that of a stem group therian. The patterns of crystallites, together with the arrangement of incremental lines establish that this tissue is solely an ectodermal product; monotypic enamel, in contrast to bitypic enamel with two cell products contributing to it as in enameloid or acrodin. Each genus examined has a different pattern, of significance in considering relationships amongst sarcopterygians. Recent information on ganoine and some new findings on enamel in extant lungfishes have led to the conclusion that types of monotypic enamel are present in both actinopterygians and sarcopterygians, and challenges the use of monotypic enamel as a synapomorphy of sarcopterygians in cladistic analyses.     

Intraspecific variation in M1 enamel development in modern humans: implications for human evolution

The timing and sequence of enamel development, as well as enamel thickness, was documented for individual cusps (protoconid, hypoconid, metaconid, entoconid) in 15 unworn permanent lower first molars (M(1)s) from a sample of modern human juveniles. These data were compared with previously published data for modern and fossil species reported in the literature. Crown formation in all teeth was initiated in the protoconid and completed in the hypoconid. These cusps had significantly longer formation times (2.91 and 2.96 yrs, respectively) than the metaconid and entoconid (2.52 and 2.38 yrs, respectively), as well as thicker enamel, and each represented between 92-95% of the total crown formation time. Rates of enamel secretion in all cusps increased significantly from 2.97 microm in the inner enamel to 4.47 microm in the outer enamel. Two cusps of one individual were studied in more detail and did not follow this typical trajectory. Rather, there was a sharp decrease in the middle of enamel formation and then a slow recovery of secretion rates from the mid- to outer enamel. This anomalous trajectory of enamel formation is discussed in the context of other nondental tissue responses to illness. Neither secretion rates nor periodicity differed significantly when compared between the cusps of each molar. Differences in cusp formation times, initiation, and completion suggest a relationship between the rates of enamel formation and enamel thickness. This fits with expectations about the mechanics of the chewing cycle and general lower molar morphology. A comparison with similar data for some nonhuman primates and fossil hominoids suggests this relationship may hold true across several primate taxa. Other aspects of enamel growth differed between this human sample and certain fossil species. The lower molars formed slowly over a longer period of time, which may reflect the extended growth period of modern humans. The methodological approach adopted in this study is discussed in the context of that used in other studies.     

Three-dimensional molar enamel distribution and thickness in Australopithecus and Paranthropus

Thick molar enamel is among the few diagnostic characters of hominins which are measurable in fossil specimens. Despite a long history of study and characterization of Paranthropus molars as relatively 'hyper-thick', only a few tooth fragments and controlled planes of section (designed to be proxies of whole-crown thickness) have been measured. Here, we measure molar enamel thickness in Australopithecus africanus and Paranthropus robustus using accurate microtomographic methods, recording the whole-crown distribution of enamel. Both taxa have relatively thick enamel, but are thinner than previously characterized based on two-dimensional measurements. Three-dimensional measurements show that P. robustus enamel is not hyper-thick, and A. africanus enamel is relatively thinner than that of recent humans. Interspecific differences in the whole-crown distribution of enamel thickness influence cross-sectional measurements such that enamel thickness is exaggerated in two-dimensional sections of A. africanus and P. robustus molars. As such, two-dimensional enamel thickness measurements in australopiths are not reliable proxies for the three-dimensional data they are meant to represent. The three-dimensional distribution of enamel thickness shows different patterns among species, and is more useful for the interpretation of functional adaptations than single summary measures of enamel thickness.     

Fructooligosaccharides consumption inhibits the loss of iron from the incisor enamel surface in gastrectomized rat

We examine the effects of fructooligosaccharides (FOS) on the reduction in the incisor iron content in gastrectomized rat. Twenty-eight 5-wk-old male Sprague-Dawley rats were divided into two groups: sham operated (bSH) and gastrectomized (bGX). After 4 wk each group was divided into two subgroups according to the presence or absence of 7.5% FOS in the synthetic diet (SH, SH+FOS, GX, and GX+FOS). At 10 wk wafter surgery, the maxilla was prepared to examine the iron content of the incisor enamel surface at four points. These points corresponded to the iron content at 6,7,8, and 10 wk, respectively. Blood was collected to determine serum iron levels at 4 and 10 wk. The serum iron level significantly decreased at 4 and 10 wk the GX group. At 10 wk, the level in the GX+FOS group significantly increased but did not reaach that in the SH group. The iron content of the enamel surface time-dependently increased and no significant differences were seen between SH and GX+FOS at 8 and 10 wk. These results suggest that FOS consumption impaired the loss of enamel content following gastrectomy, and this effect preceded the effect on the serum iron level.     

A freeze-fracture study of the papillary layer of the rat incisor enamel organ

After tooth enamel has been secreted it undergoes maturation or hardening. This process is mediated by ruffled and smooth-ended ameloblasts and associated papillary layer cells. The cells of the papillary layer are characterized by large numbers of mitochondria, coated vesicles, microvilli, and gap junctions. These features have led numerous investigators to speculate that the papillary layer is an ion-transporting epithelium. We have conducted freeze-fracture studies of the rat papillary layer in order to better characterize the surface features of these cells. The cell membranes of the papillary cells contained large numbers of intramembrane particles of various sizes ranging from 4 to 9 nm in diameter. Gap junctions were present at the cell surface and in the cytoplasm in the form of annular gap junctions. The intramembrane particles or connexons of both types of gap junctions were about 8-9 nm wide and were either packed randomly or present in the so-called 'crystallized' state. At the interface between smooth-ended ameloblasts and papillary layer cells, a well-developed zonula occludens was present along the basal surfaces of the ameloblasts and several large gap junctions were formed between the two cell types. The capillary network associated with the papillary layer was characterized by a thin endothelium containing large numbers of fenestrations.     

Phylogenetic implications of the Crustacean nauplius

The plesiomorphic mode of crustacean development is widely accepted to be via a larva called the nauplius. Extant taxa like the Cephalocarida, Branchiopoda, Ostracoda, Mystacocarida, Copepoda, Cirripedia, Ascothoracida, Facetotecta, Euphausiacea and Penaeidea hatch from an egg as a free-living nauplius. Other crustaceans show an embryonic phase of development suggestive of a naupliar organization. Several features of the nauplius larva have been proposed as diagnostic characters for the Crustacea: a median (nauplius) eye; at least three pairs of head appendages (antennules, antennae, mandibles); a posteriorly directed fold (the labrum) extending over the mouth and a cephalic (nauplius) shield. The relationship between trilobite protaspis with at least four appendages and the crustacean nauplius remains unclear, but reports of a copepod orthonauplius with four appendages are rejected. Swimming is suggested to represent the underived mode of locomotion for the crustacean nauplius, and that naupliar swimming directly results in naupliar feeding which also is underived.     

Variation in hominoid molar enamel thickness

Enamel thickness has figured prominently in discussions of hominid origins for nearly a century, although little is known about its intra-taxon variation. It has been suggested that enamel thickness increases from first to third molars, perhaps due to varying functional demands or developmental constraints, but this has not been tested with appropriate statistical methods. We quantified enamel cap area (c), dentine area (b), and enamel-dentine junction length (e) in coronal planes of sections through the mesial and distal cusps in 57 permanent molars of Pan and 59 of Pongo, and calculated average (c/e) and relative enamel thickness (([c/e]/ radicalb) * 100). Posteriorly increasing or decreasing trends in each variable and average (AET) and relative enamel thickness (RET) were tested among molars in the same row. Differences between maxillary and mandibular analogues and between mesial and distal sections of the same tooth were also examined. In mesial sections of both genera, enamel cap area significantly increased posteriorly, except in Pan maxillary sections. In distal sections of maxillary teeth, trends of decreasing dentine area were significant in both taxa, possibly due to hypocone reduction. Significant increases in AET and RET posteriorly were found in all comparisons, except for AET in Pongo distal maxillary sections. Several significant differences were found between maxillary and mandibular analogues in both taxa. Relative to their mesial counterparts, distal sections showed increased enamel cap area and/or decreased dentine area, and thus increased AET and RET. This study indicates that when AET and RET are calculated from samples of mixed molars, variability is exaggerated due to the lumping of tooth types. To maximize taxonomic discrimination using enamel thickness, tooth type and section plane should be taken into account. Nonetheless, previous findings that African apes have relatively thinner enamel than Pongo is supported for certain molar positions.     

Cultural evolution: implications for understanding the human language faculty and its evolution

Human language is unique among the communication systems of the natural world: it is socially learned and, as a consequence of its recursively compositional structure, offers open-ended communicative potential. The structure of this communication system can be explained as a consequence of the evolution of the human biological capacity for language or the cultural evolution of language itself. We argue, supported by a formal model, that an explanatory account that involves some role for cultural evolution has profound implications for our understanding of the biological evolution of the language faculty: under a number of reasonable scenarios, cultural evolution can shield the language faculty from selection, such that strongly constraining language-specific learning biases are unlikely to evolve. We therefore argue that language is best seen as a consequence of cultural evolution in populations with a weak and/or domain-general language faculty.     

Age at first molar emergence in early Miocene Afropithecus turkanensis and life-history evolution in the Hominoidea

Among primates, age at first molar emergence is correlated with a variety of life history traits. Age at first molar emergence can therefore be used to broadly infer the life histories of fossil primate species. One method of determining age at first molar emergence is to determine the age at death of fossil individuals that were in the process of erupting their first molars. This was done for an infant partial mandible of Afropithecus turkanensis (KNM-MO 26) from the approximately 17.5 Ma site of Moruorot in Kenya. A range of estimates of age at death was calculated for this individual using the permanent lateral incisor germ preserved in its crypt, by combining the number and periodicity of lateral enamel perikymata with estimates of the duration of cuspal enamel formation and the duration of the postnatal delay in the inception of crown mineralization. Perikymata periodicity was determined using daily cross striations between adjacent Retzius lines in thin sections of two A. turkanensis molars from the nearby site of Kalodirr. Based on the position of the KNM-MO 26 M(1)in relation to the mandibular alveolar margin, it had not yet undergone gingival emergence. The projected time to gingival emergence was estimated based on radiographic studies of M(1)eruption in extant baboons and chimpanzees.The estimates of age at M(1)emergence in KNM-MO 26 range from 28.2 to 43.5 months, using minimum and average values from extant great apes and humans for the estimated growth parameters. Even the absolute minimum value is well outside the ranges of extant large Old World monkeys for which there are data (12.5 to <25 months), but is within the range of chimpanzees (25.7 to 48.0 months). It is inferred, therefore, that A. turkanensis had a life history profile broadly like that of Pan. This is additional evidence to that provided by Sivapithecus parvada (Function, Phylogeny, and Fossils: Miocene Hominoid Evolution and Adaptations, 1997, 173) that the prolonged life histories characteristic of extant apes were achieved early in the evolutionary history of the group. However, it is unclear at present whether life-history prolongation in apes represents the primitive catarrhine pace of life history extended through phyletic increase in body mass, or whether it is derived with respect to a primitive, size-adjusted life history that was broadly intermediate between those of extant hominoids and cercopithecoids. Life history evolution in primates as a whole may have occurred largely through a series of grade-shifts, with the establishment of fundamental life-history profiles early in the histories of major higher taxa. These may have included shifts that were largely body mass dependent, as well as those that occurred in the absence of significant changes in body mass.     

Phylogenetic distribution and evolution of mycorrhizas in land plants

A survey of 659 papers mostly published since 1987 was conducted to compile a checklist of mycorrhizal occurrence among 3,617 species (263 families) of land plants. A plant phylogeny was then used to map the mycorrhizal information to examine evolutionary patterns. Several findings from this survey enhance our understanding of the roles of mycorrhizas in the origin and subsequent diversification of land plants. First, 80 and 92% of surveyed land plant species and families are mycorrhizal. Second, arbuscular mycorrhiza (AM) is the predominant and ancestral type of mycorrhiza in land plants. Its occurrence in a vast majority of land plants and early-diverging lineages of liverworts suggests that the origin of AM probably coincided with the origin of land plants. Third, ectomycorrhiza (ECM) and its derived types independently evolved from AM many times through parallel evolution. Coevolution between plant and fungal partners in ECM and its derived types has probably contributed to diversification of both plant hosts and fungal symbionts. Fourth, mycoheterotrophy and loss of the mycorrhizal condition also evolved many times independently in land plants through parallel evolution.     

Incisor Schmelzmuster Diversity in South America's Oldest Rodent Fauna and Early Caviomorph History

Investigation of the enamel microstructure of 20 isolated rodent incisors from the ?Eocene Santa Rosa local fauna (Peru) yielded exclusively schmelzmuster with multiserial Hunter–Schreger bands (HSB). All three subtypes of multiserialHSB with parallel, acute angular, and rectangular interprismatic matrix (IPM) that were previously reported for caviomorph rodents are present. Two lower incisors with rectangular IPM can be attributed to the Octodontoidea, a caviomorph superfamily exhibiting this highly derived enamel type. The plesiomorphic pauciserial condition that characterizes early Paleogene rodents such as North American Ischyromyoidea (including “Franimorpha”) has not been detected. It is therefore probable that the founder populations of South American Caviomorpha already possessed a derived incisor schmelzmuster with multiserial HSB that is shared with African Thryonomyoidea. Because on the North American continent a possible stem-lineage representative of Caviomorpha with multiserial HSB has never been detected, incisor enamel microstructure supports the hypothesis of an African origin of Caviomorpha from a common ancestor shared with Thryonomyoidea.     

Electron microscopy of dental enamel: Analysis of crystal lattice images

Summary Thin sections of rat incisor enamel were studied with the electron microscope. Fringe patterns having repeat periods in the range 3.1–8.2 Å were seen in individual enamel crystals. These images were interpreted as representing the resolution of corresponding planes in the hydroxyapatite crystal lattice. The lattice spacings and interplanar angles were identified by comparing the observations with available data derived from X-ray diffraction analysis.     

A ctenodactylid rodent (Mammalia: Rodentia) from the Middle Miocene of Chios Island (Greece)

Field work carried out in 1991 and 1993 at the Middle Miocene locality of Thymiana (Chios Island, Greece), produced much rodent material including a number of ctenodactylid dental specimens. They represent a single taxon whose upper and lower cheek tooth morphology clearly differs from Prosayimys flynni and all Sayimys species known so far, except for Sayimys intermedius. The ctenodactylid teeth from the Middle Miocene of Chios are identified as pertaining to the latter species, despite minor differences from the Pakistani and Saudi Arabian representatives of S. intermedius.     

Early Miocene cricetids (Rodentia) from the Junggar basin (Xinjiang, China) and their biochronological implications

Among the 14 small mammal species from the early Miocene locality of the northern area of the Junggar basin (northern Xinjiang, China), four species are cricetids (Rodentia): unnamed species of Cricetodon and Eumyarion, and two new species, Karydomys debruijni nov. sp. and Megacricetodon beijiangensis nov. sp. Some aspects of the morphology of Cricetodon sp. are shared by Eucricetodon from the late Oligocene, suggesting that these specimens could be of intermediate form between Eucricetodon and Cricetodon. One tooth of Eumyarion sp. was found, making its determination uncertain, but its morphology is clearly differentiated from the one of Cricetodon sp. The species K. debruijni nov. sp. is established based on its primitive features compared to the species known in Europe and Anatolia, and its specific association of characters compared to Karydomys dzerzhinskii. M. beijiangensis nov. sp. shows many plesiomorphic features compared to the species already described in the middle Miocene of China. Based on both the whole assemblage of rodents and the species of cricetids, the biochronologic position and the age of the locality are discussed. The locality appears to be biochronologically very close to the fauna from the Chul’adyr Formation in Aktau Mountains, but we propose an age slightly older than the one proposed for this fauna, probably equivalent to the MN3 biozone in Europe.     

Antimicrobial strength increases with group size: implications for social evolution

We hypothesize that aggregations of animals are likely to attract pathogenic micro-organisms and that this is especially the case for semisocial and eusocial insects where selection ultimately led to group sizes in the thousands or even millions, attracting the epithet 'superorganism'. Here, we analyse antimicrobial strength, per individual, in eight thrips species (Insecta: Thysanoptera) that present increasing innate group sizes and show that species with the largest group size (100-700) had the strongest antimicrobials, those with smaller groups (10-80) had lower antimicrobial activity, while solitary species showed none. Species with large innate group sizes showed strong antimicrobial activity while the semisocial species showed no activity until group size increased sufficiently to make activity detectable. The eusocial species behaved in a similar way, with detectable activity appearing once group size exceeded 120. These analyses show that antimicrobial strength is determined by innate group size. This suggests that the evolution of sociality that, by definition, increases group size, may have had particular requirements for defences against microbial pathogens. Thus, increase in group size, accompanied by increased antibiotic strength, may have been a critical factor determining the 'point of no return', early in the evolution of social insects, beyond which the evolution of social anatomical and morphological traits was irreversible. Our data suggest that traits that increase group size in general are accompanied by increased antimicrobial strength and that this was critical for transitions from solitary to social and eusocial organization.