Effects of macrophyte heterogeneity and food availability on structural parameters of the macroinvertebrate community in a Pampean stream

Environmental heterogeneity in natural ecosystems influences several parameters at the population and community levels. In freshwater ecosystems, habitat heterogeneity can be provided by macrophyte species with different structural shapes. Previous studies suggest that aquatic plants with more complex architectures will support higher number, biomass, and taxon richness of macroinvertebrates than plants with simpler shape. We investigated the influence of macrophyte structural heterogeneity (quantified by fractal dimension) and food availability (represented by epiphytic biomass) on several parameters (number of individuals, biomass, body size distribution, taxon richness, and diversity) of the macroinvertebrate community in a Pampean stream. Four submerged macrophyte species (Egeria densa, Elodea ernstae, Ceratophyllum demersum, and Stuckenia striata) and associated macroinvertebrates were sampled in late spring, summer, and autumn. Plants were photographed and fractal dimension was estimated from the images by the box-counting method. Fractal dimension was independent of plant surface area per unit of macrophyte biomass and differed significantly among species. Mean fractal dimension varied between 1.29 and 1.62, and increased following the sequence E. densa → S. striata → E. ernstae → C. demersum. Macrophyte species with higher fractal dimension supported a greater abundance of macroinvertebrates, especially those of small body size (500–1,000 μm); but fractal dimension was unrelated to macroinvertebrate biomass, richness, and diversity. However, overall animal biomass was significantly associated to the epiphytic abundance. Consequently, macrophyte heterogeneity influences macroinvertebrate density and body size distribution, while animal biomass depends on epiphytic food resources provided by plants.     

The <Emphasis Type="Italic">Eumyarion</Emphasis> (Mammalia,Rodentia, Muridae) assemblage from Sandelzhausen (Miocene,Southern Germany): a test on homogeneity

A large sample of Eumyarion cheek teeth (N = 569) from the Early/Middle Miocene boundary locality Sandelzhausen (MN5, Southern Germany), type locality E. bifidus Fahlbusch, 1964, is studied. It is concluded that this collection contains two species: E. bifidus and E. weinfurteri Schaub and Zapfe, 1953. The similarity in size and morphology of the cheek teeth of these two species is so great that only the M1 and M2 can be recognised with certainty. Eumyarion bifidus seems to be a descendant of E. orhani de Bruijn et al., 2006 from the Early Miocene (MN3) of Southwestern Anatolia and is therefore considered to be an immigrant into Central Europe.      

Tachyoryctes makooka (Tachyoryctini,Spalacidae, Rodentia) and its bearing on the phylogeny of the Tachyoryctini

Abstract: The Rhizomyinae is a subfamily of myomorph rodents within the family Spalacidae. It is subdivided into two tribes: the Asian Rhizomyini and the Tachyoryctini. Even though the origin of the Tachyoryctini is to be found in Asia, they are nowadays restricted to Africa. African Tachyoryctini are known from the Late Miocene and include a single genus with five species: the recently discovered Tachyoryctes makooka, which is described in detail in this work, T. pliocaenicus, T. konjiti and the two extant T. splendens and T. macrocephalus. Their closest Asian counterparts are the Late Miocene Protachyoryctes and Eicooryctes. A cladistic analysis involving all the above‐mentioned taxa was carried out. Protachyoryctes tatroti turned out to be the most basal species of the ingroup followed by the Ethiopian T. makooka. Both taxa show some primitive traits like the protosinus on the first upper molars, short but distinct mesolophids on the lower molars and the presence of the mure on the cheek teeth. T. pliocaenicus and the most derived Tachyoryctini share the synapomorphic reduction or loss of these characters. The increase in hypsodonty evidenced in this lineage is correlated with the transition from a humid to a drier climate that started at the beginning of the Pliocene.     

Prostrepsiceros and Protragelaphus (Artiodactyla,Mammalia) from the Late Miocene locality of Chomateri (Attica,Greece)

The Late Miocene fossiliferous locality of Chomateri is located close to the classic locality of Pikermi, but unfortunately its faunal context is not sufficiently known. Some fossil remains of spiral-horned antelopes from Chomateri are assigned to Prostrepsiceros rotundicornis and Protragelaphus skouzesi. The co-existence of P. rotundicornis and P. skouzesi is relatively rare in the Late Miocene, but well-documented in Pikermi. Comparisons of the Chomateri material support a middle Turolian age for the Chomateri locality and suggest a close chronological relation to the Pikermi fauna.     

Extracting complexity waveforms from one-dimensional signals

Background  

Nonlinear methods provide a direct way of estimating complexity of one-dimensional sampled signals through calculation of Higuchi's fractal dimension (1<FD<2). In most cases the signal is treated as being characterized by one value of FD and consequently analyzed as one epoch or, if divided into more epochs, often only mean and standard deviation of epoch FD are calculated. If its complexity variation (or running fractal dimension), FD(t), is to be extracted, a moving window (epoch) approach is needed. However, due to low-pass filtering properties of moving windows, short epochs are preferred. Since Higuchi's method is based on consecutive reduction of signal sampling frequency, it is not suitable for estimating FD of very short epochs (N < 100 samples).     

Power scaling of ammonitic suture patterns from Cretaceous Ancyloceratina: constraints on septal/sutural complexity

The spatial scaling of 77 hemisutures from 65 species of Cretaceous heteromorphic ammonites was quantified with the fractal box‐counting method. Fractal dimensions within Baculites compressus did not significantly differ between adult hemisutures; however, the juvenile suture of this species did exhibit a significantly lower fractal dimension. This suggests that variation in sutural complexity between explicitly adult ontogenetic stages may not contribute to significant noise in comparisons between other species/morphotypes. High‐spired, three‐dimensionally coiled heteromorphs with a larger degree of septal asymmetry exhibit higher fractal dimensions in outer whorl hemisutures than inner whorl hemisutures due to their elongation and improved space occupation over a larger whorl surface. Three‐dimensionally coiled ammonites also have higher fractal dimensions on average (mean D_b = 1.45) with respect to their 2‐D coiled counterparts (mean D_b = 1.38). All ammonites in this study exhibit a positive trend between sutural complexity and shell size (proxied by whorl height). These relationships suggest that septal frilling is constrained by shell morphology and whorl section geometry during septal morphogenesis. This, in turn, influences the scaling, space‐filling properties and scaling limits of ammonitic suture patterns. Sutural/septal complexity is also found to positively influence the amount of liquid retained in marginal septal recesses. However, as these septa approach larger scales, less cameral liquid is retained per septal mass. This may further explain the positive relationship between sutural complexity and shell size.     

Habitat complexity in aquatic systems: fractals and beyond

Despite the intensity with which ecological information involving habitat complexity has been amassed to date, much remains to be revealed for a comprehensive understanding of the mechanics and implications of the structural complexity of habitats and its influences on ecological communities. This review examines the multi-faceted characteristics of habitat complexity, focusing in particular on aquatic ecosystems. Habitat complexity in aquatic systems is characterised by at least five different traits of physical structure: (1) spatial scales, (2) diversity of complexity-generating physical (structural) elements, (3) spatial arrangement of elements, (4) sizes of elements, (5) abundance/density of elements. Of these five traits, the concept of fractal dimension fully encompasses only the last one; in this sense, habitat complexity is more complex than what fractal measures represent. It is therefore important to investigate exactly which traits of habitat structure are exerting influences on organisms/communities. We hypothesise that, where an entire range of possible fractal dimension D is considered, intermediate levels of D are most likely to be associated with the highest level of biodiversity, to which the body size spectra of assemblages would have a close bearing. In most aquatic ecosystems, broadly two-dimensional structures of bottom substrate at the scale of 1–10 m mean that the addition of vertical, three dimensional structures almost always implies an increase in both the ‘diversity’ and ‘abundance’ components of structural elements, resulting in more habitats being made available to organisms of different sizes and functional designs. The conservation and management of aquatic ecosystems would be facilitated by rigorous assessments of linkages between habitat complexity and aquatic communities, for which an integrative approach to habitat complexity seems to offer a useful and versatile framework.     

Phylogenetic analysis of Aphanius from the endorheic Kor River Basin in the Zagros Mountains,South‐western Iran (Teleostei: Cyprinodontiformes: Cyprinodontidae)

Morphologically similar populations of Aphanius that are currently considered as A. sophiae inhabit the endorheic Kor River Basin in the Zagros Mountains. Using genetic analysis based on mtDNA (cytochrome b), combined with examination of morphology (morphometry, meristics, otoliths), we discovered that what is thought to be A. sophiae is actually two distinct species, one of which is described as A. shirini sp. n. The males of the new species can be distinguished from those of all other Iranian inland Aphanius species by having only 7–10 clearly defined white flank bars, which is the lowest number of flank bars among the Iranian inland Aphanius species. Both males and females differ from all other Iranian inland Aphanius species by having a significantly longer caudal peduncle and a smaller dorsal fin depth. Based on the PhyML and Bayesian likelihood trees, A. shirini is sister to A. vladykovi from the Karoun Basin in the Zagros Mountains. Our results indicate that an ancient exorheic Kor River Basin existed in the Late Miocene and Pliocene. The close phylogenetic relationship between A. shirini and A. vladykovi suggests that the pre‐Pliocene drainage in the ancient Kor River Basin was directed to the north‐west (to the Karoun Basin), and not to the south‐east as in the present‐day Kor Basin. Both A. shirini and A. vladykovi represent the highest altitude records for Aphanius. We conclude that the splits of A. shirini and A. vladykovi can be linked to tectonic events in the Middle to Late Miocene, which created the highest altitudes (>3000 m) in the Zagros Mountains, and led to isolation of populations. The present‐day endorheic Kor Basin is known to have formed in the Late Pleistocene or Early Holocene, and the ‘young’ age of A. sophiae is clearly related to this history. Our results contribute to elucidate the link between geological history and the present‐day species diversity in the tectonically still active Zagros Mountains of Iran.     

Impact of climate changes from Middle Miocene onwards on evolutionary diversification in Eurasia: Insights from the mesobuthid scorpions

The aridification from Middle Miocene onwards has transformed the Asian interior into an arid environment, and the Pleistocene glacial–interglacial oscillations exerted further ecological impact. Therefore, both aridification and glaciation would have considerably influenced the evolution of many mid‐latitude species in temperate Asia. Here, we tested this perspective by a phylogeographic study of the mesobuthid scorpions across temperate Asia using one mitochondrial and three nuclear genes. Concordant mitochondrial and nuclear gene trees were obtained, which are consistent with species tree inferred using a Bayesian approach. The age of the most recent common ancestor (MRCA) of all the studied scorpions was estimated to be 12.49 Ma (late Middle Miocene); Mesobuthus eupeus diverged from the clade composing Mesobuthus caucasicus and Mesobuthus martensii in early Late Miocene (10.21 Ma); M. martensii diverged from M. caucasicus at 5.53 Ma in Late Miocene. The estimated MRCA ages of M. martensii and the Chinese lineage of M. eupeus were 2.37 and 0.68 Ma, respectively. Central Asia was identified as the ancestral area for the lineage leading to M. martensii and M. caucasicus and the Chinese lineage of M. eupeus. The ancestral habitat of the genus Mesobuthus is likely to have been characterized by an arid environment; a shift towards more humid habitat occurred in the MRCA of M. martensii and a lineage of M. caucasicus, finally leading to the adaptation of M. martensii to humid environment. Our data strongly support the idea that the stepwise intensified aridifications from Mid‐Miocene onwards drove the diversification of mesobuthid scorpions, and suggest that M. martensii and M. eupeus observed today in China originated from an ancestral lineage distributed in Central Asia. Both the colonization and the ensuing evolution of these species in East Asia appear to have been further moulded by Quaternary glaciations.     

A new stem-sperm whale (Cetacea, Odontoceti, Physeteroidea) from the Latest Miocene of Peru

A finely preserved skull with mandible and teeth associated, from the Latest Miocene beds (ca. 6 Ma) of the Pisco Formation, Sud-Sacaco, Peru, represents a new physeteroid genus and species, Acrophyseter deinodon. This moderate size sperm whale is characterized, among others, by: the short rostrum, the mandible distinctly curved upwards, large teeth very close together (12 on each upper tooth row and 13 on each lower tooth row), the lateral margin of the maxilla along the rostrum base much lower than the orbit roof, a wide supracranial basin dorsally overhanging the right orbit and limited to the cranium and a large temporal fossa dorsomedially elevated. A preliminary cladistic analysis provides a phylogenetic position of Acrophyseter nested within the stem-Physeteroidea, more basal than the clade Kogiidae + Physeteridae. The morphology of the oral apparatus and of the temporal fossa suggests that Acrophyseter was able to feed on large preys.     

Protein surface roughness accounts for binding free energy of Plasmepsin II‐ligand complexes

The calculation of absolute binding affinities for protein‐inhibitor complexes remains as one of the main challenges in computational structure‐based ligand design. The present work explored the calculations of surface fractal dimension (as a measure of surface roughness) and the relationship with experimental binding free energies of Plasmepsin II complexes. Plasmepsin II is an attractive target for novel therapeutic compounds to treat malaria. However, the structural flexibility of this enzyme is a drawback when searching for specific inhibitors. Concerning that, we performed separate explicitly solvated molecular dynamics simulations using the available high‐resolution crystal structures of different Plasmepsin II complexes. Molecular dynamics simulations allowed a better approximation to systems dynamics and, therefore, a more reliable estimation of surface roughness. This constitutes a novel approximation in order to obtain more realistic values of fractal dimension, because previous works considered only x‐ray structures. Binding site fractal dimension was calculated considering the ensemble of structures generated at different simulation times. A linear relationship between binding site fractal dimension and experimental binding free energies of the complexes was observed within 20 ns. Previous studies of the subject did not uncover this relationship. Regression model, coined FD model, was built to estimate binding free energies from binding site fractal dimension values. Leave‐one‐out cross‐validation showed that our model reproduced accurately the absolute binding free energies for our training set (R² = 0.76; <|error|> =0.55 kcal/mol; SD_error = 0.19 kcal/mol). The fact that such a simple model may be applied raises some questions that are addressed in the article.     

Phylogeny and diversification of the cloud forest Morpho sulkowskyi group (Lepidoptera,Nymphalidae) in the evolving Andes

The monophyletic Morpho sulkowskyi butterfly group, endemic of Andean cloud forests, was studied to test the respective contributions of Mio‐Pliocene intense uplift period and Pleistocene glacial cycles on Andean biodiversity. We sampled nine taxa covering the whole geographical range of the group. Two mitochondrial and two nuclear genes were analysed using a Bayesian method. We established a dated phylogeny of the group using a relaxed clock method and a wide‐outgroup approach. To discriminate between two hypotheses, we used a biogeographical probabilistic method. Results suggest that the ancestor of the M. sulkowskyi group originated during the Middle–Late Miocene uplift of the Eastern Cordillera in northern Peru. Biogeographical inference suggests that the M. sulkowskyi and Morpho lympharis clades diverged in the northern Peruvian Andes. The subsequent divergences, from the Late Miocene to the Late Pliocene, should have resulted from a dispersal towards the Northern Andes (M. sulkowskyi clade), after the closure of the West Andean Portal separating the Central and Northern Andes, and a southwards dispersal along the Peruvian and Bolivian Eastern Cordilleras (M. lympharis clade). Only a few divergences occurred at the very end of the Pliocene or during the Pleistocene, a period when the more recent uplifts interfered with Pleistocene glacial cycles.     

Discovery of fossil Ayu (<Emphasis Type="Italic">Plecoglossus altivelis</Emphasis>) from the Upper Miocene of Shimane Prefecture,Japan

The Ayu, Plecoglossus altivelis, comprising three subspecies, is the only species in the genus Plecoglossus in the family Osmeridae. Here, we describe for the first time the discovery of fossil specimens of P. altivelis from the Upper Miocene sediment of Matsue, Shimane Prefecture, Japan. The fossil individuals are subadult and have comb teeth in the second and third generations. These fossil specimens are clearly similar to the recent P. altivelis, the former’s meristic characters being included in the three recent subspecies. In terms of numbers of pectoral fin rays and vertebrae, the fossil specimens are more similar to the subspecies Plecoglossus altivelis ryukyuensis than the other subspecies, but they also show different dorsal fin shapes and anal fin ray counts. This discovery of fossil P. altivelis indicates the persistence of the species for at least 10 million years, from the Late Miocene to the present.     

Origine et évolution des hominidés : Toumaï, une confirmation éclatante de la prédiction de Darwin

The earliest known hominid Sahelanthropus tchadensis Brunet et al., 2002, nicknamed Toumaï, from the Late Miocene (7 Ma) of Chad, displays a unique combination of primitive and derived characters which clearly shows that it is probably temporally close to the common ancestor of chimpanzees and humans. The Late Miocene Hominids: Sahelanthropus (Chad), Orrorin (Kenya), Ardipithecus (Ethiopia) are probably the ancestral group of Australopithecins from which the genus Homo appears between 2 and 3 Ma and then for the first time widespread in Eurasia. So, Toumaï seven millions years old confirms in a spectacular way the prediction Charles Darwin made in 1871.     

Reciprocal intraguild predation and predator coexistence

Intraguild predation is a mix of competition and predation and occurs when one species feeds on another species that uses similar resources. Theory predicts that intraguild predation hampers coexistence of species involved, but it is common in nature. It has been suggested that increasing habitat complexity and the presence of alternative food may promote coexistence. Reciprocal intraguild predation limits possibilities for coexistence even further. Habitat complexity and the presence of alternative food are believed to promote coexistence. We investigated this using two species of predatory mites, Iphiseiodes zuluagai and Euseius concordis, by assessing co‐occurrence in the field and on arenas differing in spatial structure in the laboratory. The predators co‐occured on the same plants in the field. In the laboratory, adults of the two mites fed on juveniles of the other species, both in the presence and the absence of a shared food source, showing that the two species are involved in reciprocal intraguild predation. Adults of I. zuluagai also attacked adults of E. concordis. This suggests limited possibilities for coexistence of the two species. Indeed, E. concordis invariably went extinct extremely rapidly on arenas without spatial structure with populations consisting of all stages of the two predators and with a shared resource. Coexistence was prolonged on host plant leaves with extra food sources, but E. concordis still went extinct. On small, intact plants, coexistence of the two species was much longer, and ended with the other species, I. zuluagai, often going extinct. These results suggest that spatial structure and the presence of alternative food increase the coexistence period of intraguild predators.     

Orycteropus (Tubulidentata, Mammalia) from Langebaanweg and Baard's Quarry, Early Pliocene of South Africa

Fossil teeth and bones of aardvarks are relatively common at Langebaanweg, an Early Pliocene site in western Cape Province, South Africa. The remains are compatible in size and most details of morphology to extant Orycteropus afer, and are the earliest fossils attributed to this species. Other Late Miocene to Early Pliocene localities in Africa have yielded smaller species of aardvarks, suggesting that the extant lineage evolved in southern Africa. Morphologically the genus Orycteropus has been remarkably conservative since at least the Early Miocene but it witnessed an overall increase in size through the Neogene. The species O. afer has been morphometrically stable since the Early Pliocene. These observations indicate that the evolutionary process in aardvarks is extremely bradytelic. To cite this article: M. Pickford, C. R. Palevol 4 (2005).     

Late Miocene (Turolian,MN13) squirrels from Moncucco Torinese,NW Italy

Squirrel remains are rare in the fossil record, which hampers a properly documented interpretation of their evolutionary history. The Late Miocene sciurids from Moncucco Torinese (NW Italy) are described based on a diverse tooth assemblage. The data presented herein confirm the presence of Sciurus warthae and Pliopetaurista pliocaenica, and also include the first unquestionable occurrence of Hylopetes hungaricus plus an indeterminate Sciurinae. The report of Sciurus warthae from Moncucco Torinese represents the earliest record of the genus Sciurus in Europe. Moreover, Moncucco Torinese is the first European locality recording the co-occurrence of Sciurini and Pteromyini in the Late Miocene. Comparative analyses support a close relationship between S. warthae and Sciurus vulgaris corroborating the hypothesis of an earliest divergence of the “Sciurus vulgaris clade” in the Late Miocene. Biochronological, paleobiogeographical and paleoecological implications are also discussed.     

Meaningful measures of enamel hypoplasia: Prevalence and comparative intensity of developmental stress

Objectives

Developmental stress causing enamel thinning is an important topic in primate biology. Because taxa differ in growth rates and enamel thickness, the goal is to provide a new method allowing direct comparison of prevalence and salience of enamel defects among samples.

Materials and Methods

Casts of ape teeth spanning the Late Pleistocene to Late Miocene from three site areas of increasing seasonality, equator (Sumatra) to 20° (Vietnam) and 25°N latitude (China), were examined for enamel defects among paleo-orangutans (n = 571, 222, respectively) and Lufengpithecus lufengensis (n = 198). Frequency of affected teeth and number of linear enamel hypoplasia were recorded. Defect dimensions were measured with a confocal microscope. Simple prevalence is compared to weighted prevalence (%), calculated by dividing “number of LEH from specific tooth groups” by “specific tooth sample size”; this quantity divided by “tooth-specific years of imbricational enamel formation.” Defect dimensions are reduced to a dimensionless index termed “enamel deficit ratio” through dividing “daily enamel deficit” by “daily secretion rate.”

Results

Weighted prevalence increases to the North, highlighting latitudinal similarities. In contrast, “enamel deficit ratio,” designed to express comparative severity of developmental stress among samples, was least in the high latitude sample and differed little between paleo-orangutan samples.

Discussion

The actual numbers generated are not as important as efficacy of the proposed methods for other taxa. Developmental stress appears least severe in the high latitude (Lufengpithecus) sample but affects a greater proportion, compared to paleo-orangutans. Regardless of findings, the proposed solutions to improve comparability of disparate samples, yield reasonable results.     

Fractal analysis of the Orce skull sutures

Methods of fractal geometry (Mandelbrot, 1983) are used here to analyse the relative complexity of the sagittal and lambdoid sutures visible in the skull fragment formed by parts of an occipital squame and parietals found in a sealed deposit at the early Lower Pleistocene site of Venta Micena (Orce, Granada, Spain), generally regarded as human bone but occasionally suggested as belonging to an equid. For comparison with the fossil, corresponding sutures of various primates (hominids, pongids and cercopithecids) and two other groups of mammals (equids and ruminants) were analysed using the computer program FRACTAL-D (Slice, 1989) in order to determine their fractal dimensions as a measure of differential sutural design complexity. The results show that the fractal dimension of the Venta Micena skull sutures lies within the range of variation for infant specimens of both modern and Plio-Pleistocene hominids. Sutural complexity in young pongids and cercopithecids overlaps the range of fractal dimensions found in hominids, whereas values obtained from equids and ruminants are significantly greater than those for all the primates analysed here. Therefore, in terms of fractal dimension measures of relative complexity, the sutures preserved in the Venta Micena fossil could not have belonged to an equid (pace Agusti & Moyà-Sola, 1987); rather, its fractal dimension is consistent with the attribution of the fossil to an infant of Homo sp.     

New observations on the Late Miocene–Early Pliocene Lutrinae (Mustelidae: Carnivora,Mammalia) from the Middle Awash,Afar Rift,Ethiopia

New observations on the Late Miocene and Earliest Pliocene mustelids from the Middle Awash of Ethiopia are presented. The Middle Awash study area samples the last six million years of African vertebrate evolutionary history. Its Latest Miocene (Asa Koma Member of the Adu-Asa Formation, 5.54–5.77 Ma) and Earliest Pliocene (Kuseralee and Gawto Members of the Sagantole Formation, 5.2 and 4.85 Ma, respectively) deposits sample a number of large and small carnivore taxa among which mustelids are numerically abundant. Among the known Late Miocene and Early Pliocene mustelid genera, the Middle Awash Late Miocene documents the earliest Mellivora in eastern Africa and its likely first appearance in Africa, a new species of Plesiogulo, and a species of Vishnuonyx. The latter possibly represents the last appearance of this genus in Africa. Torolutra ougandensis is known from both the Late Miocene and Early Pliocene deposits of the Middle Awash. The genus Sivaonyx is represented by at least two species: S. ekecaman and S. aff. S. soriae. Most of the lutrine genera documented in the Middle Awash Late Miocene/Early Pliocene are also documented in contemporaneous sites of eastern Africa. The new observations presented here show that mustelids were more diverse in the Middle Awash Late Miocene and Early Pliocene than previously documented.