Long bone microstructure gives new insights into the life of pachypleurosaurids from the Middle Triassic of Monte San Giorgio,Switzerland/Italy

The long bone microstructure of four pachypleurosaurid taxa from Monte San Giorgio (Switzerland/Italy) was studied. Pachypleurosaurids are secondarily aquatic reptiles that lived during the Middle Triassic in varying marine environments of the Tethys. All four pachypleurosaurids show high compactness values in their long bones based on a thick cortex and a calcified cartilaginous core, which remains in the medullary region throughout the ontogeny. Parts or even the entire embryonic bone layer composed of a mixture of woven-fibered bone tissue and parallel-fibered bone tissue is preserved in both pachypleurosaurid genera. The rest of the cortex consists of lamellar-zonal bone tissue type. Differences in the microstructure of the bones between the pachypleurosaurids are reflected in the occurrence of remodelling processes, which, if present, affect the innermost growth marks of the cortex or the calcified cartilaginous core. Further variation is present in the spacing pattern of the growth cycles, as well as in the degree of vascularisation of the lamellar-zonal bone tissue type. Our data on the microstructure of the long bones support previous studies on morphology and facies distribution, which indicated different habitats and adaptation to a secondary aquatic lifestyle for each pachypleurosaurid taxon. Life history data furthermore reflect different longevities and ages at sexual maturity. The bone histological data of the stratigraphically youngest and oldest pachypleurosaurid species might indicate possible climate-dependant reproductive seasons similar to Recent lacertilian squamates.     

Mikadocephalus gracilirostris n. gen., n. sp., a new ichthyosaur from the Grenzbitumenzone (Anisian-Ladinian) of Monte San Giorgio (Switzerland)

An almost complete skull of a new ichthyosaur from the Middle Triassic Grenzbitumenzone Beds of Monte San Giorgio (Kanton Tessin, Switzerland) represents one of the most complete and best preserved finds of a large Triassic ichthyosaur cranium. Its affinites with other Triassic ichthyosaur taxa are discussed and it is demonstrated to represent a new genus and species,Mikadocephalus gracilirostris, which does not fit into any of the currently recognized families of Triassic ichthyosaurs. Remarkable similarities in cranial structure exit to postTriassic ichthyosaurs, with which a number of important apomorphies are shared.     

Taxonomic significance of biométric characters and the consequences for classification and biostratigraphy, exemplified through moussoneliform daonellas (Daonella, Bivalvia; Triassic)

The middle Triassic bivalve genusDaonella Mojsisovics, 1874 would be an ideal guide fossil (worldwide distribution, high evolutionary rates); however, the classification of daonellas is difficult. Most daonellas lack distinct characters and have a wide range of morphologic variation. A new taxonomic attempt was necessary to get a coherentDaonella classification. The new taxonomic concept proposed herein consists of four steps. (1) Biometric characters are defined unambiguously, based on landmarks and pseudolandmarks. (2) The taxonomic significance of biometric characters is analyzed. Several characters depend significantly on growth stage of specimens or on the lithology of fossil localities. In addition, the intercorrelations between biometric characters are studied to exclude weighting in some morphologic features. The study of the taxonomic significance of biometric characters shows, how important a careful selection of taxonomic characters is. (3) Most daonellas are not determinable by univariate or bivariate methods. The principle component analysis (PCA) is applied to detect groups of similar holomorphs in multivariate space. (4) The interpreted groups from PCA are applied to a canonical discriminant function analysis. This analysis helps in deciding whether two groups are sufficiently distinct to be separated as different species. This new taxonomic concept was applied to moussoneliform daonellas of the middle Triassic from the Southern Alps. Three different forms were used for the study herein:Daonella moussoni (Mérian, 1853) from the Varenna Limestone (Perledo-Varenna Formation, Grigna, Italy),D. aff.moussoni (Mérian)Rieber, 1969 from the lower Meride Limestone (Monte San Giorgio, Switzerland) andD. pseudomoussoni Rieber, 1969 from the Grenzbitumenzone (Monte San Giorgio, Switzerland). The new classification shows thatD. pseudomoussoni is a separate species and distinguishable fromD. moussoni andD. aff.moussoni, which belong to the same species. A coherent systematics ofDaonella becomes possible on the basis of newly defined taxonomic characters. Daonellas can now be used as relevant guide fossils for the middle Triassic. The presented example demonstrates a biostratigraphic correlation between the lower Meride Limestone at Monte San Giorgio and the Varenna Limestone at Grigna. The stratigraphic distribution ofD. moussoni can be assigned to thegredleri Zone.     

It's all relative: allometry and variation in the baculum (os penis) of the harp seal, Pagophilus groenlandicus (Carnivora: Phocidae)

We compared allometry and variation in the baculum (os penis), mandible, and humerus of the harp seal, Pagophilus groenlandicus. This species is presumed to have a promiscuous mating system in which choice of mate by females during intromission with different males is likely. The baculum is large and grows throughout life so may be an honest indicator of males' quality (size) or viability (age). We predicted that bacular size would exhibit stronger allometry relative to body size than mandibles or humeri. The baculum is less functionally (mechanically) constrained than mandibles or humeri so we also predicted it would be more variable, though less variable than sexually selected traits which do not function as honest indicators. Our sample (N=67 seals) represented broad ranges of size and age (0–35 yr) so we compared variation using residuals from allometric regressions of skeletal measurements on body length. Bacular size was isometric to body length until ∼ 137 cm (when some seals enter puberty) in body length then was highly positively allometric; mandibular and humeral size were negatively allometric to body length throughout growth. Bacula were more variable than mandibles or humeri. Bacular size in large specimens (>137 cm in body length) was related strongly to body length and weakly to age. We interpret bacular size to be an uncheatable honest indicator of male quality and viability. High bacular variation conforms with theoretical predictions of females' asymmetrical choice of mate and choice of extremes, and may reflect corresponding anatomical variation among females. Some bacular variation may also result incidentally from positive allometry coupled with lifelong bacular growth, which can amplify early differences between reproductive and somatic growth, enabled by weak selection on bacular form in relation to function.     

Two new species of Saurichthys (Actinopterygii: Saurichthyidae) from the Middle Triassic of Monte San Giorgio,Switzerland, with implications for character evolution in the genus

Saurichthys, characterized by a long slender body and an elongated rostrum, is one of the most iconic genera of Late Paleozoic–Early Mesozoic fishes. The genus was particularly speciose in the Triassic, with a global distribution in both marine and freshwater habitats. Here, we describe two new species from the Middle Triassic Besano Formation of Monte San Giorgio, Switzerland, Saurichthys breviabdominalis sp. nov. and Saurichthys rieppeli sp. nov. S. breviabdominalis is characterized by a proportionately long operculum, short abdominal region and rib‐like mid‐lateral scales, whereas S. rieppeli is divergent from other Middle Triassic saurichthyids in the block‐like haemal arches, fringing fulcra on the pelvic and unpaired fins, and reduction of the squamation to a single row in the abdominal region. Phylogenetic analysis places S. rieppeli in a basal position relative to congeners from the Alpine Triassic, and supports previous hypotheses regarding the convergent evolution of reduced squamation within saurichthyids. S. breviabdominalis forms a monophyletic group with species from the same locality, suggesting divergence in sympatry. This finding has implications for our understanding of disparity and character evolution in saurichthyid fishes, as well as ecomorphological divergence and resource partitioning between closely related fishes in Triassic marine ecosystems. © 2015 The Linnean Society of London     

Scaling of the appendicular skeleton of the giraffe (Giraffa camelopardalis)

Giraffes have remarkably long and slender limb bones, but it is unknown how they grow with regard to body mass, sex, and neck length. In this study, we measured the length, mediolateral (ML) diameter, craniocaudal (CC) diameter and circumference of the humerus, radius, metacarpus, femur, tibia, and metatarsus in 10 fetuses, 21 females, and 23 males of known body masses. Allometric exponents were determined and compared. We found the average bone length increased from 340 ± 50 mm at birth to 700 ± 120 mm at maturity, while average diameters increased from 30 ± 3 to 70 ± 11 mm. Fetal bones increased with positive allometry in length (relative to body mass) and in diameter (relative to body mass and length). In postnatal giraffes bone lengths and diameters increased iso‐ or negatively allometric relative to increases in body mass, except for the humerus CC diameter which increased with positive allometry. Humerus circumference also increased with positive allometry, that of the radius and tibia isometrically and the femur and metapodials with negative allometry. Relative to increases in bone length, both the humerus and femur widened with positive allometry. In the distal limb bones, ML diameters increased isometrically (radius, metacarpus) or positively allometric (tibia, metatarsus) while the corresponding CC widths increased with negative allometry and isometrically, respectively. Except for the humerus and femur, exponents were not significantly different between corresponding front and hind limb segments. We concluded that the patterns of bone growth in males and females are identical. In fetuses, the growth of the appendicular skeleton is faster than it is after birth which is a pattern opposite to that reported for the neck. Allometric exponents seemed unremarkable compared to the few species described previously, and pointed to the importance of neck elongation rather than leg elongation during evolution. Nevertheless, the front limb bones and especially the humerus may show adaptation to behaviors such as drinking posture. J. Morphol. 276:503–516, 2015. © 2014 Wiley Periodicals, Inc.     

A sequence stratigraphic approach to a Middle Triassic shelf-slope complex of the Ligurian Alps (Ligurian Briançonnais,Monte Carmo-Rialto unit,Italy)

The Mesozoic sedimentary cover belonging to the Monte Carmo-Rialto unit of the Ligurian Briançonnais domain is composed of Scythian clastics and Anisian to Carnian carbonate rocks over 300 m thick. This paper focuses on the stratigraphy of this carbonate complex, its environmental significance, and its evolution in light of dynamic stratigraphy. Our facies analysis of limestones and dolomites of the Triassic complex allowed us to reconstruct an environmental model. Data support a distally steepened carbonate ramp of Anisian age evolving to a more diversified Ladinian platform with an oolitic sand-bar belt separating the lagoon from the slope. The Monte Carmo-Rialto slope facies are the only witnesses of deep sedimentation in the Triassic terrains of the Ligurian Briançonnais domain, otherwise represented by shallow-water carbonate deposits. On the basis of facies succession, we have identified nine medium-scale cycles (third-order sequences) in the study area, comparable to those evidenced in the Briançonnais s.s. domain by the French authors. Small-scale cycles analysis evidenced mainly shallowing-upward trends in the examined sequences; although a few evidences of transgression-related deposits (deepening upward cycles) have been found at the base three sequences, they have been mostly obliterated by dolomitization and masked by local tectonics. For this reason, we can undoubtedly distinguish only the part of each sequence belonging to HST, while the TST, though present, still remains a partition that cannot be precisely characterized. In the same way, LSTs are not present in the Monte Carmo-Rialto unit, due to the original relative landward position of the examined area. Sequence stratigraphy analysis indicates different long-term dynamics for the two evolutionary stages of the Triassic Ligurian platform: a general landward backstepping to moderate progradation during the Early Anisian and true progradation during the latest Anisian and Ladinian. In addition, a good fit with the sequences proposed by the SEPM chart has been found, indicating a correspondence for the third-order sequences of the Middle Triassic.     

Facies and biota of Anisian to Carnian carbonate platforms in the Northern Calcareous Alps (Tyrol and Bavaria)

Summary During the Middle and early Late Triassic carbonate ramps and rimmed platforms developed at the northwestern margin of the Tethys ocean. In the Northern Calcareous Alps, Anisian stacked homoclinal ramps evolved through a transitional stage with distally steepened ramps to huge rimmed platforms of Late Ladinian to Early Carnian age. Middle Triassic to early Late Triassic facies and biota of basin, slope and platform depositional systems are described. Special emphasis is given to foraminifers, sponges, microproblematic organisms and algae. The Ladinian to early Carnian reef associations are characterized by the abundance of segmented sponges, microproblematica, biogenic crusts and synsedimentary cements. Among the foraminifers, recifal forms likeHydrania dulloi andCucurbita infundibuliformis (Carnian in age) are reported from the Northern Calcareous Alps for the first time. Some sphinctozoid sponges likeParavesicocaulis concentricus were known until now only from the Hungarian and Russian Triassic.     

Re‐evaluation of the ontogeny and reproductive biology of the Triassic fish Saurichthys (Actinopterygii,Saurichthyidae)

Viviparity has evolved independently at least 12 times in ray‐finned fishes. However, the fossil record of actinopterygian viviparity is poor, with only two documented occurrences. Both of these are from the non‐teleost actinopterygian Saurichthys, and include S. curionii and S. macrocephalus from the Middle Triassic Meride Limestone (Monte San Giorgio, Switzerland). Here, we present new data on the reproductive biology of these species, giving unprecedented insights into their life‐history. Based on positional and preservational criteria, six specimens were identified as unambiguously gravid. Embryos were positioned dorsal to the gastrointestinal tract, parallel to the axial skeleton and to each other, in the posterior two‐thirds of the abdominal region. A minimum of 16 embryos are preserved in the most fecund females and, based on the largest preserved embryos and smallest preserved neonates, birth must have occurred at 7–12% of maternal fork length. Embryonic crania and teeth are relatively well‐ossified, however ossification of the parietal region is delayed. In the postcranium, the median scale rows and lepidotrichia are ossified, but not the lateral scale rows. Ossified squamation and gradual allometric growth suggests that neonates did not undergo metamorphosis and were relatively precocial. When considered in a phylogenetic context, neither live birth nor internal fertilization appears to represent the primitive state for saurichthyid fishes.     

Taxonomical and palaeoecological investigations of the chondrichthyan and osteichthyan fish remains from the Middle-Late Triassic deposits of the Villány Hills (Southern Hungary)

Two sites in the Villány Hills, Hungary, have yielded rich fish assemblages from Middle to Late Triassic shallow marine deposits. The collected material comes from the Ladinian Templomhegy Dolomite Member and from the Carnian Mészhegy Sandstone Formation. The ichthyofauna is composed of both chondrichthyans (Hybodontidae indet., Palaeobates angustissimus, ‘Polyacrodus’ sp., ?Lissodus sp.) and osteichthyans (Gyrolepis sp., Birgeria sp., and further indeterminate actinopterygians). Despite the large sample size, no remains of neoselachians have been found. The Ladinian Templomhegy Dolomite is dominated by durophagous hybodontiforms (Palaeobates angustissimus, ?Lissodus sp.), but the piscivorous hybodontid and the generalist ‘Polyacrodus’ sp. are missing, while in the fish fauna collected from the Carnian Mészhegy Formation indeterminate piscivorous hybodontids are the most common elements and durophagous forms are much less abundant. The dominance of piscivorous hybodontids in the Carnian Mészhegy Sandstone could be related to the global decrease of diversity of marine fish-eating reptiles (e.g., nothosaurs) or to a change of paleoenvironmental conditions. The present study improves our knowledge on the poorly known Triassic vertebrate faunas of the Tisza Mega-unit, which formed a segment of the passive Neotethys margin of the European Plate and shows an important example of a potential vertebrate faunal shift during the Middle to Late Triassic.     

Floral Assemblages and Patterns of Insect Herbivory during the Permian to Triassic of Northeastern Italy

To discern the effect of the end-Permian (P-Tr) ecological crisis on land, interactions between plants and their insect herbivores were examined for four time intervals containing ten major floras from the Dolomites of northeastern Italy during a Permian–Triassic interval. These floras are: (i) the Kungurian Tregiovo Flora; (ii) the Wuchiapingian Bletterbach Flora; (iii) three Anisian floras; and (iv) five Ladinian floras. Derived plant–insect interactional data is based on 4242 plant specimens (1995 Permian, 2247 Triassic) allocated to 86 fossil taxa (32 Permian, 56 Triassic), representing lycophytes, sphenophytes, pteridophytes, pteridosperms, ginkgophytes, cycadophytes and coniferophytes from 37 million-year interval (23 m.yr. Permian, 14 m.yr. Triassic). Major Kungurian herbivorized plants were unaffiliated taxa and pteridosperms; later during the Wuchiapingian cycadophytes were predominantly consumed. For the Anisian, pteridosperms and cycadophytes were preferentially consumed, and subordinately pteridophytes, lycophytes and conifers. Ladinian herbivores overwhelming targeted pteridosperms and subordinately cycadophytes and conifers. Throughout the interval the percentage of insect-damaged leaves in bulk floras, as a proportion of total leaves examined, varied from 3.6% for the Kungurian (N = 464 leaves), 1.95% for the Wuchiapingian (N = 1531), 11.65% for the pooled Anisian (N = 1324), to 10.72% for the pooled Ladinian (N = 923), documenting an overall herbivory rise. The percentage of generalized consumption, equivalent to external foliage feeding, consistently exceeded the level of specialized consumption from internal feeding. Generalized damage ranged from 73.6% (Kungurian) of all feeding damage, to 79% (Wuchiapingian), 65.5% (pooled Anisian) and 73.2% (pooled Ladinian). Generalized-to-specialized ratios show minimal change through the interval, although herbivore component community structure (herbivore species feeding on a single plant-host species) increasingly was partitioned from Wuchiapingian to Ladinian. The Paleozoic plant with the richest herbivore component community, the coniferophyte Pseudovoltzia liebeana, harbored four damage types (DTs), whereas its Triassic parallel, the pteridosperm Scytophyllum bergeri housed 11 DTs, almost four times that of P. liebeana. Although generalized DTs of P. liebeana were similar to S. bergeri, there was expansion of Triassic specialized feeding types, including leaf mining. Permian–Triassic generalized herbivory remained relatively constant, but specialized herbivores more finely partitioned plant-host tissues via new feeding modes, especially in the Anisian. Insect-damaged leaf percentages for Dolomites Kungurian and Wuchiapingian floras were similar to those of lower Permian, north-central Texas, but only one-third that of southeastern Brazil. Global herbivore patterns for Early Triassic plant–insect interactions remain unknown.     

Geological significance of the discovery of Middle Triassic (Ladinian) radiolarians from the Hong Hoi Formation of the Lampang Group,Sukhothai Zone,northern Thailand

This paper presents a newly found fauna of Ladinian (Middle Triassic) radiolarians from the Hong Hoi Formation of the Lampang Group, Sukhothai Zone, northern Thailand. The Ladinian age determined by the radiolarian fauna concords with the age previously determined by mollusks. The study section consists of intercalated sandstone and shale in the lower part and a thick conglomerate in the upper part. The radiolarian-bearing siliceous beds are intercalated within the lower unit. The lithic sandstone of the lower unit consists mainly of volcanic rock fragments, quartz, and feldspar, whereas the thickly bedded conglomerate of the upper unit is characterized by abundant gravel-sized clasts of volcanic rock and limestone. These lithic features of the study section suggest that during deposition the Hong Hoi Formation was located near a supply of volcanic materials. A forearc basin close to the Sukhothai Arc would be the most suitable environment for the deposition. The Ladinian age determined by the radiolarian fauna supports the occurrence of intensive volcanic activity within the Sukhothai Arc during the Middle Triassic.     

Paraplacodus and the phylogeny of the Placodontia (Reptilia: Sauropterygia)

The skeletal anatomy of Paraplacodus broilii Peyer from the Grenzbitumen-horizon (Anisian-Ladinian boundary) of Monte San Giorgio (Switzerland) is described and compared with that of other placodonts. Paraplacodus is found to share a number of potential synapomorphies with Placodus which could potentially corroborate the monophyly of the Placodontoidea, but Placodus also shares an number of potential synapomorphies with the armored placodonts (Cyamodontoidea) which are absent in Paraplacodus. Parsimony analysis rejects the monophyly of the Placodontoidea, and places Paraplacodus at the root of the placodont tree, as the sister-taxon of all the other representatives of the clade. This correlates with a configuration of the temporal region of the skull that suggest the loss of the lower temporal arch in a diapsid skull. The loss of the lower temporal arch is therefore recognized as a sauropterygian synapomorphy, and might even be a lepidosauromorph synapomorphy.     

中国陕西中三叠世中蝎蛉科三新属新种(昆虫纲,长翅目)

记述中蝎蛉科Mesopanorpodidae 4属4种,其中包括3新属4新种：Longifurcula hejiafangensi gen.et sp.nov,Ladinochorista lata gen.et sp.nov.,Allochorista erdosensis gen.et sp.nov.,Itaphlebia tongchuanensis sp.nov..化石标本采自陕西铜川中三叠统铜川组下段上部的灰绿色泥岩和页岩.这些新属、种系陕西昆虫群铜川昆虫组合的新成员（属陕西动物群、陕西生物群的一个类别）.根据前人的植物、孢子、双壳、叶肢介、介形虫、昆虫化石等的研究结果,铜川组的时代划归中三叠世,相当于欧洲中三叠世拉丁尼期（Ladinian stage）.     

Comment on Phylogenetic analyses with four new Cretaceous bristletails reveal inter-relationships of Archaeognatha and Gondwana origin of Meinertellidae

Archaeognatha (Machilida plus †Monura), with ~450 extant species worldwide, represent a key lineage for the understanding of insect evolution. In a recently published phylogenetic analysis on Archaeognatha, based on a total-evidence approach, a Machilidae key fossil (Gigamachilis triassicus) from the Middle Triassic Lagerstätte of Monte San Giorgio and a representative of †Monura (Dasyleptus triassicus) from the same fossiliferous site, were not considered. In the present study, the phylogenetic analyses performed by Zhang et al. (2018) using maximum parsimony and Bayesian inference were repeated including these two key taxa; in addition, Tricholepidion gertschi and Lepisma saccharina were included as representatives of Zygentoma. In the total-evidence Bayesian analysis, †Monura was found to be paraphyletic, whereas in the maximum parsimony analysis, performed only on morphological characters, they were monophyletic. Moreover, the attribution of G. triassicus to Machilidae is now confirmed by these analyses, indicating that members of the extant group of Machilidae were already present in the Middle Triassic, at least ~100 Myr before the previous estimate.     

贵州中三叠世安顺龙(爬行纲:海龙目)一新种

中国的海龙类以前仅发现于法郎组瓦窑段,包括安顺龙和新铺龙两属。最近在法郎组竹杆坡段也发现了海龙化石(完整的骨架,中国科学院古脊椎动物与古人类研究所标本编号V 13782),被归入安顺龙属。标本产自贵州兴义乌沙,时代是中三叠世拉丁期。此标本小于黄果树安顺龙两个已知标本,但是根据以下特征判断应该是个成年个体：头骨相对较小,许多骨缝愈合,髓椎线(neurocentral suture)封闭,肱骨上各种结构已经显著分化,掌骨和跗骨骨化程度高。根据以下特征建立一新种,乌沙安顺龙(Anshunsaurus wushaensis sp．nov．)：头骨相对较小(相对肩臼至髋臼长度);轭骨后突短,向后延伸不超过下颞孔之半;后背区的神经棘高度小于宽度,其背缘有垂向沟、脊;间锁骨十字形,前突基部宽;肱骨外髁沟明显,内髁很发育,在内腹侧有脊但无孔;第五掌骨比第四掌骨稍长;第四指失去一个指节;髂骨板向后背向展开;7个跗骨骨化。     

陕西中三叠世中蝎蛉科两新属和新种(昆虫纲,长翅目)

记述中蝎蛉科Mesopanorpodidae 2新属、新种：Triassochoristites jinsuoguanensis gen．et sp．nov．,Forcinerva tongchuanensis gen．et sp．nov．化石标本采自陕西铜川中三叠世铜川组下段上部的灰绿色泥岩和页岩。这些新属、种系陕西昆虫群(陕西生物群的一个类别)铜川昆虫组合的新成员。铜川组的时代相当于欧洲拉丁尼期(Ladinian Stagc)。     

Comparative limb bone scaling in turtles: Phylogenetic transitions with changes in functional demands?

Several terrestrial vertebrate clades include lineages that have evolved nearly exclusive use of aquatic habitats. In many cases, such transitions are associated with the evolution of flattened limbs that are used to swim via dorsoventral flapping. Such changes in shape may have been facilitated by changes in limb bone loading in novel aquatic environments. Studies on limb bone loading in turtles found that torsion is high relative to bending loads on land, but reduced compared to bending during aquatic rowing. Release from torsion among rowers could have facilitated the evolution of hydrodynamically advantageous flattened limbs among aquatic species. Because rowing is regarded as an intermediate locomotor stage between walking and flapping, rowing species might show limb bone flattening intermediate between the tubular shapes of walkers and the flattened shapes of flappers. We collected measurements of humeri and femora from specimens representing four functionally divergent turtle clades: sea turtles (marine flappers), softshells (specialized freshwater rowers), emydids (generalist semiaquatic rowers), and tortoises (terrestrial walkers). Patterns of limb bone scaling with size were compared across lineages using phylogenetic comparative methods. Although rowing taxa did not show the intermediate scaling patterns we predicted, our data provide other functional insights. For example, flattening of sea turtle humeri was associated with positive allometry (relative to body mass) for the limb bone diameter perpendicular to the flexion-extension plane of the elbow. Moreover, softshell limb bones exhibit positive allometry of femoral diameters relative to body mass, potentially helping them maintain their typical benthic position in water by providing additional weight to compensate for shell reduction. Tortoise limb bones showed positive allometry of diameters, as well as long humeri, relative to body mass, potentially reflecting specializations for resisting loads associated with digging. Overall, scaling patterns of many turtle lineages appear to correlate with distinctive behaviors or locomotor habits.     

Anisian (middle triassic) buildups of the Northern Dolomites (Italy): The recovery of reef communities after the permian/triassic crisis

Summary After the end-Permian crisis and a global ‘reef gap’ in the early Triassic, reefs appeared again during the early Middle Triassic. Records of Anisian reefs are rare in the Tethys as well as in non-Tethyan regions. Most Anisian reefs are known from the western part of the Tethys but there are only very few studies focused on biota, facies types and the paleogeographical situation of these reefs. From the eastern part of the Tethys, Anisian reefs, reefal buildups or potential reef-building organisms have been reported from different regions of southern China. Most of the Anisian reefs known from western and central Europe as well as from southern China seem to be of middle and late Pelsonian age. The study area is situated in the northern Dolomites (South Tyrol, Italy) southeast of Bruneck (Brunico). It comprises the area between Olang (Valdaora) and Prags (Braies). The study is based on detailed investigations of the regional geology, stratigraphy and lithofacies (R. Zühlke, T. Bechst?dt) as well as on a comprehensive inventory of Anisian reef organisms (B. Senowbari-Daryan, E. Flügel). These data are used in the discussion of the controls on the recovery of reefs during the early Middle Triassic. Most late Anisian reef carbonates studied are represented by allochthonous talus reef blocks of cubicmeter size. Small biostromal autochthonous mounds are extremely rare (Piz da Peres). The reef mounds as well as most of the reef blocks occur within the middle to late Pelsonian Recoaro Formation. They were formed on the middle reaches of carbonate ramps in subtidal depths, slightly above the storm wave base with only moderate water energy. Most lithotypes observed in the reef blocks correspond to sponge and/or algal bafflestones. Low-growing sessile organisms (Olangocoelia (sponge, alga?), sphinctozoan sponges, bryozoans, soleno-poracean algae, corals) and encrusting epibionts (sponges, porostromate algae, cyanophycean crusts, foraminifera, worms, microproblematica) created low cm-sized biogenic structures (bioconstructions) which baffled and bound sediment. Organic framework was only of minor importance; it is restricted to theOlangocoelia lithotype. Framework porosity was small in these reef mounds. Submarine carbonate cements, therefore, are only of minor importance s compared with Permian or Ladinian reefs. The relatively high number of lithotypes encountered in the reef blocks indicates a high biofacies diversity. Regarding the relative frequency, the diverse biota consist in descending order ofOlangocoelia, sponges (sphinctozoans, inozoans, siliceous sponges), bryozoans, porostromate algae and worm tubes. The sphinctozoans are characterized by small, mostly incrusting forms. The numerical diversity (species richness) is low compared with late Permian or Ladinian and late Triassic sphinctozoan faunas occurring within reefs. Following the sponges, monospecific bryozoans (Reptonoditrypa cautica Sch?fer & Fois) are the most common organisms in the reef limestones. Porostromate algae were restricted to areas within the bioconstructions not inhabited by sponges. The low-diverse corals had no importance in the construction of an organic framework. Surprisingly, microbial crusts are rare or even lacking in the investigated Anisian bioconstructions. This is in contrast to late Permian and Ladinian as well as Carnian reefs which are characterized by the abundance of specific organic crusts. The same comes true for‘Tubiphytes’ which is a common constituent in Permian, Ladinian and Carnian reef carbonates but is very rare in the Anisian of the Olang Dolomites. Instead of‘Tubiphytes’ different kinds of worm tubes (spirorbid tubes, Mg-calcitic tubes and agglutinated tubes) were of importance as epifaunal elements. Macrobial encrustations consisting of characteristic successions of sponges, bryozoans, algae, worm tubes and microproblematica seem to be of greater quantitative importance than in Ladinian reefs. Destruction of organic skeletons (predominantly of bryozoans) by macroborers (cirripedia?) is a common feature. The Anisian reef organisms are distinctly different from late Permian and from most Ladinian reef-builders. No Permian Lazarus taxa have been found. New taxa: Sphinctozoan sponges—Celyphia? minima n.sp.,Thaumastocoelia dolomitica n. sp.,Deningeria tenuireticulata n. sp.,Deningeria crassireticulata n. sp.,Anisothalamia minima n.g. n.sp., Inozoan sponges-Meandrostia triassica n.sp. Microproblematica-Anisocellula fecunda n.g. n.sp., Porostromate alga-Brandneria dolomitica n.g. n.sp. Most of our data are in agreement with the model described byFois & Gaetani (1984) for the recovery of reef-building communities during the Ansian but the biotic diversity seems to be considerably higher than previously assumed. Anisian deposition and the formation of the reef mounds within the Pelsonian Recoaro Formation of the Dolomites were controlled by the combined effects of synsedimentary tectonics and eustatic changes in sea-level. During several time intervals, especially the early Anisian (northern and western Dolomites: tectonic uplift), the early Pelsonian (eastern Dolomites: drowning) and the late Illyrian (wide parts of the Dolomites: uplift and drowning), the sedimentation was predominantly controlled by regionally different tectonic subsidence rates. The amount of terrigenous clastic input associated with synsedimentary tectonics (tectonic uplift of hinterlands) had a major influence on carbonate deposition and reef development. The re-appearance of reef environments in the Olang Dolomites was controlled by a combination of regional and global factors (paleogeographic situation: development of carbonate ramps; decreasing subsidence of horst blocks; reduced terrigenous input; moderate rise in sea-level).