Empirical 3D model of the conch of the Middle Jurassic ammonite microconch Normannites: its buoyancy,the physical effects of its mature modifications and speculations on their function

A 3D model of the Middle Jurassic ammonoid Normannites with an apertural modification from Thürnen, Switzerland, was constructed using physical–optical tomography. It was tested to determine whether the formation of the apertural modification affected shell orientation, to estimate buoyancy regulation and to reconstruct the mode of life of this ammonoid. No drastic postural changes occurred between the 3D models that excluded and included lappets, suggesting that the lappets were not formed to change the syn vivo shell orientation and, in turn, locomotion. We speculate that these adult shell modifications served to protect the soft parts during the reproduction period. Buoyancy calculations based on the model assume that ammonoids were positively buoyant when the phragmocone was devoid of liquid. When 31% of the entire phragmocone was filled with liquid, the living animal would have reached neutral buoyancy in contrast to 27% of cameral liquid filling when the weight of the aptychi is included. Provided that smaller ammonoids had more cameral liquid than bigger ammonoids, such as the modern Nautilus, Normannites examined in this study would have been able to maintain neutral buoyancy and might have had a demersal, nektobenthic or nektonic habitat somewhere in the water column.     

Soft‐tissue imprints in fossil and Recent cephalopod septa and septum formation

Several soft‐tissue imprints and attachment sites have been discovered on the inside of the shell wall and on the apertural side of the septum of various fossil and Recent ectocochleate cephalopods. In addition to the scars of the cephalic retractors, steinkerns of the body chambers of bactritoids and some ammonoids from the Moroccan and the German Emsian (Early Devonian) display various kinds of striations; some of these striations are restricted to the mural part of the septum, some start at the suture and terminate at the anterior limit of the annular elevation. Several of these features were also discovered in specimens of Mesozoic and Recent nautilids. These structures are here interpreted as imprints of muscle fibre bundles of the posterior and especially the septal mantle, blood vessels as well as the septal furrow. Most of these structures were not found in ammonoids younger than Middle Devonian. We suggest that newly formed, not yet mineralized (or only slightly), septa were more tightly stayed between the more numerous lobes and saddles in more strongly folded septa of more derived ammonoids and that the higher tension in these septa did not permit soft‐parts to leave imprints on the organic preseptum. It is conceivable that this permitted more derived ammonoids to replace the chamber liquid faster by gas and consequently, new chambers could be used earlier than in other ectocochleate cephalopods, perhaps this process began even prior to mineralization. This would have allowed faster growth rates in derived ammonoids.     

Post-pathological keel-loss compensation in ammonoid growth

Among the various pathologies documented in ammonoids, impairs affecting the apertural margin may have long-lasting sequelae on subsequent shell geometry. An interesting healing pattern, known as sculptural compensation, led to the permanent replacement of an ornament by adjacent sculptural elements. Moreover, in several ventrally impaired individuals the symmetry was preserved. Those developed annular ribs in place of any previous ventral ornamentation (keel, sulcus or smooth area). This phenomenon is known from diverse ammonite families. Monestieria resouchei (Monestier 1931), type species of 'Monestieriinae' Sapunov 1965, displays exactly that type of annularly-ribbed morphology and has been shown to be otherwise similar to species of Grammoceratinae Buckman 1904 occurring in the same beds, thus corroborating its pathological nature and leading to the rejection of that taxon. Now, keel absence in Praehaploceras Monestier 1931 and Buckmanites Guex 1973 cannot be explained by the same process as they do not have annular ribs. Moreover, the absence of any clue of malformation, their relative frequency and specific characteristics exclude the previously suggested synonymies with Pseudolioceras Buckman 1889 as equivalent pathological forms. In consequence, their rehabilitation is herein proposed. They should be included within Harpoceratinae Neumayr 1875.     

Bemerkungen zur Systematik der Lituolidae

In his recent publication onLituola grandis,Joseph H. Ziegler also discusses the classification of the Lituolidae in general. Since such evident morphological criteria of the test as type of coiling and apertural character are held to be of no generic value, some ofZiegler's conclusions differ considerably from those of the author and others. Attention is drawn to all the aspects concerning the morphological features of the group which should be taken into account for taxonomic purposes. Some misinterpretations and errors are pointed out and a supplementary list of publications dealing with lituolid Foraminifera is given.     

Paedomorphosis of ammonoids as a result of sealevel fluctuations in the Late Devonian Wocklumeria Stufe

A remarkable diversification of several independent ammonoid lineages with high evolutionary rates occurred in the Late Devonian Wocklumeria Stufe. Many speciation events led to paedomorphic ammonoids that display a striking range of conch shapes, sculpture, and ornamentation. In the goniatite family Prionoceratidae, the transition from normal Mimimitoceras species to paedomorphic Balvia species provides an example of rapid size decrease combined with an early character developmental offset arising from progenesis. Adults of early Balvia species largely have the preadult ancestral morphology of Mimimitoceras , but later evolving species acquire distinct conch and ornamentation types. Progenetic ammonoid species also appeared within the clymeniid family Kosmoclymeniidae and probably in the Glatzielliidae. In the clymeniid family Parawocklumeriidae, evolution is characterized by the extension of tri-segmented and triangularly coiled whorls found only in juveniles of earlier species, to the adults of later species. This is interpreted as resulting from neoteny. The distribution of paedomorphic ammonoids in the Late Devonian Wocklumeria Stufe is closely correlated to relative sealevel changes. The regressive trend in the lower two-thirds of the Wocklumeria Stufe is interpreted as the cause of a diversification of the pelagic habitat during unstable conditions, and as an extrinsic factor inducing heterochronic change. Some ammonoids reacted by rapid maturation and faster reproductive rates, giving the opportunity to exploit a wider range of niches. The apparent consequence was the formation of several allopatric species. □ Ammonoidea, Late Devonian, evolution, heterochrony, sealevel changes.     

The phylogeny of Hildoceratidae (Cephalopoda,Ammonitida) resolved by an integrated coding scheme of the conch

Ammonite phylogeny has mainly been established based on a stratigraphic approach, with cladistics underconsidered. The main arguments against the use of cladistics are the supposed large amount of homoplasy and the small number of characters. Resolving the phylogeny of the Hildoceratidae (Early Jurassic) is especially challenging because of its large diversity and disparity. Many forms that have not been determined as closely related in previous studies exhibit very similar shapes. Moreover, some groups are morphologically very different, adding difficulties to building a unique coding scheme at a low taxonomic resolution (i.e. species). Here we propose an integrated coding scheme of the peristome shape and the ornamentation, allowing an increased level of comparison. The shape of the peristome is used as a new reference to locate ornamental features and propose new homology hypotheses. In total, 105 taxa have been analysed for 47 characters. We code continuous characters by their means and ranges ± one standard deviation. We test two weighting schemes: equal weights standardized by unit range and implied weighting with several concavity constants. This work has led to redefinition of the phylogenetic inclusivenesses of all the hildoceratid subfamilies. The new coding scheme based on peristome shapes provides the fewest homoplastic characters. The schemes appear promising to improve phylogenetic analyses in ammonoids as well as molluscs as a whole by creating a general coding framework.     

Untangling phylogenetic,geometric and ornamental imprints on Early Triassic ammonoid biogeography: a similarity‐distance decay study

Ammonoids are diverse and widespread fossil, externally shelled cephalopods that flourished for more than 300 Myr before their total extinction 65 Ma ago. In spite of two centuries of intensive scientific studies, their mode(s) of life and long‐distance dispersal abilities remain poorly known. Here, we address this by focusing on the latitudinal distribution of Early Triassic (approximately 250 Myr) ammonoids through similarity‐distance decay analyses. We examine and compare rates of similarity‐distance decay between various groups with respect to systematics, shell geometry and ornamentation to untangle phylogenetic, geometric and ornamental imprints on the observed biogeographical pattern. Our data do not support any phylogenetic and shell ornamentation influence, but rather demonstrate the significant effect of (sub‐)adult shell geometry on the similarity–distance decay: most evolute morphs tend to have been more endemic than most involute forms. This contrasts with the classic hypothesis that long‐distance ammonoid dispersal mainly occurred during the earliest planktonic stages, and thus that (sub‐)adult morphological characteristics should not constrain large‐scale biogeographical patterns of ammonoids. Although direct control by Sea Surface Temperature can be discarded, this result may indicate that at least some adult Triassic ammonoid morphs were skilled active swimmers capable of achieving long‐distance migration, as observed for some present‐day coleoid cephalopods. □Ammonoid, dispersal, similarity‐distance decay, morphology, phylogeny, biogeography, Triassic.     

Developmental evolution of sexual ornamentation: model and a test of feather growth and pigmentation

A tremendous diversity of avian color displays has stimulatednumerous studies of natural and sexual selection. Yet, the developmentalmechanisms that produce such diversification, and thus the proximatetargets of selection pressures, are rarely addressed and poorlyunderstood. In particular, because feathers are colored duringgrowth, the dynamics of feather growth play a deterministicrole in the variation in ornamentation. No study to date, however,has addressed the contribution of feather growth to the expressionof carotenoid-based ornamentation. Here, we examine the developmentalbasis of variation in ornamental feather shapes in male housefinches (Carpodacus mexicanus)—a species in which carotenoiddisplays are under strong natural and sexual selection. First,we use geometric morphometrics to partition the observed shapevariation in fully grown feathers among populations, ages, degreesof elaboration, ornamental body parts, and individuals. Second,we use a biologically informed mathematical model of feathergrowth to predict variation in shape of ornamental feathersdue to simulated growth rate, angle of helical growth of featherbarbs, initial number of barb ridges, rate of addition of newbarbs, barb diameter, and ramus-expansion angle. We find closeconcordance between among-individual variation in feather shapeand hue of entire ornament, and show that this concordance canbe attributed to a shared mechanism—growth rate of featherbarbs. Predicted differences in feather shape due to rate ofaddition of barbs and helical angle of feather growth explainedobserved variation in ornamental area both among individualsand between populations, whereas differences in helical angleof growth and the number of barbs in the feather follicle explaineddifferences in feather shape between ornamental parts and amongmales of different ages. The findings of a close associationof feather growth dynamics and overall ornamentation identifythe proximate targets of selection for elaboration of sexualdisplays. Moreover, the close association of feather growthand pigmentation not only can reinforce condition-dependencein color displays, but can also enable phenotypic and geneticaccommodation of novel pigments into plumage displays providinga mechanism for the observed concordance of within-populationdevelopmental processes and between-population diversificationof color displays.     

Mitochondrial function,ornamentation, and immunocompetence

Understanding the mechanisms that link ornamental displays and individual condition is key to understanding the evolution and function of ornaments. Immune function is an aspect of individual quality that is often associated with the expression of ornamentation, but a general explanation for why the expression of some ornaments seems to be consistently linked to immunocompetence remains elusive. We propose that condition‐dependent ornaments may be linked to key aspects of immunocompetence through co‐dependence on mitochondrial function. Mitochondrial involvement in immune function is rarely considered outside of the biomedical literature, but the role of mitochondria as the primary energy producers of the cell and the centres of biosynthesis, the oxidative stress response, and cellular signalling place them at the hub of a variety of immune pathways. A promising new mechanistic explanation for correlations between a wide range of ornamental traits and the properties of individual quality is that mitochondrial function may be the ‘shared pathway’ responsible for links between ornament production and individual condition. Herein, we first review the role of mitochondria as both signal transducers and metabolic regulators of immune function. We then describe connections between hormonal pathways and mitochondria, with implications for both immune function and the expression of ornamentation. Finally, we explore the possibility that ornament expression may link directly to mitochondrial function. Considering condition‐dependent traits within the framework of mitochondrial function has the potential to unify central tenets within the study of sexual selection, eco‐immunology, oxidative stress ecology, stress and reproductive hormone biology, and animal physiology.     

Speciation is associated with changing ornamentation rather than stronger sexual selection

Although sexual ornamentation mediates reproductive isolation, comparative evidence does not support the hypothesis that stronger sexual selection promotes speciation. Prior analyses have neglected the possibility that decreases in ornamentation may also promote speciation, such that both increases and decreases in the strength of sexual selection and associated changes in ornamentation promote speciation. To test this hypothesis, we studied color ornamentation in one of the largest and fastest avian radiations, the estrildid finches. We show that more ornamented lineages do not speciate more, even though they tend to have faster rates of ornamental evolution, whereas changes in ornamentation (i.e., increases or decreases) are associated with speciation. This indicates that divergence in sexually selected ornamentation, rather than stronger sexual selection, promotes or is otherwise associated with speciation. We also show that gregariousness and investment in reproduction are related to the elaboration of some ornamental traits, suggesting ecological influences on speciation mediated by ornamentation. We conclude that past work focusing specifically on the strength of sexual selection may have greatly underestimated the importance of sexual ornamentation for speciation.     

祖先性状重建法揭示铁线蕨属植物孢子表面纹饰的形态多样性及其演化

蕨类植物孢子表面的纹饰形态复杂多样, 在分类学和古生物学研究中具有重要意义, 目前探索祖先性状演化常用形态学与系统发育学结合的方法。本文选取一回羽叶铁线蕨类植物为材料, 用扫描电子显微镜观察其孢子形态, 用5个叶绿体基因序列构建系统树, 用最大似然法和最大简约法来分析孢子性状的演化。结果表明: 一回羽叶铁线蕨类的孢子纹饰有5种类型, 即光滑、粗糙、颗粒、疣状及瘤状; 这一类群的祖先孢子纹饰有很大的可能是瘤状纹饰; 疣状纹饰可能是由瘤状纹饰演化而来; 孢子纹饰由简单向复杂演化, 越进化的物种其孢子纹饰越复杂。     

Pollen Morphology and Wall Stratification in Rubus L. (Rosaceae) in the Iberian Peninsula

A palynological study using LM and SEM was made of the morphology, ornamentation, apertural system and structure of the pollen wall, of the 18 species of the genus Rubus which are present in the Iberian Peninsula. We demonstrate: a) the stenopalynous character of the genus (isopolar, subprolate, 3-lobate, 3-zonocolporate and striate-perforate, except for R. genevierii whose ornamentation is striate-gemmate-perforate and with a striated margin); b) the lalongate shape of the endoaperture, due to the disappearance or thinning of the nexine; and c) the existence of two structural types, Type I granular and Type II columellar with clearly-defined columellar spaces or compact without intercolumellar spaces.     

Caractères ultrastructuraux du pollen de quelques Podostémales. Affinités avec les Rosidae évoluées

The order Podostemales are two pantropical families of aquatic plants living in running water: Tristichaceae (five genera, ten species) and Podostemaceae (35 genera, 200 species). Pollen of the 26 genera and 62 species studied is characterized by: a granular infratectum in which the granules are sometimes organized as columellae like units, and a lamellar and/or granular endexine in all pollen types, single or in dyads. Most of the apertural characters and the ornamentation of the exine allow us to distinguish the previously established taxonomic groups: Weddellinoideae have tricolporate rugulo-areolate, pollen with a smooth apertural membrane; in Tristichoideae, pollen is periporate and the microspines of the tectum and of the apertural membrane are massive; in Podostemaceae, the tectal spines with their broad bulbous base are formed from numerous masses of sporopollenin, the endexine is microfibrillar at the base, the apertural membrane is constituted of structured ectexinous masses, and the endexine is granular at the aperture. Presence of the tricolporate pollen type associates the Podostemales with the higher eudicotyledons. Most of the pollen characters of Podostemales and their variations are found among advanced Rosidae (Hamamelidales, Polygalales, Santalales, Violales, Euphorbiaceae).     

Ontogenetic Torsion and Protoconch Form in the Archaeogastropod Haliotis kamtschatkana: Evolutionary Implications

Ontogenetic torsion in archaeogastropods has strongly influenced theories about early gastropod evolution, but the seminal studies by Crofts (1937, 1955) remain the major source of information about tissue movements during this developmental process. Computer-generated reconstructions of histological sections indicate that the cephalopodium of Haliotis kamtschatkana Jonas, 1845 rotates by a full 180° relative to the shell and visceral lobe during the first quarter of pre-metamorphic development. However, a portion of pallial epithelium, including some of the shell field, accompanies the rotating cephalopodium; a process facilitated by detachment of the pallium from the apertural rim of the protoconch. Transmission electron microscopy indicates that a tract of the larval retractor muscle, which Crofts (1955) implicated in the mechanism of torsion, inserts on both pedal and pallial cells. A deep invagination of shell field epithelium is a major focus of rotational torque. As a result of pallial deformation during cephalopodial rotation, the anus and gill rudiment are restricted to the right half of the larval body for 2 days after cephalopodial rotation by 180° has been completed. Scanning and transmission electron microscopy indicate that grooves in the lateral flanks of the protoconch correspond to the deep invagination of shell field epithelium. The grooves are not created by a coiling type of accrelionary shell growth or by flexion of the protoconch. A calcareous shelf is secondarily added to the periostracal template of the protoconch along its visceral apertural rim. Morphogenetic movements during ontogenetic torsion in this species are more complex than a simple rotation between cephalopodium and visceropallium and the protoconch shows no evidence of exogastric coiling. © 1997 Published by Elsevier Science Ltd on behalf of The Royal Swedish Academy of Sciences.     

Parental Effort in Relation to Structural Plumage Coloration in the Mountain Bluebird (Sialia currucoides)

Indicator models of sexual selection suggest that costly ornaments signal reliable information regarding an individual's quality to potential mates. In species that produce altricial offspring, the amount of parental care provided by both males and females can impact reproductive success. The Good Parent Hypothesis proposes that ornamentation in biparental species can act as an honest signal of parental ability to potential mates. We tested this hypothesis using the mountain bluebird (Sialia currucoides), a sexually dichromatic, socially monogamous species in which both sexes have structurally based ornamental plumage coloration. A male's plumage color predicted neither the rate at which it provisioned nestlings nor brood growth rate. The same was true for females. We also found no indication of assortative mating by color or body condition. Feeding rates within pairs were positively correlated, which we suggest may be due to pairs responding similarly to the perceived needs of nestlings or to local area prey availability. In sum, our results do not support the Good Parent Hypothesis as an explanation for the evolution of ornamental plumage color in mountain bluebirds. We suggest alternative hypotheses for the evolution of ornamental plumage color in this species.     

Differentiation and growth of bone ornamentation in vertebrates: A comparative histological study among the Crocodylomorpha

Bone ornamentation, that is, hollow (pits and grooves) or protruding (ridges) repetitive reliefs on the surface of dermal bones, is a frequent, though poorly studied and understood, feature in vertebrates. One of the most typical examples of this characteristic is given by the Crurotarsi, a taxon formed by the crocodilians and their closest allies, which generally display deep ornamentation on skull roof and osteoderms. However, the ontogenetic process responsible for the differentiation and development of this character remains controversial. This study was conducted to settle the question on histological and microanatomical evidence in several crurotarsan taxa. Observational and experimental data in extant and extinct crocodyliforms show that bone ornamentation is initially created, and later maintained during somatic growth (that is indefinite in crocodilians), by a complex process of bone remodeling comprising local resorption of superficial bone cortices, followed by partial reconstruction. The superficial reliefs of crocodilian dermal bones are thus permanently modified through pit enlargement, drift, stretching, shrinking, or complete filling. Ridges are also remodeled in corresponding ways. These processes allow accommodation of unitary ornamental motifs to the overall dimensions of the bones during growth. A parsimony optimization based on the results of this study, but integrating also published data on bone histology in non‐crocodyliform crurotarsans and some non‐crurotarsan taxa, suggests that the peculiar mechanism described above for creating and maintaining bone ornamentation is a general feature of the Crurotarsi and is quite distinct from that attributed by previous authors to other vertebrates. J. Morphol. 276:425–445, 2015. © 2014 Wiley Periodicals, Inc.     

Large‐scale evolutionary trends of Acrochordiceratidae Arthaber, 1911 (Ammonoidea,Middle Triassic) and Cope’s rule

Abstract: Directed evolution of life through millions of years, such as increasing adult body size, is one of the most intriguing patterns displayed by fossil lineages. Processes and causes of such evolutionary trends are still poorly understood. Ammonoids (externally shelled marine cephalopods) are well known to have experienced repetitive morphological evolutionary trends of their adult size, shell geometry and ornamentation. This study analyses the evolutionary trends of the family Acrochordiceratidae Arthaber, 1911 from the Early to Middle Triassic (251–228 Ma). Exceptionally large and bed‐rock‐controlled collections of this ammonoid family were obtained from strata of Anisian age (Middle Triassic) in north‐west Nevada and north‐east British Columbia. They enable quantitative and statistical analyses of its morphological evolutionary trends. This study demonstrates that the monophyletic clade Acrochordiceratidae underwent the classical evolute to involute evolutionary trend (i.e. increasing coiling of the shell), an increase in its shell adult size (conch diameter) and an increase in the indentation of its shell suture shape. These evolutionary trends are statistically robust and seem more or less gradual. Furthermore, they are nonrandom with the sustained shift in the mean, the minimum and the maximum of studied shell characters. These results can be classically interpreted as being constrained by the persistence and common selection pressure on this mostly anagenetic lineage characterized by relatively moderate evolutionary rates. Increasing involution of ammonites is traditionally interpreted by increasing adaptation mostly in terms of improved hydrodynamics. However, this trend in ammonoid geometry can also be explained as a case of Cope’s rule (increasing adult body size) instead of functional explanation of coiling, because both shell diameter and shell involution are two possible paths for ammonoids to accommodate size increase.     

Buckman's first law of covariation–a case of proportionality

Buckman's first law of covariation states that in ammonoids, coarseness of ribs and other ornamentation correlates negatively with degree of compression (whorl height/width ratio) and involution. This ‘law’ has escaped several attempts at functional or physiological explanation. However, Buckman's first law becomes self‐evident, and almost collapses to a tautology, when we reword it as follows: Size of lateral and ventral ornamentation correlates with size of the aperture (its width and height, respectively). In a shell that is strongly compressed laterally, the diameter of the aperture, and hence the soft parts within, is small in the lateral direction. For such a shell, coarse lateral ribbing would imply impossibly strong relative compression and expansion of the soft body through the formation of a rib. Conversely, if the shell is depressed (compressed dorsoventrally), the diameter of the soft parts is small in the dorsoventral direction, and the ventral ribbing correspondingly small. Buckman's first law is thus simply a statement of proportionality that needs no special explanation. On the contrary, any departure from this ‘law’ would imply strong deformations of the soft body as ornamentation was formed. Measurements on the Triassic ammonite Pseudodanubites halli show that ventral ribs get relatively stronger on compressed specimens, in accordance with this view of Buckman's law. ? Ammonoids, intraspecific variation, morphometry, ornamentation.     

Sexually selected sexual selection: Can evolutionary retribution explain female ornamental colour?

By preferring mates with increasingly costly ornaments or courtship displays, females cause an escalation of male reproductive costs. Such increased costs should promote male selectivity based on fecundity‐linked female attributes, leading to female ornamentation in species with traditional sex roles. Consequently, female ornamentation should evolve more frequently in taxa where male reproduction is costly than in comparable taxa where it is cheaper. We assessed the prevalence of female ornamental colouration in two clades of viviparous cyprinodontid fish: the Goodeinae, where stringent female choice imposes male mating costs, and the Poeciliinae, whose males can circumvent female mate choice. We found that although in the Poeciliinae female ornamental colour is a correlated, but paler version of male coloration, females of the Goodeinae often display vivid ornamental colours that are distinct from those of males. Thus, male and female ornaments are not (phylo)genetically correlated in the Goodeinae. Furthermore, phylogenetic signal on male and female colour is clearly detectable in the Poeciliinae, but absent in the Goodeinae, suggesting that ornamental colour of males and females in the latter may be the consequence of selection. Given that enforceable female choice has promoted male ornaments, we propose that evolutionary retribution has promoted distinct female ornaments in the Goodeinae.