Buckman's Paradox: variability and constraints on ammonoid ornament and shell shape

Buckman's Law of Covariation states that ammonoid shell shape and ornamentation are typically correlated, such that compressed, involute forms have light ornament while more inflated, evolute forms have heavier ornament. Such covariation has been observed in many ammonoid groups, and implies a link between the morphogenesis of shell shape and ornamentation. However, other evidence suggests that while ornament growth is controlled by the genetic‐developmental program of the ammonoid, shell shape is strongly influenced by environmental factors. These differing viewpoints lead to Buckman's Paradox – are ornamentation and shell shape tightly linked, as implied by Buckman's covariation, or is the morphogenesis of ornament controlled genetically, while shell shape is controlled environmentally? To address this issue, the variability of shell shape and rib morphology has been compared for a group of endemic acanthoceratid ammonites from the Cretaceous Western Interior Seaway of North America. If Buckman's Law holds due to a morphogenetic connection between shell shape and ornamentation, we would expect taxa with more variable shell shapes to also show more variable rib features and growth. Morphometric analysis of seven shell shape and two rib characters for the Western Interior acanthoceratids finds no such correlation, suggesting that shell shape and rib growth are controlled by different processes. Indeed, rib growth appears to be more constrained than shell shape, consistent with the view that ornamentation is more tightly controlled by the developmental‐genetic growth program of the ammonoid. These results emphasize the complexity of ammonoid morphogenesis and highlight our limited understanding of the causes underlying Buckman's Law.     

Systematic revision and sexual dimorphism in Choffatia (Ammonoidea: Perisphinctoidea) from the Callovian of Kutch,India

The genus Choffatia (Siemiradzki) is an important Callovian ammonite in Kutch biostratigraphy. Several species of the genus were described by the early workers who did not recognize intraspecific variability and sexual dimorphism in terms of the modern concepts. We have described here three dimorphic species based on material reposited in the Geological Survey of India, Kolkata as well as systematically collected additional specimens with precise stratigraphic information. These species are: Choffatia recuperoi (Gemmellaro, 1873), Choffatia cobra (Waagen, 1875), and Choffatia perdagata (Waagen, 1875). The microconchs of the three species, described here for the first time, are lappeted whereas the macroconchs have a simple aperture. However, these species have similar shell morphologies and they intergrade in shell shape and sculpture; thus it is difficult to distinguish them qualitatively. Statistical analyses (both bivariate and multivariate) support that they are distinct species and there is a good correlation among the degree of involution, the degree of inflation, and the strength of ornamentation. C. cobra is the most evolute, depressed, and strongly ribbed whereas C. perdagata is the most involute, compressed, and weakly ribbed. C. recuperoi occupies the intermediate position. This interspecific diversity of Choffatia thus supports Buckman's Law of Covariation.     

The origin and geometry of radial ribbing patterns in articulate brachiopods

Ackerly, S. C. 1992 07 15: The origin and geometry of radial ribbing patterns in articulate brachiopods.
Geometric models for simple. radial ribbing in articulate brachiopods include (1) ribs radiating isometrically from the shell umbo. (2) divergence of thc ribs from some 'point' within the shell, and (3) reorientation of the ribs at right angles to the shell margin. Analyses of the Orthida, the ancestral taxon of articulate brachiopods, indicate that rib geometries are isometric in Early Cambrian taxa (model 1). but that by the Early Ordovician rib orientations are generally perpendicular to the shell margin (model 3). A combination of functional and morphogenetic Factors explains the ribbing geometries observed in orthide brachiopods.     

Functional morphological analysis of evolution of ribbing in pliocene viviparid shells from Croatia

Posilović, H., & Bajraktarević, Z. 2010: Functional morphological analysis of evolution of ribbing in pliocene viviparid shells from Croatia. Lethaia, 10.1111/j.1502‐3931.2009.00209.x. This paper provides a functional analysis of Pliocene freshwater gastropods from the phylogenetic lineage of Viviparus neumayri to Viviparus ornatus. On this section of the phylogenetic line, it is possible to follow the continuous evolution of the shell ornamentation (shouldering and rib formation) from ancestral V. neumayri with smooth shell to V. ornatus with spiral ribs. The shell morphology and possible structural function of rib development is discussed from the analytical point of view, but also by Finite Element Modelling. Spiral rib development in the viviparids is not correlated with shell thickening, but rather with strengthening of the shell is achieved through development of spiral ribs, with direct biomechanical and evoloutionary significance. □Croatia, functional morphology, gastropod shell evolution, Pliocene, viviparid evolution.     

Vicariance and convergence in Magellanic and New Zealand long‐looped brachiopod clades (Pan‐Brachiopoda: Terebratelloidea)

Phylogenetic reconstructions using parsimony, maximum likelihood (ML), and Bayesian relaxed molecular clock analyses of ～2850 nucleotides of nuclear‐encoded small and large subunit ribosomal DNAs (SSU and LSU rDNAs) from 14 long‐looped (terebratelloid) ingroup and four short‐looped (terebratulidine) outgroup brachiopod taxa, together with ML analyses of ～663 nucleotides of mitochondrial cytochrome oxidase subunit 1 (cox1) from 12 terebratelloid taxa, show that deep divergence separates taxa endemic to waters in the vicinity of New Zealand from those with a Magellanic distribution around South America and Antarctica. This deep divergence also separates Magellanic Terebratella dorsata from New Zealand Terebratella sanguinea, showing that they are not congeneric. Instead, they belong to separate ‘Magellanic’ (MAG) and ‘New Zealand’ (NZ) clades that first diverged about 82 Mya (95% highest posterior density, 48–120 Mya), correlating with separation between the NZ and Antarctic tectonic plates. Sequence analyses also reveal (1) that the Antarctic endemic taxa Magellania fragilis and Magellania joubini are not congeneric with Magellania venosa, suggesting that their previous placement in Aerothyris should be restored, and (2) that divergence between Antarctic and NZ species of the terebratulide Liothyrella occurred much later than plate separation, perhaps because of continuing gene flow caused by long‐lived larvae. The topology of the rDNA and cox1 gene trees implies that radial ornament of the shell (‘ribbing’) has been gained (and/or lost) independently within the MAG and NZ clades. Radial ribs are widespread in articulate brachiopods throughout the Phanerozoic, but no comparisons of brachiopod rib morphology and morphogenesis have been published. Our comparisons of transverse shell mid‐sections in the scanning electron microscope reveal no obvious evidence of differences in morphology between independently gained ribs. We also consider several ways in which ribs may affect fitness, including effects on hydrodynamics. Only scanty and inconclusive evidence is available, but we suggest that effects (if any) are likely to be of small magnitude; adaptive value of brachiopod shell ribs remains to be demonstrated. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 162 , 631–645.     

The ammonoid genus <Emphasis Type="Italic">Spiroceras</Emphasis> (Spiroceratidae,Ammonoidea) from the Upper Bajocian of the Northern Caucasus

Ammonites of the genus Spiroceras are described from the Upper Bajocian of the Kyafar River Basin (Karachay-Cherkessia), including macroconchs with preserved apertures and (for the first time in the entire history of the study of Middle Jurassic heteromorphs) microconchs with lateral lappets. The microconchs of S. bispinatum (Baugier et Sauzé), mainly represented by cyrtocones with one ventral row of nodes, correspond to microconchs with a gyroconic shell and a single row of nodes. Macroconchs and microconchs of S. annulatum (Deshayes), the ribs of which lack prominent nodes or spines, are found together. The assemblage also contains small-sized slightly uncoiled shells of Spiroceras aff. S. fourneti Roman et Pétouraud with widely spaced ribs and two prominent rows of large nodes, presumably macro- and microconchs. The ornamentation of this species resembles that of the monomorph Bajocia rarinoda Sturani from the upper part of the Lower Bajocian; perhaps the latter was ancestral to the Middle Jurassic heteromorph. The above species of Spiroceras, as well as S. obliquecostatum (Quenstedt), are figured.     

Opisoma excavatum Boehm,a Lower Jurassic photosymbiotic alatoform‐chambered bivalve

Alatoform bivalves are a polyphyletic group characterized by antero‐posteriorly compressed shells and a ventro‐lateral wing originating from a tight fold of the shell wall. This bizarre shell morphology is interpreted as an adaptation for algal photosymbiosis in heliophilous bivalves. The group contains the living heart cockle Corculum together with four extinct genera ranging in age from the Permian to the Jurassic. The Jurassic alatoform bivalve is Opisoma, which has an aragonitic shell that is divided into two regions, both with different functions: one for stabilization, the other for hosting symbionts. The dorsal part of the shell is massive and played the stabilization role. The ventral part has a very thin and fragile shell that permitted the transmission of light into the internal tissues harbouring photosymbionts. The morphology of this delicate ventral part has thus far remained obscure, due to lack of preservation. Accumulations of Opisoma excavatum Boehm with exquisitely preserved shells containing the fragile winged ventral parts are common within the Pliensbachian shallow‐water, lagoonal carbonate succession of the Rotzo Formation of northern Italy. The wings have internal curved chambers limited by septa parallel to the wing edge. The shell of the ventral part consists of irregular fibrous prismatic and homogeneous structures which progressively infill the chambers. As the chambered wings are analogous structures among alatoform bivalves, they are no longer considered a taxonomic character. According to the observed shell orientation in the field and the consequent organization of the soft parts, Opisoma had an opisthogyrate shell.     

中国伞形科5个引种栽培的模式种果实比较解剖学研究

对中国伞形科中长期引种栽培的旱芹（Apium graveolens L．）、茴香（Foeniculum vulgare Miller）、欧当归（Levisticum of ficinale Koch）、欧防风（Pastinaca sativa L．）和莳萝（Anethum graveolens L．）5个模式种的果实进行了比较解剖学研究,详细描述了各种类分生果的解剖结构及主要特征,并从分生果横切面的轮廓形状、果棱、维管束、油管和合生面等解剖学性状系统分析了各种类的异同,提出果体腹面及合生面的长度及其比值、果棱间距离及其比值、果体侧面与腹面间的角度、侧棱与腹面间的角度、果体长宽比等新的数量化的伞形科解剖性状特征,并按果体压扁程度将5个种类分成几不压扁（旱芹）、轻度压扁（茴香）、中度压扁（莳萝和欧当归）和高度压扁（欧防风）4大类,同时探讨了果实解剖特征与它们的演化程度的关系。     

The Middle–Late Ordovician brachiopod Plectorthis from North America and its paleobiogeographic significance

Plectorthis is a commonly reported Late Ordovician brachiopod genus from Laurentia and several other tectonic plates. Examination of shells from the Cincinnati Arch type area shows that species previously assigned to the genus have a considerable range of variability, such as ribbing style, shell outline, and musculature. The type species, Orthis plicatella Hall, 1847 is characterized by generally simple costae, with double-rowed aditicules on each rib, and a cordate ventral muscle field. Some species previously attributed to Plectorthis have a subtriangular ventral muscle field and hence a closer affinity to the plaesiomyid genus Austinella. Orthis fissicosta is here demoted to a subspecies of Plectorthis plicatella based on its similar ribbing style to some specimens of the type species. In Laurentia, Plectorthis first appeared in the Darriwilian–Sandbian boundary interval, achieved a relatively high species diversity and wide paleogeographic distribution in pericratonic seas during the early Katian; the subequatorial species attained a notably larger shell than those in the higher paleotropics. The genus did not invade the intracratonic basins of Laurentia and became extinct by the late Katian.     

Rib fabrication in Ostreoidea and Plicatuloidea (Bivalvia,Pteriomorphia) and its evolutionary significance

Ribs of Ostreoidea and Plicatuloidea are defined as antimarginal, that is, perpendicular to the margin throughout growth. Morphogenetically, these ribs are unique, since, unlike radial ribs, they are secreted by a homogeneous mantle margin. Based also on the reconstructed shell secretion cycle in Bivalvia, we propose that ribs of Ostreoidea and Plicatuloidea are formed by a mantle margin which, upon extension from the shell margin, stretches and folds by taking the preformed ribs as templates. In extending perpendicular to the margin (as in all Bivalvia growing isometrically), such a mantle extends the rib pattern antimarginally. Ribs of this kind are purely mechanical structures, as their arrangement depends on the mechanical properties of the mantle and on the environmental conditions. This explains the high irregularity of such ribbing patterns. The presence of antimarginal ribs in both the Ostreoidea and Plicatuloidea sheds light on their origin. The first known oyster, Actinostreon cristadifformis, probably derived from an antimarginally ribbed Prospondylidae gen. indet. in the Late Permian or Early Triassic. Antimarginally ribbed Triassic species formerly included in Placunopsis originated both the Dimyidae Atreta in the Late Triassic and Enantiostreon in the Mid Triassic, which was transitional to Plicatulidae. Therefore, Dimyidae and Plicatulidae are closely connected and grouped under Plicatuloidea, to which Ostreoidea is phylogenetically unrelated.     

河北秦皇岛地区奥陶纪奇壁角石科分子的发现及意义

奇壁角石科（Ａｌｌｏｔｒｉｏｃｅｒａｔｉｄａｅ）分子在我华北东部属首次发现。它产于马家沟组下部。界于Ｄｅｆｏｒｍｏｎ－ｃｅｒａｓ－Ｐｅｒｉｐａｔｏｃｅｒａｓ与Ｐｏｌｙｄｅｓｍｉａ－Ｗｕｔｉｎｏｃｅｒａｓ－Ｅｏｓｉｏｔｅｌｕｓ组合带之间偏下。奇壁角石科是Ｆｌｏｗｅｒ,Ｒ．Ｈ．１９９５年创立的,至今科内已建立６属１３种和３个相似种。但以往所描述标本绝大多数仅保存有部分内体管,对其它特征所知甚少。文中描     

Mechanics of sculpture formation in Magadiceramus? rangatira rangatira (Inoceramidae,Bivalvia) from the Upper Cretaceous of New Zealand

The surface sculpture of the inoceramid bivalve Magadiceramus? rangatira rangatira consists of commarginal ribs and curious, transverse wrinkles. The wrinkles typically are at a high angle or orthogonal to the shell margin (‘antimarginal’) and thus differ from purely radial structures. They show features of distribution and morphology that reveal them to be products of margin‐parallel compression of the shell‐secreting mantle and its adjacent, flexible, uncalcified periostracum. The interaction of wrinkles with commarginal ribs indicates that the ribs also formed as folds of the mantle margin. During growth, commarginal folding caused withdrawal of the entire mantle margin towards the umbo, with a consequent reduction in perimeter length. Measurement of specimens indicates that fabrication of the commarginal ribs resulted in the magnitude of commarginal shortening that is required for the formation of transverse wrinkles. We infer that early in ontogeny, at the first development of these sculptures, the wrinkles resulted entirely from mantle contraction and resultant commarginal shortening. With subsequent growth, total wrinkling included a component of ‘pre‐wrinkling’ inherited from the preceding growth stage; the contribution of pre‐wrinkling to total wrinkling increased with shell size. The proposed mechanical model is two‐phase. First, the transversely corrugated (pre‐wrinkled) mantle and periostracum advanced and secreted a slightly concave growth increment. Secondly, the mantle subsequently contracted to create a commarginal rib and increase the number and amplitude of transverse wrinkles. This model is consistent with a homogeneous mantle lacking any differentiated and specialized rib‐constructing segments.     

The taxonomic value of fruit structure in the Chinese endemic genus Dickinsia (Apiaceae)

The monotypic Dickinsia Franch. is misplaced in its current position in the tribe Hydrocotyleae and is transferred to the tribe Mulineae. Its mericarps are strongly dorsally compressed and the lateral ribs of each mericarp are much more prominent than the dorsal rib. The carpophore is free. These characters are typical for genera of the Mulineae but are obviously different from other genera of Hydrocotyleae, which all have laterally compressed mericarps, dorsal ribs which are more expanded than the lateral ribs and an absence of free carpophores.     

Editorial policy and Notes for authors

Modern accounts of evolutionary mechanism pay little attention to the pattern and logic of Darwin's explanation, but recognition of the logic of Darwin's explanation has significance for our understanding of evolution and for our appreciation of the extent to which Darwin's concept of evolutionary mechanism accords with modern evolutionary thought. Also, Darwin's explanation exemplifies the concept of a scientific ‘law’ and thus is of considerable value in helping learners to understand the nature of scientific explanation.     

Sudden origin of ribbing in Jurassic Paracenoceras (Nautiloidea) and its bearing on the evolution of ribbing in post‐Triassic Nautiloids

Ribs appeared cryptically in the Middle Jurassic nautiloid Paracenoceras. These ribs were produced by crowding of growth lirae as a corollary of change in body size during paedomorphic evolution. Initially, they had no direct functional significance. Some other contemporary genera are found to have similar ribbing patterns, partially developed on either the flanks or venter of the adult body chamber.

Subsequently in nautiloid phylogeny, ribs spread all around the whorl, becoming analogous to those of contemporary ammonites. Shell rugosity is observed to occur with increasing frequency in post‐Triassic nautiloids, paralleling the trend in ammonites. This is believed to be an outcome of the ‘arms race’ known as the Mesozoic marine revolution. Ribbing that was not at first adaptive in these nautiloids was subsequently co‐opted as a defensive adaptation. The evolution of this structure is a good example of exaptation.     

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.     

Ribbing patterns in the brachiopod Diceromyonia

Ribbing patterns in two collections of the Ordovician brachiopod Diceromyonia from central North America are compared using the system invented by Bancroft (1928; Manchester Memoirs 72 ) for naming ribs. Simple and nearly symmetric arrangements of frequently occurring branches characterize both collections, with much of the complexity evident in larger valves due to irregular occurrence of low-frequency branches on either side of major ribs. Examination of median sectors reveals little support for Bancroft's derivation of the rhipidomelloid midrib from isorthoid forms, and an alternative non-evolutionary interpretation is proposed in which median patterns appear as predictable outcomes of a simple space-filling process. The approximately 2500 ribs identified in this study are treated as a population in their own right: orderly patterns which emerge are not inferred but are simple summaries of complete counts, obtained by hand. This approach is quick and painless compared to methods using order relations to assess ribbing differences in dalmanellid brachiopods. For this reason, it may encourage systematic study of ribbing patterns among these taxa and lead ultimately to better understanding of the utility of ribbing in their taxonomy.     

The heteromorph ammonite genus Ancyloceras (Ancyloceratidae) in the Paris Basin (lower Aptian,Lower Cretaceous,NE France)

In the present work, we examine a sample of 54 specimens of the heteromorph ammonite genus Ancyloceras d’Orbigny, 1842 from the Argiles à Plicatules Formation (lower Aptian, Lower Cretaceous) of the eastern Paris Basin (NE France). The results are as follows: (1) the sample is regarded as monospecific and assigned to the well-named species Ancyloceras varians d’Orbigny, 1842; (2) its variability is continuous and concerns as well as adult size, coiling and shell ornamentation. Adult size varies by a factor of two from one individual to another. The angle between the proversum and the retroversum is variable. The ornamentation on the spiral and the proversum varies between robust morphotype with trituberculate strong ribs separated by few smooth intercalaries ribs and slender morphotype with feebly tuberculate ribs. There are no arguments in favor of sexual dimorphism in Ancyloceras varians.     

Syn vivo hydrostatic and hydrodynamic properties of scaphitid ammonoids from the U.S. Western Interior

Scaphitid ammonoids were ubiquitous and significant components of the Western Interior Seaway during the Late Cretaceous. This group is characterized by a recurved hook at maturity that deviates from the juvenile whorls. Such a modification seems counterproductive to active locomotion and to manage a biologically effective orientation that facilitates efficient feeding and swimming. Virtually reconstructed 3D hydrostatic models reveal that the examined mature scaphitids had the capacity for neutral buoyancy while assuming a stable, upward-facing orientation in the water column during life. Models of juvenile Hoploscaphites nicolletii suggest that scaphitid apertures were oriented only slightly more horizontal than adults. The hydrostatic influence of sexual dimorphism was explored with the species Hoploscaphites crassus. The inflated macroconch has a lower stability and higher hydrodynamic drag compared to its microconch counterpart. The effect of shell compression was investigated by comparing H. crassus and the more compressed H. nicolletii. The latter species has a relatively high stability and much less hydrodynamic drag during movement. The mature U-shaped body chamber distributes organismal mass in a way that increases stability, and simultaneously orients the soft body so that propulsive energy is efficiently transmitted into horizontal backwards movement with minimal rocking. Swimming velocities computed from hydrodynamic drag experiments suggest that scaphitids were relatively slow swimmers with compressed forms attaining slightly higher velocities (when scaled by mass). Hydrodynamic lift was investigated with computational fluid dynamics simulations. These experiments revealed that the overall shape of the shell is responsible for significant lift in the upwards direction, which is not heavily influenced by ornamentation. This explains how a reduced soft body can overcome and manage a slightly negatively buoyant condition during life. Therefore, the seemingly cumbersome shape and orientation of the scaphitid morphotype may not have been a hindrance during locomotion.