Development of the amentiferous concept

The amentiferous concept developed in pre-Linnaean times, and early botanists clearly recognized the topical similarities among plants bearing aments. Among the amentiferous plants placed side by side in early times were many that would not be so situated today—e.g., gymnosperms intermixed with dicotyledons. By the time of Linnaeus, only dicotyledons were included among the ament-bearing groups. J. G. Gmelin was first to recognize ament-bearing plants (including some gymnosperms) under a single category, “Amentaceae.” Linnaeus, A. L. de Jussieu, W. J. Hooker, Lindley, and Eichler, at one time or another, placed these plants in a separate amentaceous category, but never under the term “Amentiferae.” The name was never used by Engler although he did place the ament-bearing plants among the first families of his Archichlamydeae. The category “Amentiferae” appears to have entered the literature in British publications and through British/English translations from the German.     

Introduction and Notes to R. Dahlgren's System of Classification of the Angiosperms

The present paper aims at introducting Dahlgren’s system of classification of the angiosperms. Phenetic and phylogenetic classifications are discussed. The basic principles and methods used by Dahlgren are explained. Dahlgren’s opinions on some important problems, such as the origin of angiosperms, the flowers of primitive angiosperms, the relation between the dicotyledons and monocotyledons, the origin of the monocotyledons, the treatment of the “Amentiferae” and of the orders of the “Sympetalae”, are all expressed. A brief comparison between Dahlgren’s system and three other current systems, viz. those of Takhtajan, Cronquist and Thorne is also given.     

Extant‐only comparative methods fail to recover the disparity preserved in the bird fossil record

Most extant species are in clades with poor fossil records, and recent studies of comparative methods show they have low power to infer even highly simplified models of trait evolution without fossil data. Birds are a well‐studied radiation, yet their early evolutionary patterns are still contentious. The fossil record suggests that birds underwent a rapid ecological radiation after the end‐Cretaceous mass extinction, and several smaller, subsequent radiations. This hypothesized series of repeated radiations from fossil data is difficult to test using extant data alone. By uniting morphological and phylogenetic data on 604 extant genera of birds with morphological data on 58 species of extinct birds from 50 million years ago, the “halfway point” of avian evolution, I have been able to test how well extant‐only methods predict the diversity of fossil forms. All extant‐only methods underestimate the disparity, although the ratio of within‐ to between‐clade disparity does suggest high early rates. The failure of standard models to predict high early disparity suggests that recent radiations are obscuring deep time patterns in the evolution of birds. Metrics from different models can be used in conjunction to provide more valuable insights than simply finding the model with the highest relative fit.     

Deconstructing the long-standing a priori assumption that serial homology generally involves ancestral similarity followed by anatomical divergence

It has long been assumed that serial homologues are ancestrally similar—polysomerism resulting from a “duplication” or “repetition” of forms—and then often diverge—anisomerism, for example, as they become adapted to perform different tasks as is the case with the forelimb and hind limbs of humans. However, such an assumption, with crucial implications for comparative, evolutionary, and developmental biology, and for evolutionary developmental biology, has in general not really been tested by a broad analysis of the available empirical data. Perhaps not surprisingly, more recent anatomical comparisons, as well as molecular knowledge of how, for example, serial appendicular structures are patterned along with different anteroposterior regions of the body axis of bilateral animals, and how “homologous” patterning domains do not necessarily mark “homologous” morphological domains, are putting in question this paradigm. In fact, apart from showing that many so-called “serial homologues” might not be similar at all, recent works have shown that in at least some cases some “serial” structures are indeed more similar to each other in derived taxa than in phylogenetically more ancestral ones, as pointed out by authors such as Owen. In this article, we are taking a step back to question whether such assumptions are actually correct at all, in the first place. In particular, we review other cases of so-called “serial homologues” such as insect wings, arthropod walking appendages, Dipteran thoracic bristles, and the vertebrae, ribs, teeth, myomeres, feathers, and hairs of chordate animals. We show that: (a) there are almost never cases of true ancestral similarity; (b) in evolution, such structures—for example, vertebra—and/or their subparts—for example, “transverse processes”—many times display trends toward less similarity while in many others display trends toward more similarity, that is, one cannot say that there is a clear, overall trend to anisomerism.     

On the systematic position of the Early Cretaceous fern genus “Athyrium”

The oldest fossils assigned to Athyrium (mostly based on the sorus morphology) comprise fronds and spores from the Lower Cretaceous of Northeast Asia. However, most molecular dating suggests that extant Athyrium diverged from its sister genus during the Eocene or later, implying that the Cretaceous fossils probably belong to another polypodiaceous taxon. By examining the sorus morphology of extant genera related to the family Athyriaceae, we found that the primary diagnostic feature for assigning the Cretaceous fossils to Athyrium, i.e., the sorus shape, is common to the entire extant family, or plesiomorphic for the genus. As the fronds are more commonly preserved than the reproductive parts, we compared the fossil frond morphology with those of living taxa of the family that is divided into two types. The Cretaceous fossil we examined here bears the frond’s costal groove characters on adaxial side, which is more closely related to that of the Deparia-clade instead of the clade including Athyrium and other genera of the family. The observation is further confirmed by the cladistic analysis using morphological characters. The systematic position of the Early Cretaceous “Athyrium” was resolved as a stem member of the total Athyriaceae using a tip-dating approach with the Fossilized Birth-Death model in a Bayesian framework. Our study suggests that Early Cretaceous fossils previously assigned to Athyrium require taxonomic revision.     

The chelifores of sea spiders (Arthropoda,Pycnogonida) are the appendages of the deutocerebral segment

SUMMARY Within the last decade, gene expression patterns and neuro‐anatomical data have led to a new consensus concerning the long‐debated association between anterior limbs and neuromeres in the arthropod head. According to this new view, the first appendage in all extant euarthropods is innervated by the second neuromere, the deutocerebrum, whereas the anterior‐most head region bearing the protocerebrum lacks an appendage. This stands in contrast to the clearly protocerebrally targeted “antennae” of Onychophora and to some evidence for protocerebral limbs in fossil euarthropod representatives. Yet, the latter “frontal appendages” or “primary antennae” have most likely been reduced or lost in the lineage, leading to extant taxa. Surprisingly, a recent neuro‐anatomical study on a pycnogonid challenged this evolutionary scenario, reporting a protocerebral innervation of the first appendages, the chelifores. However, this interpretation was soon after questioned by Hox gene expression data. To re‐evaluate the unresolved controversy, we analyzed neuro‐anatomy and neurogenesis in four pycnogonid species using immunohistochemical techniques. We clearly show the postprotocerebral innervation of the chelifores, which is resolved as the plesiomorphic condition in pycnogonids when evaluated against a recently published comprehensive phylogeny. By providing direct morphological support for the deutocerebral status of the cheliforal ganglia, we reconcile morphological and gene expression data and argue for a corresponding position between the anterior‐most appendages in all extant euarthropods. Consequently, other structures have to be scrutinized to illuminate the fate of a presumptive protocerebral appendage in recent euarthropods. The labrum and the “frontal filaments” of some crustaceans are possible candidates for this approach.     

Species and paleoanthropology

The biotic world is self-evidently “packaged” into units, of which the most basic is the species. It is necessary to develop an accurate understanding of what species are and how they are to be identified before we can proceed to more complex analyses of the evolutionary histories and relationships of extinct and extant taxa at all levels of the systematic hierarchy. In this article, we review the major species concepts current today among paleoanthropologists, and examine the limitations of their applicability to practical studies of extant and extinct faunas. The primary such limitation for paleoanthropologists is the fact that all major species definitions stress reproductive continuity (whether by exclusionary or inclusionary mechanisms), a quality that is inferential at best among forms known only as fossils (and, in many cases, in the extant fauna as well). The only reliable signal as to species status in the fossil record is morphology, yet speciation carries with it no specifiable quantity of morphological innovation. Some groups with autapomorphies are not species, and some species do not bear autapomorphies. How, then, are we to recognize species in the hominid and other fossil records? Noting that osteodental differences among congeneric primate species tend to be subtle, and that when consistent identifiable “morphs” can be found at least as many species are present, we recommend equating morphs based on several characters with species—realizing that only one or two distinctive characters may not make a morph. In this way, our views of the phylogenetic histories of higher taxa may be oversimplified, but their essential patterns will not be distorted.     

Vegetative anatomy and morphology of Amentiferae

Some of the important vegetative characters of amentiferous families described in research papers concerned with interfamilial relationships and published since 1947–1948 are briefly summarized. Important conclusions by the writers are also reviewed. Thus, Bataceae are considered derivatives of the Centrospermae ; Betulaceae, Casuarinaceae, and Ulmaceae are hamamelidaceous derivatives; Garryaceae and Cornaceae are closely related; and Julianiaceae and Anacardiaceae are plausible relatives. A more extensive tabulation of vegetative characters of essentially all amentiferous and ranalean families and Dilleniaceae is also presented, and the data are utilized to compare the percentages of advanced and primitive characters in these same taxa. The logic for considering trends whose evolutionary directions were initiated previous or incidental to angiospermous origin as more valid than other evolutionary trends is offered. The following principal conclusions are based upon the analysis of the tabulation: (1) the Amentiferae have sufficient advanced vegetative characters to preclude consideration of them as earliest angiosperms or their direct derivatives unless a large number of primitive features of other sorts offers strong counter evidence; (2) they also have sufficient primitive characters to discount the concept that they are highly evolved; (3) most Amentiferae clearly possess many more advanced vegetative characters than do most Magnoliales, Laurales, Trochodendrales, or Dilleniaceae ; and (4) relationships of several Amentiferae to various, unrelated angiospermous families have been demonstrated.     

Dicotyledonous woods from the Upper Cretaceous of southern Illinois

Dicotyledonous woods from the Upper Cretaceous of Southern IllinoiS. Five species of fossil dicotyledonous wood are described from an Upper Cretaceous (Maestrichtian; locality in Alexander County, IllinoiS. U.S. A. Paraquercimum cretaceum has structure similar to the Fagaceae (evergreen Oak- Lithocarpus ) and Casuarinaceae and represents the earliest known occurrence of this structural type (large solitary pores and uniseriate and large multiseriale rays). Paraphyltanthoxyhin illirioisense and Icacinoxylon alternipunctata are species of genera represented at other Cretaceous and Early Tertiary localities In large diameter trees. Parabombacaceoxylon magniporosum has large diameter pores and scalariform perforation plates, a combination of characters that is extremely rare in the extant flora. Paraapocynaceoxylon barghoorni has a combination of characters represented in extant Apocynaceae. These five species lack growth rings, have high vulnerability indices (mean vessel diameter divided by mean number of vessels per square millimeter, and a relatively high proportion of ray parenchyma. They lack specialized wood anatomical characters, and a compilation of vessel element lengths in these and other Cretaceous woods indicates that short vessel elements (a derived character) were less frequent in the Cretaceous than in extant dicotyledonous trees.     

When Earth started blooming: insights from the fossil record

Recent palaeobotanical studies have greatly increased the quantity and quality of information available about the structure and relationships of Cretaceous angiosperms. Discoveries of extremely well preserved Cretaceous flowers have been especially informative and, combined with results from phylogenetic analyses of extant angiosperms (based mainly on molecular sequence data), have greatly clarified important aspects of early angiosperm diversification. Nevertheless, many questions still persist. The phylogenetic origin of the group itself remains as enigmatic as ever and, in some cases, newly introduced techniques from molecular biology have given confusing results. In particular, relationships between the five groups of extant seed plants remain uncertain, and it has sometimes proved difficult to reconcile estimates of the time of divergence between extant lineages made using a 'molecular clock' with the fossil record. One result, however, is becoming increasingly clear: a great deal of angiosperm diversity is extinct. Some groups of angiosperms were evidently more diverse in the past than they are today. In other cases, fossils defy assignment to extant groups at the family level or below. This raises the possibility that evolutionary conclusions based solely upon extant taxa that are merely relics of groups that were once much more diverse might be misled by the effects of extinction. It also introduces the possibility that some early enigmatic fossils might represent lineages that diverged from the main line of angiosperm evolution below the most recent common ancestor of all extant taxa. These, and other questions, are among those that need to be addressed by future palaeobotanical research.     

Tarsiers,adapids and the integrity of Strepsirhini

Although strepsirhine primates can be described by their narial configuration, this and most other definable features are probably primitive retentions; only the development of a grooming claw of the second pedal digit and of a toothcomb (the latter of which has been lost in Daubentonia) emerge as potential apomorphies of the group. Within this assemblage lemurids, Lepilemur, the indriids, and Daubentonia can be argued to constitute a monophyletic group whose relationships cladistically are in the sequence listed; Lemuridae and Indriidae can themselves be delineated as monophyletic groups. The remaining strepsirhine primates—the cheirogaleids, galagids, and lorisids—also appear to constitute a definable clade, with the former group representing the sister taxon of the latter two families; cach family can be united on the basis of distinct synapomorphies. Although there are features—especially of the ear region—which present themselves as potentially reflective of the sister relationship of Tarsius + Anthropoidea, other characters, including the possession of the grooming claw, are suggestive of an alternative scheme: Tarsius may be the sister of the extant lorisiform group, thereby reconstituting, albeit in a novel form, the primate suborder Prosimii. It also appears that fossil “tarsioids” may in fact be more closely related to the extant lorisiforms than to Tarsius. A reconsideration of the so-called fossil lemurs, the adapids, leads to the conclusion that Adapis-like primates are a clade apart from Pelycodus, Notharctus, Smilodectes and their most immediate relatives, and may themselves constitute a clade that is related as the primitive sister to all other “prosimians” by virtue of the development of the so-called free intrabullar tympanic ring.     

THE CONTRIBUTION OF GENE MOVEMENT TO THE “TWO RULES OF SPECIATION”

The two “rules of speciation”—the Large X‐effect and Haldane's rule—hold throughout the animal kingdom, but the underlying genetic mechanisms that cause them are still unclear. Two predominant explanations—the “dominance theory” and faster male evolution—both have some empirical support, suggesting that the genetic basis of these rules is likely multifarious. We revisit one historical explanation for these rules, based on dysfunctional genetic interactions involving genes recently moved between chromosomes. We suggest that gene movement specifically off or onto the X chromosome is another mechanism that could contribute to the two rules, especially as X chromosome movements can be subject to unique sex‐specific and sex chromosome specific consequences in hybrids. Our hypothesis is supported by patterns emerging from comparative genomic data, including a strong bias in interchromosomal gene movements involving the X and an overrepresentation of male reproductive functions among chromosomally relocated genes. In addition, our model indicates that the contribution of gene movement to the two rules in any specific group will depend upon key developmental and reproductive parameters that are taxon specific. We provide several testable predictions that can be used to assess the importance of gene movement as a contributor to these rules in the future.     

Cretaceous Promecognathinae (Goleoptera: Garabidae): a new genus, phylogenetic reconstruction and zoogeography

A fossil carabid (Coleoptera) belonging to the subfamily Promecognathinae is described from Cretaceous crater lake deposits at Orapa, Botswana. The specimen, which is the first ever fossil promecognathine, is placed in a new genus Palaeoxinidium. The fossil supports a Cretaceous age determination for the sediments, and it indicates that the Orapa crater was, at least in part, heavily vegetated at the time of deposition. Cladistic analysis of the fossil and extant Promecognathinae indicates that the fossil represents the sister group of the extant taxa and that the latter, in turn, comprise a pair of sister groups. It is, therefore, proposed that the Promecognathinae should be divided into two tribes: the Palaeoaxinidiini, represented by the fossil, and the Promecognathini, represented by the extant groups. The Promecognathini should be subdivided further into the subtribes Promecognathina and Axinidiina, represented by the American and African species, respectively. Three alternative zoogeographic hypotheses that explain the distributions of the fossil and extant promecognathines are discussed and tests for the hypotheses are proposed.     

Fossils and seed plant phylogeny reanalyzed

In a cladistic analysis of Recent seed plants, Loconte and Stevenson (1990) obtained results that conflict with our 1986 analysis of both extant and fossil groups and argued that fossil data had led us to incorrect conclusions. To explore this result and the general influence of fossils on phylogeny reconstruction, we assembled new “Recent” and “Complete” (extant plus fossil) data sets incorporating new data, advances in treatment of characters, and those changes of Loconte and Stevenson that we consider valid. Our Recent analysis yields only one most parsimonious tree, that of Loconte and Stevenson, in which conifers are linked with Gnetales and angiosperms (anthophytes), rather than with Ginkgo, as in our earlier Recent and Complete analyses. However, the shortest trees derived from our Complete analysis show five arrangements of extant groups, including that of Loconte and Stevenson and our previous arrangements, suggesting that the result obtained from extant taxa alone may be misleading. This increased ambiguity occurs because features that appear to unite extant conifers and anthophytes are seen as convergences when fossil taxa are interpolated between them. All trees found in the Complete analysis lead to inferences on character evolution that conflict with those that would be drawn from Recent taxa alone (e.g., origin of anthophytes from plants with a “seed fern” morphology). These results imply that conclusions on many aspects of seed plant phylogeny are premature; new evidence, which is most likely to come from the fossil record, is needed to resolve the uncertainties.     

Variations of Mesozoic feathers: Insights from the morphogenesis of extant feather rachises

The rachises of extant feathers, composed of dense cortex and spongy internal medulla, are flexible and light, yet stiff enough to withstand the load required for flight, among other functions. Incomplete knowledge of early feathers prevents a full understanding of how cylindrical rachises have evolved. Bizarre feathers with unusually wide and flattened rachises, known as “rachis-dominated feathers” (RDFs), have been observed in fossil nonavian and avian theropods. Newly discovered RDFs embedded in early Late Cretaceous Burmese ambers (about 99 million year ago) suggest the unusually wide and flattened rachises mainly consist of a dorsal cortex, lacking a medulla and a ventral cortex. Coupled with findings on extant feather morphogenesis, known fossil RDFs were categorized into three morphotypes based on their rachidial configurations. For each morphotype, potential developmental scenarios were depicted by referring to the rachidial development in chickens, and relative stiffness of each morphotype was estimated through functional simulations. The results suggest rachises of RDFs are developmentally equivalent to a variety of immature stages of cylindrical rachises. Similar rachidial morphotypes documented in extant penguins suggest that the RDFs are not unique to Mesozoic theropods, although they are likely to have evolved independently in extant penguins.     

Mountain gorilla life histories,reproductive competition,and sociosexual behavior and some implications for captive husbandry

New data on modal patterns of, and variability in, mountain gorilla life history tactics and sociosexual behavior, collected during long-term fieldwork at the Karisoke Research Centre, are presented here. These data show that immature males and females develop sociosexual relationships with both peers and adults, and that these relationships—which contribute to more complex social relationships—often lead to mating between natal females and males who are “familiar” partners. They also show that within-group reproductive competition between males can reach considerable levels, but that immature males are sometimes able to copulate with both nulliparous and parous females; that unrelated silverbacked males can reside in the same all-male group but become intolerant of each other if females join the group; and that, whereas females sometimes compete with each other for opportunities of copulate, their probability of conceiving is probably not decreased by any inability to control the mating situation. This information may contribute to continued improvement in the husbandry of captive gorillas.     

The origin of angiosperms

The claim of monophyletic origin of angiosperms arose from the confusion of phylogenetic and taxonomic concepts. Unpreconceived studies of extant angiosperms point to more than one archetype. Several lines of angiosperms have simultaneously entered the fossil record; the monocotyledons, proto-Hamamelidales, proto-Laurales and “proteophylls” (possibly ancestral to the Rosidae) are recognized among them. Three groups of Mesozoic seed plants — the Caytoniales, Czekanowskiales and Dirhopalostachyaceae — are distinguished as major sources of angiosperm characters (proangiosperms). Other Mesozoic lineages probably also contributed to the angiosperm character pool. Angiospermization is related to Mammalization and other processes involved in development of the Cenozoic lithosphere and biosphere.     

The Paleobiosphere: a novel device for the in vivo testing of hydrocarbon producing-utilizing microorganisms

The construction and testing of a unique instrument, the Paleobiosphere, which mimics some of the conditions of the ancient earth, is described. The instrument provides an experimental testing system for determining if certain microbes, when provided an adequate environment, can degrade biological materials to produce fuel-like hydrocarbons in a relatively short time frame that become trapped by the shale. The conditions selected for testing included a particulate Montana shale (serving as the “Trap Shale”), plant materials (leaves and stems of three extant species whose origins are in the late Cretaceous), a water-circulating system, sterile air, and a specially designed Carbotrap through which all air was passed as exhaust and volatile were hydrocarbons trapped. The fungus for initial testing was Annulohypoxylon sp., isolated as an endophyte of Citrus aurantifolia. It produces, in solid and liquid media, a series of hydrocarbon-like molecules. Some of these including 1,8-cineole, 2-butanone, propanoic acid, 2-methyl-, methyl ester, benzene (1-methylethyl)-, phenylethyl alcohol, benzophenone and azulene, 1,2,3,5,6,7,8,8a-octahydro-1,4-dimethyl-7-(1-methylethenyl), [1S-(1α,7α,8aβ)]. These were the key signature compounds used in an initial Paleobiosphere test. After 3 weeks, incubation, the volatiles associated with the harvested “Trap Shale” included each of the signature substances as well as other fungal-associated products: some indanes, benzene derivatives, some cyclohexanes, 3-octanone, naphthalenes and others. The fungus thus produced a series of “Trap Shale” products that were representative of each of the major classes of hydrocarbons in diesel fuel (Mycodiesel). Initial tests with the Paleobiosphere offer some evidence for a possible origin of hydrocarbons trapped in bentonite shale. Thus, with modifications, numerous other tests can also be designed for utilization in the Paleobiosphere.