The phylogenetic significance of the carpophore in Apiaceae

Background and aims

Fruit structural characters have traditionally been important in the taxonomy of the family Apiaceae. Previous investigations using a limited number of taxa have shown that the carpophore may be especially useful in helping to circumscribe subfamily Azorelloideae. The present study examines, for the first time, carpophore structure in 92 species from 43 genera, representing all subfamilies of Apiaceae, and including all genera assigned to subfamily Azorelloideae. Phylogenetic interpretations are made for the first time, using all available information, and a standard terminology is proposed to describe the various character states found in carpophores.

Methods

Carpophore structure was studied in detail using light microscopy.

Key Results

Carpophores, when present, may be categorized into two main groups (B and C) based mainly on the arrangement of the vascular bundles in transverse section, and further divided into six sub-types according to the length of the carpophore (short in B1 and C1) and whether they are entire (B1–B3 and C1) or bifurcate (B4 and C2). Free carpophores are absent in subfamily Mackinlayoideae, and in tribes Lichtensteinieae and Phlyctidocarpeae, which have two opposite vascular bundles (Group A). Entire carpophores with one or two vascular bundles, or bifurcate carpophores with lateral vascular bundles (arranged side by side within the commissural plane), are the main types characterizing Azorelloideae. The short, hygroscopic carpophores found in Choritaenia are unique in Apiaceae and provide additional evidence for the exclusion of this genus from Azorelloideae. Carpophore type C2 is typical for most Apioideae sensu lato (exceptions are, for example, Arctopus and Alepidea, which have type B2).

Conclusions

A single carpophore and ventral vascular bundles not forming free carpophores are proposed to be the ancestral conditions in Apiaceae, while bifurcate carpophores with opposite vascular bundles are the derived state, present in most Apioideae. Secondary reductions seem to have occurred in several unrelated lineages in all major groups, e.g. many Azorelloideae, several protoapioids (including nearly all members of the tribe Saniculeae) and 29 euapioid genera (e.g. some Oenantheae).     

The phylogenetic significance of strepsirhinism in Paleogene primates

Two lines of evidence reviewed here argue against the recent proposal that strepsirhinism is an apomorphous feature in primates, shared only by adapiforms and lemuriforms. These are (1) the presence of strepsirhinism in several extant taxa of nonprimate mammals, including Tupaiidae, Tenrecidae, Erinaceus,and Didelphis,and (2) the inferred presence of strepsirhinism not only in adapiforms, but also in all plesiadapiforms and omomyids for which the relevant anatomical regions are known. Therefore, strepsirhinism cannot be invoked as an adaptive innovation underlying the initial strep-sirhine/haplorhine dichotomy. Likewise, the apparent retention of strepsirhinism in omomyids suggests either that the haplorhine oronasal configuration of extant tarsiids and anthropoids was acquired independently or that tarsiids and anthropoids form a clade to the exclusion of omomyids.     

The anatomy, affinity, and phylogenetic significance of Markuelia

The fossil record provides a paucity of data on the development of extinct organisms, particularly for their embryology. The recovery of fossilized embryos heralds new insight into the evolution of development but advances are limited by an almost complete absence of phylogenetic constraint. Markuelia is an exception to this, known from cleavage and pre-hatchling stages as a vermiform and profusely annulated direct-developing bilaterian with terminal circumoral and posterior radial arrays of spines. Phylogenetic analyses have hitherto suggested assignment to stem-Scalidophora (phyla Kinorhyncha, Loricifera, Priapulida). We test this assumption with additional data and through the inclusion of additional taxa. The available evidence supports stem-Scalidophora affinity, leading to the conclusion that scalidophorans, cyclonerualians, and ecdysozoans are primitive direct developers, and the likelihood that scalidophorans are primitively metameric.     

The phylogenetic significance of bone types in euteleost fishes

The paradentary is a small, sometimes dentigerous element in the lower jaw of some atherinomorph neoteleost fishes. Identification of the paradentary as a neomorphic, perichondrally ossified bone prompted re-examination of theories of the association of bone and teeth in teleost fishes. Teeth on a chondral lower jaw bone might be explained simply by epidermal-mesodermal interactions. Since the work of Kolliker in 1859, it has been known that there are two basic types of bone in teleost fishes: cellular bone, characterized by a matrix that has enclosed osteoblasts or osteocytes; and acellular bone, characterized by a relatively featureless matrix that lacks these bone-forming cells. Cellular bone is typical of lower teleosts, whereas acellular bone is typical of higher teleosts. Ontogenetic data indicate that acellular bone is derived relative to cellular bone. Even though identification of cellular and acellular bone can be made readily with histological preparations, acellular bone has been used infrequently as a character in analyses of teleost phylogeny. Acellular bone is considered here to be a derived character within teleost fishes. It is found in all Neoteleostei as well as some, but not all Salmoniformes. Independent of studies of bone, derived types of teeth in teleosts have been described in terms of their failure to become completely mineralized. Acellular bone and teeth of higher teleosts share several properties, including a large fraction of collagen. Teleosts lack a parathyroid gland; bone type is critical to the mechanism of calcium regulation. It is proposed that the character of acellular bone be incorporated into phylogenetic analyses of teleost fishes by correlating it with derived types of tooth structure.     

The phylogenetic significance of sperm morphology in the Platyhelminthes

The phylogenetic significance of flatworm sperm morphology is discussed against the background of general spermatology. The modified type of spermatozoon of the Nemertodermatida, a group of primitive flatworms, indicates that the Platyhelminthes evolved from forms characterized by the primitive type of metazoan sperm and by the primitive mode of fertilization, implying the release of sperm freely into sea water.The occurrence of aberrant types of spermatozoa in most platyhelminths is obviously a consequence of early evolution of the internal mode of fertilization, which characterizes all true members of this group. It can be concluded, from the ultrastructure of these aberrant spermatozoa that higher metazoans cannot have evolved from seriated flatworms related to the recent Seriata (Proseriata and Tricladida). Even the seemingly primitive Acoela have such aberrant spermatozoa that evolution of higher metazoans from acoels related to the recent Acoela seems highly improbable.The ultrastructure of the spermatozoa of the parasitic groups of flatworms (Monogenea, Digenea, Cestoda) is very similar to that found in the Kalyptorhynchia, a further indication that the parasitic groups are related to the rhabdocoel turbellarians.     

The sound-transmitting apparatus in primitive snakes and its phylogenetic significance

Summary The structure of the sound-transmitting apparatus in primitive snakes (Scolecophidia, Henophidia) is reviewed and compared with that of advanced snakes (Caenophidia) and of some fossorial lizards. Assuming a constant course of the chorda tympani, the ophidian stylohyal can be homologized with the intercalary cartilage of lizards while the cartilaginous distal portion of the ophidian stapes represents the internal process. The cladistic significance of the stapes-quadrate-articulation in the Henophidia and Caenophidia is discussed. The Booidea and the Caenophidia show a shift of the stapes-quadrate-articulation which is correlated with changes in the suspensorium as an adaptation to relatively larger prey. However, convergence cannot be ruled out. Dibamus is shown to be the only lizard known so far which approaches the ophidian middle ear structure. Convergence has to be assumed since there is no sign of a stylohyal in Dibamus and since the course of the ramus communicans externus n. facialis cum glossopharyngeo supports the hypothesis that snakes are to be derived from a pre-lacertilian stage of lepidosaurian evolution.

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Zusammenfassung Die Struktur des schalleitenden Apparates im Mittelohr primitiver Schlangen (Scolecophidia, Henophidia) wird beschrieben und mit höheren Schlangen (Caenophidia) sowie mit einigen grabenden Lacertiliern verglichen. Unter der Annahme eines konstanten Verlaufes der Chorda tympani lässt sich das Stylohyale der Schlangen mit dem Processus dorsalis homologisieren, während das knorpelige distale Ende des Stapes der Schlangen dem Processus internus entspricht. Die kladistischen Implikationen der Struktur des Mittelohres werden diskutiert. Die Booidea und die Caenophidia zeigen eine Verschiebung das Stapes-Quadratum-Gelenkes, welche möglicherweise mit Änderungen der Proportionen des Suspensoriums als Anpassung an relativ grössere Beute zusammenhängt. Konvergenz kann hierbei nicht ausgeschlossen werden. Dibamus ist die einzige bislang bekannte Echse welche im Bau des Mittelohres den Schlangen nahe kommt. Allerdings ist Konvergenz anzunehmen, da Dibamus keinerlei Hinweis auf das Vorhandensein eines Stylohyale liefert, und da der Verlauf des Ramus communicans externus n. facialis cum glossopharyngeo die Hypothese stützt, dass Schlangen nicht von rezenten Unterordnungen der Lacertilier abgeleitet werden können.

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Abbreviations (used in the figures) chr chorda tympani - ci carotis interior - crst cartilaginous distal extension of the stapedial shaft - dper ductus perilymphaticus - fc carotis facialis - par articulatory process of quadrate - pt pterygoid - q quadrate - rce ramus communicans externus - rci ramus communicans internus - st supratemporal - stp stapes - sty stylohyal - vcl vena capitis lateralis - lX n. glossopharyngeus - Vll_hy ramus hyomandibularis of facial nerve     

The origin of the coelom in Brachiopoda and its phylogenetic significance

The origin of the mesoderm and the subsequent formation of the coelom in the larvae of the brachiopod species Notosaria nigricans and Calloria inconspicua is documented in detail at the ultrastructural level. During gastrulation, the blastocoel is completely displaced by the invaginating archenteron. Initial mesoderm formation was observed in late wedge-shaped to early three-lobed stages in both species. Proliferation of mesodermal cells from the archenteral epithelium mainly occurs in the dorsolateral (C. inconspicua) and caudolateral (N. nigricans) parts of the archenteral wall. Thus, a compact mesodermal cell mass pushes its way towards the subepidermal basal lamina. During further development of the larva, the mesoderm is separated from the archenteral epithelium by an extracellular matrix secreted frontad from behind. As a result, a single coelomic anlage is formed. The initial mesoderm in both species is of archenteral/endodermal origin. Considering endodermal origin as the crucial character for enterocoely, coelom formation through proliferation of a compact, endodermally derived mesodermal cell mass in Brachiopoda is clearly identified as enterocoely. Endodermal origin of mesoderm and, therefore, of the coelomic epithelium is hypothesised as a synapomorphy of Brachiopoda and Deuterostomia. As a consequence: (1) Brachiopoda and Deuterostomia are considered sister groups, (2) Brachiopoda group within Radialia and (3) lophophorates (”Tentaculata”) remain as a paraphyletic grouping. Accepted: 26 November 1999     

The phylogenetic significance of colour patterns in marine teleost larvae

Ichthyologists, natural‐history artists, and tropical‐fish aquarists have described, illustrated, or photographed colour patterns in adult marine fishes for centuries, but colour patterns in marine fish larvae have largely been neglected. Yet the pelagic larval stages of many marine fishes exhibit subtle to striking, ephemeral patterns of chromatophores that warrant investigation into their potential taxonomic and phylogenetic significance. Colour patterns in larvae of over 200 species of marine teleosts, primarily from the western Caribbean, were examined from digital colour photographs, and their potential utility in elucidating evolutionary relationships at various taxonomic levels was assessed. Larvae of relatively few basal marine teleosts exhibit erythrophores, xanthophores, or iridophores (i.e. nonmelanistic chromatophores), but one or more of those types of chromatophores are visible in larvae of many basal marine neoteleosts and nearly all marine percomorphs. Whether or not the presence of nonmelanistic chromatophores in pelagic marine larvae diagnoses any major teleost taxonomic group cannot be determined based on the preliminary survey conducted, but there is a trend toward increased colour from elopomorphs to percomorphs. Within percomorphs, patterns of nonmelanistic chromatophores may help resolve or contribute evidence to existing hypotheses of relationships at multiple levels of classification. Mugilid and some beloniform larvae share a unique ontogenetic transformation of colour pattern that lends support to the hypothesis of a close relationship between them. Larvae of some tetraodontiforms and lophiiforms are strikingly similar in having the trunk enclosed in an inflated sac covered with xanthophores, a character that may help resolve the relationships of these enigmatic taxa. Colour patterns in percomorph larvae also appear to diagnose certain groups at the interfamilial, familial, intergeneric, and generic levels. Slight differences in generic colour patterns, including whether the pattern comprises xanthophores or erythrophores, often distinguish species. The homology, ontogeny, and possible functional significance of colour patterns in larvae are discussed. Considerably more investigation of larval colour patterns in marine teleosts is needed to assess fully their value in phylogenetic reconstruction. © 2013 The Authors. Zoological Journal of the Linnean Society published by John Wiley & Sons Ltd on behalf of The Linnean Society of London     

The yeastKluyveromyces africanus nov. spec. and its phylogenetic significance

Summary A new budding yeast species isolated from soil is described. Its outstanding features are, firstly, the formation of asci containing up to sixteen long oval to reniform ascospores and, secondly, a fermentative as well as oxidative metabolism. The assimilation of nitrate is absent and no pseudomycelium is formed. The taxonomic position of the yeast is discussed and it is pointed out that, due to its exceptional ascospore number (1–16), it cannot be classified in any of the existing fermentative genera of theEndomycetaceae (in sensu Lodder et Kreger-van Rij). The reniform shape of its ascospores indicates, however, its close relationship with the multispored genusKluyveromyces, on the one hand, and the newly proposed one to four-spored genusDekkeromyces on the other. The species is provisionally classified as aKluyveromyces species,Kluyveromyces africanus nov. spec., until further information regarding its sexual characteristics becomes available. By virtue of its more or less intermediate ascospore number, it establishes the direct derivation of the genusDekkeromyces fromDipodascus uninucleatus via the multispored yeast genusKluyveromyces.     

The mammalian centrosome and its functional significance

Primarily known for its role as major microtubule organizing center, the centrosome is increasingly being recognized for its functional significance in key cell cycle regulating events. We are now at the beginning of understanding the centrosome's functional complexities and its major impact on directing complex interactions and signal transduction cascades important for cell cycle regulation. The centrosome orchestrates entry into mitosis, anaphase onset, cytokinesis, G1/S transition, and monitors DNA damage. Recently, the centrosome has also been recognized as major docking station where regulatory complexes accumulate including kinases and phosphatases as well as numerous other cell cycle regulators that utilize the centrosome as platform to coordinate multiple cell cycle-specific functions. Vesicles that are translocated along microtubules to and away from centrosomes may also carry enzymes or substrates that use centrosomes as main docking station. The centrosome's role in various diseases has been recognized and a wealth of data has been accumulated linking dysfunctional centrosomes to cancer, Alstrom syndrome, various neurological disorders, and others. Centrosome abnormalities and dysfunctions have been associated with several types of infertility. The present review highlights the centrosome's significant roles in cell cycle events in somatic and reproductive cells and discusses centrosome abnormalities and implications in disease.     

Divided and undivided compound eyes in Ascalaphidae (Insecta, Neuroptera) and their functional and phylogenetic significance

The external morphology of the compound eyes of 13 species of the Ascalaphidae family (Insecta, Neuroptera) from Africa, Asia and Europe was studied in relation to the habitat, phylogeny and time of activity during the day. The six species with undivided eyes (Haplogleniinae) are nocturnal; four inhabit more or less open terrain, while two inhabit more or less dense vegetation. Of the seven species with divided eyes (Ascalaphinae), three are diurnal, one is crepuscular and nocturnal, and three are nocturnal. It was found that two of the diurnal species inhabit open terrain and open forest, and one inhabits dense vegetation; the crepuscular and nocturnal species inhabits open terrain; and two of the nocturnal species inhabit open terrain, while one inhabits dense vegetation. The results are discussed in relation to the hypothesis that divided eyes evolved from undivided eyes, originally serving as an adaptation to daytime vision in open terrain.     

Embryology of the Turbellaria and its phylogenetic significance

Developmental characters — including oocyte and yolk cell structure, patterns of cleavage, and modes of gastrulation — are presented and examined in relation to the phylogeny of the Turbellaria. Eggshell granules, which have been demonstrated to occur in the oocytes of entolecithal eggs and the yolk cells of ectolecithal eggs, are compared among species, and their potential value as a taxonomic character is discussed. The quartet 4d spiral cleavage of the entolecithal egg of polyclads is described as reminiscent of the primitive pattern of early development for the Turbellaria. This is compared to duet spiral cleavage of acoels, and possible phylogenetic schemes involving the two types of spiral cleavage are reviewed. The link between the precise spiral cleavage, which characterizes development of most archoophorans, and blastomere separation (Blastomeren-Anarchie), which occurs in several neoophoran orders, is established by the occurrence of quartet 4d spiral cleavage in one neoophoran order, and of both quartet spiral cleavage and Blastomeren-Anarchie in different species of a second neoophoran order. The epibolic gastrulation of polyclads is described as primitive for the Turbellaria because of its similarity to that of other members of the Spiralia. Although no identical process occurs in neoophoran development, the earlier event of formation of the hull membrane in some neoophorans, and the later event of formation of the definitive epidermis in all neoophorans studied are presented as processes of possible homology to the epibolic gastrulation of polyclads. The lack of correspondence between polyclads and neoophorans in the relationship of the definitive body axes to the egg axis is discussed, and an hypothesis is advanced to account for the differences. The phylogenetic relationships indicated by known developmental phenomena differ only slightly from the scheme presented by Karling in 1974.     

The evolution and functional significance of incest avoidance

The origin and evolution of outbreeding is discussed in the light of the most recent genetical observations and theoretical developments in ethology. Rather than problems resulting from the expression of recessive deleterious genes, small-population genetic stochastic processes and the resulting homosis are viewed as the primary fitness-limiting features of inbreeding populations. With evolution of biparental reproduction, selection favored the reproductive potential of individuals exhibiting a genetically adopted, behavioral aversion to mating with parents, siblings, and offspring. The human incest taboo is regarded as a cultural epiphenomenon following from the general biological inclination to outbreed.