The systematics of Rigidella (Iridaceae)

Rigidella and the four species contained therein are defined primarily on characters related to, and a direct consequence of, adaptation to hummingbird pollination. The genus is probably derived from either the “multiflora” complex inTigridia subgenusHydrotaenia or is an evolutionary line parallel toTigridia. Illustrations, distribution maps, and a key to the species are included. The rediscovery ofRigidella flammea Lindl. is reported, and a new species,R. inusitata, is described.     

Claims and limits of phylogenetic systematics1

Within the methodology of phylogenetic systematics four hierarchic levels are distinguished: the “Central Claim” (to reconstruct phylogeny), methodoloical postulate (to conclude analysis with a purely dichotomous cladogram if ever possible), method (search for sister-group relationships by character analysis), and “Taxonomic Principle” (establishment of a classification reflecting merely the recognized genealoy). Certain limits of applicability and reliability of traditional phylogenetic systematics are specified: genealogy can only be analysed among taxa with perceptible evolutionary novelties; reticulated genealogy is not yet regarded; events other than cladogenetic ones cannot be recognised. Phylogenetic systematics is an independent method which has not been absorbed by any type of “pattern” or “transformed” cladism. Phylogenetic systematics relies on the theory of evolution, which does not lead into circularity, since phylogenetic systematics does not claim to prove or to explain evolution whatsoever.     

Biochemical systematics of the Savi's bat (Hypsugo savii) (Chiroptera: Vespertilionidae)

In order to assess the controversial systematic position of the Savi's bat (Hypsugo savii), 43 ves-pertilionid bats belonging to 10 species were analyzed by mean of isozyme electrophoresis. The results were interpreted in two ways: first, we found that the amount of genetic differentiation between Hypsugo and the true Pipistrellus (D # 0.719) is of the same magnitude as the mean distance which separates the other pipistrelloid genera (D = 0.738). Secondly, the cladogram based on these biochemical results confirms the splitting of these “pipistrelles”, as Hypsugo is more closely related to Vespertilio than to any other palearctic genus. The other species analyzed in this study group themselves in a different clade, including Pipistrellus, Eptesicus and Nyctalus. This phylogenetic interpretation presents a testable hypothesis, which could be confirmed by an analysis of other loci or other allied species.     

Evolutionary theory and systematics: relationships between process and patterns

The relevance of the Modern Evolutionary Synthesis to the foundations of taxonomy (the construction of groups, both taxa and phyla) is reexamined. The nondimensional biological species concept, and not the multidimensional, taxonomic, species notion which is based on it, represents a culmination of an evolutionary understanding. It demonstrates how established evolutionary mechanisms acting on populations of sexually reproducing organisms provide the testable ontological basis of the species category. We question the ontology and epistemology of the phylogenetic or evolutionary species concept, and find it to be a fundamentally untenable one. We argue that at best, the phylogenetic species is a taxonomic species notion which is not a theoretical concept, and therefore should not serve as foundation for taxonomic theory in general, phylogenetics, and macroevolutionary reconstruction in particular. Although both evolutionary systematists and cladists are phylogeneticists, the reconstruction of the history of life is fundamentally different in these two approaches. We maintain that all method, including taxonomic ones, must fall out of well corroborated theory. In the case of taxonomic methodology the theoretical base must be evolutionary. The axiomatic assumptions that all phena, living and fossil, must be holophyletic taxa (species, and above), resulting from splitting events, and subsequently that evaluation of evolutionary change must be based on a taxic perspective codified by the Hennig ian taxonomic species notion, are not testable premises. We discuss the relationship between some biologically, and therefore taxonomically, significant patterns in nature, and the process dependence of these patterns. Process-free establishment of deductively tested “genealogies” is a contradiction in terms; it is impossible to “recover” phylogenetic patterns without the investment of causal and processual explanations of characters to establish well tested taxonomic properties of these (such as homologies, apomorphies, synapomorphies, or transformation series). Phylogenies of either characters or of taxa are historical-narrative explanations (H-N Es), based on both inductively formulated hypotheses and tested against objective, empirical evidence. We further discuss why construction of a “genealogy”, the alleged framework for “evolutionary reconstruction”, based on a taxic, cladistic outgroup comparison and a posteriori weighting of characters is circular. We define how the procedure called null-group comparison leads to the noncircular testing of the taxonomic properties of characters against which the group phylogenies must be tested. This is the only valid rooting procedure for either character or taxon evolution. While the Hennig -principle is obviously a sound deduction from the theory of descent, cladistic reconstruction of evolutionary history itself lacks a valid methodology for testing transformation hypotheses of both characters and species. We discuss why the paleontological method is part of comparative biology with a critical time dimension ana why we believe that an “ontogenetic method” is not valid. In our view, a merger of exclusive (causal and interactive, but best described as levels of organization) and inclusive (classificatory) hierarchies has not been accomplished by a taxic scheme of evolution advocated by some. Transformational change by its very nature is not classifiable in an inclusive hierarchy, and therefore no classification can fully reflect the causal and interactive chains of events constituting phylogeny, without ignoring and contradicting large areas of corroborated evolutionary theory. Attempts to equate progressive evolutionary change with taxic schemes by Haeckel were fundamentally flawed. His ideas found 19th century expression in a taxic perception of the evolutionary process (“phylogenesis”), a merger of typology, hierarchic and taxic notions of progress, all rooted in an ontogenetic view of phylogeny. The modern schemes of genealogical hierarchies, based on punctuation and a notion of “species” individuality, have yet to demonstrate that they hold promise beyond the Haeckel ian view of progressive evolution.     

Molecular systematics of Iridaceae: evidence from four plastid DNA regions

Iridaceae are one of the largest families of Lilianae and probably also among the best studied of monocotyledons. To further evaluate generic, tribal, and subfamilial relationships we have produced four plastid DNA data sets for 57 genera of Iridaceae plus outgroups: rps4, rbcL (both protein-coding genes), the trnL intron, and the trnL-F intergenic spacer. All four matrices produce similar although not identical trees, and we thus analyzed them in a combined analysis, which produced a highly resolved and well-supported topology, in spite of the fact that the partition homogeneity test indicated strong incongruence. In each of the individual trees, some genera or groups of genera are misplaced relative to morphological cladistic studies, but the combined analysis produced a pattern much more similar to these previous ideas of relationships. In the combined tree, all subfamilies were resolved as monophyletic, except Nivenioideae that formed a grade in which Ixioideae were embedded. Achlorophyllous Geosiris (sometimes referred to Geosiridaceae or Burmanniaceae) fell within the nivenioid grade. Most of the tribes were monophyletic, and Isophysis (Tasmanian) was sister to the rest of the family; Diplarrhena (Australian) fell in a well-supported position as sister to Irideae/Sisyrinchieae/Tigridieae/Mariceae (i.e., Iridoideae); Bobartia of Sisyrinchieae is supported as a member of Irideae. The paraphyly of Nivenioideae is suspicious due to extremely high levels of sequence divergence, and when they were constrained to be monophyletic the resulting trees were only slightly less parsimonious (<1.0%). However, this subfamily also lacks clear morphological synapomorphies and is highly heterogeneous, so it is difficult to develop a strong case on nonmolecular grounds for their monophyly.     

Towards a better understanding of the higher systematics of Nymphalidae (Lepidoptera: Papilionoidea)

Research on the molecular systematics of higher taxa in the butterfly family Nymphalidae (Lepidoptera) is only just beginning. Outgroup selection is difficult at the moment due to the lack of consensus on the basal relationships of the major groups in Nymphalidae. We identify four major clades in the Nymphalidae based on a cladistic analysis of one mitochondrial gene sequence (COI, 1450 bp) and two nuclear gene sequences (EF-1alpha, 1064 bp, and wingless, 412-415 bp) from 54 exemplar species sampled from all currently recognized subfamilies. The COI data set was found to be highly incongruent with the nuclear data sets and a Partitioned Bremer Support analysis shows that the COI data set largely undermines support for most clades. Transitions at the third codon positions of the COI data set were highly saturated, but analyzing the combined data set with the COI third positions removed did not change the results. The major clades we found are termed the danaine clade (including Danainae), the satyrine clade (including Charaxinae, Satyrinae, Calinaginae, and Morphinae), the heliconiine clade (including Heliconiinae and Limenitidinae excluding Biblidini, Cyrestini, Pseudergolini, and Coeini) and the nymphaline clade (including Nymphalinae, Apaturinae, and Coeini, Cyrestini, Pseudergolini, and Biblidini from Limenitidinae). The heliconiine and nymphaline clades are sister groups, while the most parsimonious explanation for the combined data set places the danaine clade as the most basal large group of Nymphalidae. Our results give one of the strongest hypotheses for the subfamilial relationships within Nymphalidae. We were able to resolve the polyphyletic nature of Limenitidinae, which we recommend to be split into three subfamilies: Limenitidinae, Biblidinae, and Cyrestinae. The tribe Coeini belongs in Nymphalinae.     

Floral morphology and phylogeny of the Disinae (Orchidaceae)

Previous classification of the Disinae has been based almost entirely on floral morphological data. These data are critically assessed by various methods to determine to what extent they support a classification resolved to sectional level. The variation in the characters relative to the sections and genera is mapped, homologies are established by careful morphological observation and ontogenetic studies, and finally hypotheses of homology are tested by congruence in several cladistic analyseS. It is found that although floral morphology allows the recognition of some groups, for others the results are ambiguouS. Floral morphological data are clearly inadequate to establish the taxonomy of the Disinae on a sound footing.     

Anatomy and systematics of bathyphytophilid limpets (Mollusca, Archaeogastropoda) from the northeastern Pacific

Bathyphytophilus diegensis sp. n. is described on basis of shell and radula characters. The radula of another species of Bathyphytophilus is illustrated, but the species is not described since the shell is unknown. Both species feed on detached blades of the surfgrass Phyllospadix carried by turbidity currents into continental slope depths in the San Diego Trough. The anatomy of B. diegensis was investigated by means of semithin serial sectioning and graphic reconstruction. ‘The shell is limpet-like; the protoconch resembles that of pseudococculinids and other lepetclloids. The radula is a distinctive, highly modified rhipidoglossate type with close similarities to the lepetellid radula. The anatomy falls well into the lepetelloid bauplan and is in general similar to that of Pseudococculinidae and Pyropcltidae. Apomorphic features are the presence of gill-leaflets at both sides of the pallial roof (shared with certain pseudococculinids), the lack of jaws, and in particular many enigmatic pouches (bacterial chambers?) which open into the postcrior oesophagus. Autapomorphic characters of shell, radula and anatomy confirm the placement of Bathyphytophilus (with Aenigmabonus) in a distinct family, Bathyphytophilidac Moskalev, 1978. As revealed by a cladistic study, the Bathyphytophilidae should be classified within the Lepetelloidea close to the Lepetellidae, Pyropeltidae, and Pseudococculinidae.     

Congruence between phylogenetic and stratigraphic data on the history of life

The quality of the fossil record and the accuracy of reconstructed phylogenies have been debated recently, and doubt has been cast on how far current knowledge actually reflects what happened in the past. A survey of 384 published cladograms of a variety of animals (echinoderms, fishes, tetrapods) shows that there is good agreement between phylogenetic (character) data and stratigraphic (age) data, based on a variety of comparative metrics. This congruence of conclusions from two essentially independent sources of data confirms that the majority of cladograms are broadly accurate and that the fossil record, incomplete as it is, gives a reasonably faithful documentation of the sequence of occurrence of organisms through time.     

Further revisions of the Sabellidae subfamilies and cladistic relationships among the Fabriciinae (Annelida: Polychaeta)

Cladistic relationships of the sabellid subfamilies Fabriciinae and Sabellinae are examined in light of recent revisions of fabriciin taxa. The potential placement of Caobangia in the Sabellinae is suggested from an initial analysis of selected fabriciin species and genera. Subsequent cladistic analyses at the family level produced over 1800 cladograms, among which Caobangia is the most plesiomorphic taxon of either the Fabriciinae or Sabellinae. Successive approximations weighting reduced this ambiguity, consistently placing Caobangia in the Sabellinae. Subfamilies are emended on the basis of these results. Cladistic relationships of fabriciin taxa, exclusive of Caobangia , display topological instability among some genera and species. Genera are, however, monophyletic in all topologies. Incorporating ontogenetic data for branching ventral filamentous appendages in Augeneriella reduces ambiguity among genera and suggests alternative transformation series for ventral filamentous appendages.     

蜡梅科植物的分支分析

蜡梅科是一个仅有４属,１０种的小科,将蜡梅科的生物信息数字化,利用徐克学的和谐性分析程序,剔除了不合理的性状安排,判别关系含糊的性状极性,利用最大同步法,最小平行演化法及最大离散量分支分类法,对由性状再分析后获得的数值矩阵进行运算,推导分支图,明确各属之间的发生、发展和演化的关系。结果表明：椅子树亚科（Ｉｄｉｏｓｐｅｒｍｏｉｄｅａｅ）的椅子树属（Ｉｄｉｏｓｐｅｒｍｕｍ Ｂｌａｋｅ）在整个蜡梅科（Ｃ