Exoskeleton and Systematics: A Historical Problem in the Classification of Glyptodonts

The glyptodonts (Mammalia: Cingulata) are characterized by an exoskeleton comprising most notably an armored tail and an immobile dorsal carapace formed by a large number of osteoderms. In 1889, Florentino Ameghino published the first phylogenetic scenario for the glyptodonts, based on the sequential application of two transformation series related to the morphology of the tail armor and carapace osteoderms. From the early to mid 1900s, several authors used Ameghino’s transformation series subordinated to a model of evolution in which derived glyptodont groups had arisen independently from separate pre-middle Miocene ancestors. This approach, in which the morphological states of Ameghino’s series were considered analogous rather than homologous, provided different phylogenetic scenarios and the paraphyletic classification still in use. Two recent cladistic analyses based on cranial and postcranial (including caudal tube) characters support the monophyly of glyptodonts and suggest novel intra-clade relationships. However, neither analysis included the classic osteoderm characters used by earlier authors. Therefore, we propose new osteoderm and carapace characters and evaluate their performance in a new cladistic analysis. We found that: a) some osteoderm characters used by earlier authors to support ancestor-descendent hypotheses are in fact fully homoplastic autapomorphies (e.g., multiplication of the number of rows of peripheral figures); b) characters previously believed to have originated independently in several groups (e.g., presence of caudal tube) are synapomorphies at a wider hierarchical level; c) some ancestor–descendant pre-cladistic hypotheses are incompatible with the topology and synapomorphy distribution obtained; and d) there is no reason to favor exoskeletal characters in glyptodont systematics.     

Comparative detailed morphology of the Heteroderinae Filip'ev & Schuurmans Stekhoven, 1941, sensu Luc et al. (1988): phylogenetic systematics and revised classification

Taxonomic schemes for the Heteroderinae Filip'ev & Schuurmans Stekhoven, 1941, sensu Luc et al., (1988) have been unstable due to the large number of genera and the paucity of known reliable characters. Reliable characters are essential when using phylogenetic inference in developing a natural classification. Morphological and developmental studies using light, scanning and transmission electron microscopy have revealed the new characters of host response, en face patterns, phasmid structure and female cuticular layers. These techniques also gave us insight into the homoplasy and polarity of many characters, revealed previously undetected character states and clarified misinterpreted character states. A matrix with the 19 most reliable characters is proposed for 20 operational taxonomic units (OTUs) and we employ this matrix for comparing computer generated phylogenetic analyses of the PHYLIP and PAUP packages. PAUP was deemed the more reliable parsimony algorithm for phylogenetic analysis of the Heteroderinae (Fink, 1986; Platnick, 1987). Monophyly of Atalodera + Sherodera + Thecavermiculatus (tribe Ataloderini), and Cactodera + Heterodera + Afenestrata, as well as Punctodera + Globodera + Dolichodera is supported by both programs. Most importantly, analyses strongly support monophyly of all cyst-forming genera (tribe Heteroderini) contrary to previous hypotheses of repeated evolution of the cyst (Wouts, 1985). In addition, monophyly of the Heteroderini with the Ataloderini is demonstrated. PAUP indicates monophyly of Sarisodera + Rhizonema + Bellodera + Hylonema and Ekphymatodera (tribe Sarisoderini new rank). Monophyly of the Sarisoderini was at first only weakly supported, but, subsequently, the reduced width of the submedial lips of second stage juveniles and males was recognized as a synapomorphy which strengthened subsequent PAUP trees and monophyly of the tribe. The present study rejects as paraphyletic or polyphyletic several previously proposed combinations, including Thecavermiculatus sequoiae (versus Rhizonema sequoiae), Sarisodera africana (versus Afenestrata africana), Dolichodera andinus (versus Thecavermiculatus andinus). The question whether T. andinus is a distinct genus, was not resolved due to insufficient data. PAUP supports our previous observations that Cactodera betulae is intermediate in a transformation series between other Cactodera and Heterodera: it also indicates these species as bring monophyletic with Heterodera + Afenestrata, but not with other Cactodera. Although these phylogenetic analyses strongly support some relationships, they indicate unresolved alternative hypotheses for others. Meloidodera (tribe Meloidoderini) and Cryphodera (tribe Cryphoderini) must be investigated for consideration of a possible synapomorphy not included in the present data matrix. Future studies are proposed to more clearly define the monophyly of the Heteroderini, as well as the Sarisoderini. Tests are also proposed to clarify questions of the monophyly of Verutus (tribe Verutini new rank) with the Heteroderinae versus other Tylenchida.     

On the position of the digenean family Heronimidae: an inquiry into a cladistic classification of the Digenea

Brooks, O'Grady & Glen (1985b) placed the seemingly aberrant and highly derived family Heronimidae at the base of their cladogram of the Digenea. In the absence of arguments for the composition and polarity of the putative homologous series on which their cladogram is based, it was found necessary to consider in detail almost all of their character series. In the course of my analysis I (1) argue that the oral sucker is singular to the Digenea, and that the oral sucker is not synapomorphic nor the ventral sucker symplesiomorphic for the Digenea, but that early digeneans lacked both suckers, (2) reiterate that the amphistomate condition is secondary and has arisen more than once, and that the embryology of the excretory system proves it to be secondary in all cases, (3) offer support for Cable's view that the cercarial stylet is not of single origin, and (4) restate the argument that paramphistomes, microscaphidiids, and gyliauchenids have not an oral sucker but a pharynx. Brooks, Bandoni, Macdonald & O'Grady (1989) stated that the bifurcate gut is unambiguously a synapomorph of the Digenea, on the basis of their cladistic demonstration that Rugogaster with a bifurcate gut is a sister group to the rest of the Aspidobothria, which in turn is a sister group to the digenea. Testing this in a new cladogram of the Aspidobothria constructed from Brooks et al.'s (1989) 15 characters (corrected, especially those of the ventral sucker, for which a new phylogeny is advanced), supplemented by an additional 23 characters, reveals Rugogaster as the derived form it appeared to be and aspidogastrids, with a single caecum, as sister group to the rest of the Aspidobothria. A new analysis of the characters produced an increase in homoplasy from 18% to 37%, showed 97 out of 212 characters invalid as used, these two resulting in a decrease in resolution from 82% to 32%, and resulted in changes in polarity in 20 of the 65 putative homologous series listed by Brooks et al. (1985b). The Heronimidae cannot be at the base of their cladogram as it has not three but 13 apomorphs, which is more than have the next three groups, the paramphistomiforms, echinostomatiforms and haploporiforms. Comparison of the Heronimidae with the Bivesiculidae in the light of 48 defensible plesiomorphs shows that the Bivesiculidae, which have 39 of the 48 characters, are more plesiomorphic than the Heronimidae, which have only 23. In conclusion, what remains of the data-base is insufficient to support their cladogram and classification. Better data from the flukes themselves are necessary for a sound cladistic analysis.     

Preliminary morphological analysis of relationships between the spider wasp subfamilies (Hymenoptera: Pompilidae): revisiting an old problem

No qualitative cladistic analysis has been performed previously for the subfamily classification of Pompilidae (Hymenoptera). In 1994 Shimizu proposed six subfamilies, but their validity and relationships remain inconclusive. The objective of this study was to perform a quantitative analysis of phylogenetic relationships of the Pompilidae, with emphasis on testing the validity of proposed subfamilies. Two cladistic analyses were performed based on morphological evidence. First, a maximum-parsimony analysis of Shimizu's original morphological data matrix (72 taxa by 54 characters) was conducted, with the data subjected to a heuristic search for the first time with phylogenetic software. The resulting strict-consensus cladogram yielded a monophyletic Ceropalinae that was sister group to a large polytomy containing members of the remaining five subfamilies. In a second analysis, several of Shimizu's characters were re-examined, and new characters and more taxa were added to the data set. Terminal taxa were coded as species rather than as generic abstractions, and 20 additional morphological characters were introduced. The analysis was based on 77 morphological characters derived from the adults of 84 taxa. This second analysis suggested that Notocyphinae sensu Shimizu (1994) was nested within Pompilinae and that Epipompilinae sensu Shimizu (1994) was nested within Ctenocerinae; neither should retain their status as a separate subfamily. Lastly, Chirodamus s .s., which historically has been a member of the Pepsinae, is placed within the Pompilinae with reservations rather than erecting a new subfamily. After these allowances were made, a strict consensus tree gave the following relationships: (Ceropalinae + (Pepsinae + (Ctenocerinae + Pompilinae))).     

Changing the landscape: a new strategy for estimating large phylogenies

In this paper we describe a new heuristic strategy designed to find optimal (parsimonious) trees for data sets with large numbers of taxa and characters. This new strategy uses an iterative searching process of branch swapping with equally weighted characters, followed by swapping with reweighted characters. This process increases the efficiency of the search because, after each round of swapping with reweighted characters, the subsequent swapping with equal weights will start from a different group (island) of trees that are only slightly, if at all, less optimal. In contrast, conventional heuristic searching with constant equal weighting can become trapped on islands of suboptimal trees. We test the new strategy against a conventional strategy and a modified conventional strategy and show that, within a given time, the new strategy finds trees that are markedly more parsimonious. We also compare our new strategy with a recent, independently developed strategy known as the Parsimony Ratchet.     

Spermatozoa of tapeworms (Platyhelminthes,Eucestoda): advances in ultrastructural and phylogenetic studies

New data on spermiogenesis and the ultrastructure of spermatozoa of ‘true’ tapeworms (Eucestoda) are summarized. Since 2001, more than 50 species belonging to most orders of the Eucestoda have been studied or reinvestigated, particularly members of the Caryophyllidea, Spathebothriidea, Diphyllobothriidea, Bothriocephalidea, Trypanorhyncha, Tetraphyllidea, Proteocephalidea, and Cyclophyllidea. A new classification of spermatozoa of eucestodes into seven basic types is proposed and a key to their identification is given. For the first time, a phylogenetic tree inferred from spermatological characters is provided. New information obtained in the last decade has made it possible to fill numerous gaps in the character data matrix, enabling us to carry out a more reliable analysis of the evolution of ultrastructural characters of sperm and spermiogenesis in eucestodes. The tree is broadly congruent with those based on morphological and molecular data, indicating that convergent evolution of sperm characters in cestodes may not be as common as in other invertebrate taxa. The main gaps in the current knowledge of spermatological characters are mapped and topics for future research are outlined, with special emphasis on those characters that might provide additional information about the evolution of tapeworms and their spermatozoa. Future studies should be focused on representatives of those major groups (families and orders) in which molecular data indicate paraphyly or polyphyly (e.g. ‘Tetraphyllidea’ and Trypanorhyncha) and on those that have a key phylogenetic position among eucestodes (e.g. Diphyllidea, ‘Tetraphyllidea’, Lecanicephalidea, Nippotaeniidea).     

A classification of the Ericaceae: subfamilies and tribes

The variation shown by 60 characters (morphological, anatomical, embryological, chemical and cytological) is tabulated for the whole of the Ericaceae. The reliability of the characters and the observations on them are discussed, and it is emphasized that many characters are difficult to use because they are rarely restricted to one taxon. Six subfamilies are recognized. Within the Rhododendroideae seven tribes are recognized, the monogeneric Daboecieae being new; Epigaea is transferred to the Rhododendroideae from the Andromedeae and Diplarche from the Diapensiaceae. Calluna is placed in a new monotypic tribe within the Ericoideae. The extended Vaccinioideae includes the Arbutoideae; five tribes are recognized, of which the Enkiantheae and Cassiopeae are new. The Pyroloideae and Monotropoideae are left unchanged: a new subfamily, the Wittsteinioideae, includes the deviant monotypic genus Wittsteinia. A key to subfamilies, and to genera of the Pyroloideae, Rhododendroideae and superior-ovaried Vaccinioideae is given.     

EVOLUTIONARY CHARACTER ANALYSIS: TRACING CHARACTER CHANGE ON A CLADOGRAM

Abstract— There has been little formal discussion concerning character analysis in cladistics, even though characters and their character state trees are central to phylogenetic analyses. We refer to this field as Evolutionary Character Analysis. This paper defines the components of evolutionary character analysis: character state trees, transmodal characters, cladogram characters, attribute and character phylogenies; and the use of these components in phylogenetic inference and evolutionary studies. Character state trees and their effect on cladogram construction are discussed. A new method for numerically coding complex character state trees is described that further reduces the number of variables required to describe them. This method, ordinal coding, reduces the size of data matrices, and facilitates retrieval of state codes. This paper advocates the use of both biological evidence and evidence internal to the cladogram itself to construct character state trees (CSTs). We discuss general models of character evolution (morphocline analysis, Fitch minimum mutation model, etc.) and their role in forming CSTs. Character state trees formed with theories of character evolution are referred to as transmodal characters. These transmodal characters are contrasted with cladogram characters (Mickevich, 1982), and the place of each in a phylogenetic analysis is discussed. The method for determining cladogram characters is detailed with more complicated examples than found in previous publications. We advocate testing transmodal characters by comparing them with the resultant cladogram characters. This comparison involves transformation series analysis (TSA; Mickevich, 1982) which is viewed as an extension of reciprocal illumination. The TSA procedure and its place in hypothesis testing are reviewed. Tracing the evolution of characters interests both systematists and non-systematists alike. When character state trees (transmodal characters) are optimized on pre-existing phylogenies, character phylogenies and attribute phylogenies result. Attributes are defined as a feature that may or may not be homologous (i.e., ecological categories, plant hosts, etc.). We provide two illustrations of this approach, one involving the evolution of the anuran ear and another involving the coevolution of the butterfly Heliconius and its hostplants. Finally, the components of phylogenetic character analysis can be used to test more general evolutionary theories such as the biogenetic law and vicariance biogeography.     

Systematics, cladistics and biogeography of the American genus Farrodes (Ephemeroptera: Leptophlebiidae: Atalophlebiinae)

Cladistic and biogeographic analyses of the genus Farrodes are presented. Two species groups are delineated within Farrodes: F. caribbianus and F. bimaculatus. Three species formerly placed in other genera– Thraulus caribbianus Traver, Thraulus roundsi Traver and Homothraulus maculatus (Needham & Murphy)–are transferred to Farrodes. The species of the F. caribbianus species group are revised. Three new species are described: F. savagei from Venezuela, F. maya and F. mexicanus from Mexico. Keys to separate the two species groups of Farrodes and the species of the F caribbianus species group are provided. Successive cladistic analyses were carried out on both adult and nymphal characters using Hennig86 and CLADOS. The matrix was composed of all available data (nymphal characters were missing for some species), from nymphal and adult stages separately and on taxa represented by both adult and nymphal characters. Species of the genera Simothraulopsis and Homothraulus (components of the Farrodes lineage) were included in the analyses, and Ecuaphkbia was used as the outgroup. Results of the four analyses are compatible. The historical biogeography of Farrodes , with a distribution from nordiern Argentina to southern Texas, is analysed using the program COMPONENT. Areas of endemism are established, and some of their relationships compared with those of other groups available in the literature.     

Clade stability and the addition of data: A case study from erigonine spiders (Araneae: Linyphiidae, Erigoninae)

This study presents a new phylogeny of erigonine spiders with emphasis on genera from the Neotropics. Thirty‐nine exemplar taxa representing mostly Neotropical genera were added to a global sample of 31 erigonine and 12 outgroup exemplar taxa analyzed in a previous study. These 82 taxa were coded for 176 (172 informative) mostly morphological characters. Eighty‐one characters were identical to or modified from the 73 (67 informative) characters included in a previous study; the remaining 95 characters are new. The complete data set includes 70 erigonine exemplars representing 65 genera, seven nonerigonine linyphiid exemplars, and five exemplars representing four araneoid families in the outgroup. Cladistic analysis resulted in a single most parsimonious tree (L =904, CI = 0.23, RI = 0.58; uninformative characters excluded: L = 900, CI = 0.23). This paper explores the implications of the new topology for the evolution of several characters of interest in erigonine evolution. The phylogeny implies that the desmitracheate condition is a synapomorphy of erigonines, with a reversal to the haplotracheate condition in one large clade within Erigoninae. We infer that the loss of the paracymbium in Neotropical erigonines occurred twice and may have progressed by different evolutionary pathways. Our phylogeny differs markedly from the previous cladistic hypothesis of erigonine relationships. We investigate how the addition of characters and taxa (alone and together) have altered the earlier hypothesis of erigonine phylogeny. We conclude that topological changes from the previous study to the current one are largely the result of adding and modifying characters, not adding taxa. Continuous Jackknife Function (CJF) analysis predicts that the inclusion of additional character data will continue to imply changes in the relationships among taxa in our analysis.     

Combinatorial model for the formation of body plans in higher metazoan taxa: Paleontological insight

The body plans of higher metazoan taxa were formed during a short time (on the geological time scale) by combination of the previously developed characters. The combinations were realized as a result of manifestation of latent characters in adults through various heterochronies. This resulted in mosaic evolution and concealment of intermediate forms. Many characters of new body plans appeared in the ancestral taxon and their various combinations in the newly established taxa formed the archaic diversity. The maximum rank of newly appearing higher taxa decreased with geological time. The evolution of metazoans passed from the development of the general body plan to less significant details and appearance of body plans describing taxa of lower ranks. New body plans of higher taxa were superposed on the old body plan rather than replaced it, extending with time the subordination of body plans and respective hierarchy of taxa. Aromorphoses are always connected with the appearance of a new body plan. The appearance of new taxa and an increase in morphological diversity mostly occurred at certain boundaries in the development of the biota, which were connected with a sharp increase in the previously limited resources.     

Phylogeny of the Hymenoptera: a reanalysis of the Ronquist et al. (1999) reanalysis, emphasizing wing venation and apocritan relationships

Ronquist et al . (1999) recoded Rasnitsyn's (1988) analysis to effect an explicit numerical cladistic analysis of his data. Here we examine their analysis and reveal that much of the resolution obtained for apocritan relationships is dependant on reductional wing characters. The wing characters in their matrix are replaced with a revised set of wing characters and reanalysed using strict parsimony. For apocritan taxa the resulting strict consensus tree is considerably less resolved, but perhaps preferable as a more conservative starting point in the continuing investigation of higher level hymenopteran relationships.     

新疆鳞网衣属地衣的初步研究

采用形态解剖、化学等传统分类方法,对新疆天山的鳞网衣属地衣进行了初步研究,发现该属的3个种,其中脑状鳞网衣（Psora cerebriformis W. A. Weber）和小红褐色鳞网衣［Psora luridella (Tuck.) Fink］是中国新记录种。文中对鳞网衣属3个种进行了详细的描述,并提供了相关彩色图片。     

Variability in epidermal characters of <Emphasis Type="Italic">Ginkgo tzagajanica</Emphasis> Samylina (Ginkgoales) from the Paleocene of the Tsagayan formation (Amur Region) and the taxonomy of tertiary species of <Emphasis Type="Italic">Ginkgo</Emphasis>

The study of new material on Ginkgo tzagajanica Samylina and earlier published data has shown that the epidermal morphology of the species in highly variable; several new characters were found. Occurrence of different states of characters was analyzed, their frequency distribution was revealed, and typical and rare states of characters within variation rows were determined. The specific diagnosis of G. tzagajanica was emended. The most variable characters are degree of undulation of the anticlinal walls of cells of abaxial and adaxial epidermises (the latter is more variable) and degree of development and number of papillae on ordinary epidermal cells. Less variable characters are degree of development of papillae on subsidiary cells of stomata and how the papillae cover the stomatal apertures. These are characters that bear maximal diagnostic significance for Cenozoic Ginkgo.     

Inferring hominoid and early hominid phylogeny using craniodental characters: the role of fossil taxa

Recent discoveries of new fossil hominid species have been accompanied by several phylogenetic hypotheses. All of these hypotheses are based on a consideration of hominid craniodental morphology. However, Collard and Wood (2000) suggested that cladograms derived from craniodental data are inconsistent with the prevailing hypothesis of ape phylogeny based on molecular data. The implication of their study is that craniodental characters are unreliable indicators of phylogeny in hominoids and fossil hominids but, notably, their analysis did not include extinct species. We report here on a cladistic analysis designed to test whether the inclusion of fossil taxa affects the ability of morphological characters to recover the molecular ape phylogeny. In the process of doing so, the study tests both Collard and Wood's (2000) hypothesis of character reliability, and the several recently proposed hypotheses of early hominid phylogeny. One hundred and ninety-eight craniodental characters were examined, including 109 traits that traditionally have been of interest in prior studies of hominoid and early hominid phylogeny, and 89 craniometric traits that represent size-corrected linear dimensions measured between standard cranial landmarks. The characters were partitioned into two data sets. One set contained all of the characters, and the other omitted the craniometric characters. Six parsimony analyses were performed; each data set was analyzed three times, once using an ingroup that consisted only of extant hominoids, a second time using an ingroup of extant hominoids and extinct early hominids, and a third time excluding Kenyanthropus platyops. Results suggest that the inclusion of fossil taxa can play a significant role in phylogenetic analysis. Analyses that examined only extant taxa produced most parsimonious cladograms that were inconsistent with the ape molecular tree. In contrast, analyses that included fossil hominids were consistent with that tree. This consistency refutes the basis for the hypothesis that craniodental characters are unreliable for reconstructing phylogenetic relationships. Regarding early hominids, the relationships of Sahelanthropus tchadensis and Ardipithecus ramidus were relatively unstable. However, there is tentative support for the hypotheses that S. tchadensis is the sister taxon of all other hominids. There is support for the hypothesis that A. anamensis is the sister taxon of all hominids except S. tchadensis and Ar. ramidus. There is no compelling support for the hypothesis that Kenyanthropus platyops shares especially close affinities with Homo rudolfensis. Rather, K. platyops is nested within the Homo + Paranthropus + Australopithecus africanus clade. If K. platyops is a valid species, these relationships suggest that Homo and Paranthropus are likely to have diverged from other hominids much earlier than previously supposed. There is no support for the hypothesis that A. garhi is either the sister taxon or direct ancestor of the genus Homo. Phylogenetic relationships indicate that Australopithecus is paraphyletic. Thus, A. anamensis and A. garhi should be allocated to new genera.     

Phylogeny of the family Congiopodidae (Perciformes: Scorpaenoidea), with a proposal of new classification

The phylogenetic relationships of the family Congiopodidae are inferred based on morphological characters. The monophyly of this family is supported by 13 unambiguous apomorphic characters, including four autapomorphies among the superfamily Scorpaenoidea. The Congiopodidae shares 26 apomorphic characters with other scorpaenoid taxa, and these characters are considered to also support the monophyly of the family. Upon completion of the phylogenetic analysis using the characters in 39 transformation series, it was assumed that the family is unambiguously supported by five characters (and also by three and one characters when ACCTRAN and DELTRAN are used, respectively) and is branched into two major clades, including Congiopodus and Alertichthys plus Zanclorhynchus, respectively. Based on the phylogenetic relationships, a new classification, recognizing two subfamilies (Congiopodinae and Zanclorhynchinae) in the family Congiopodidae, is proposed. The genus Perryena, that was recently inferred being closely related to the Tetrarogidae (although many authors included it in the Congiopodidae), is provisionally placed into the Congiopodidae as incertae sedis.