Ingroup Relationships Research Articles

Systematic revision, phylogeny and host plant associations of the Australian endemic genus Mcateella Drake (Hemiptera:Heteroptera:Piesmatidae)

A systematic revision of Mcateella is given including the description of seven new species: M. austera, sp. nov., M. coolgardie, sp. nov., M. esperancensis, sp. nov., M. exocarposa, sp. nov., M. kwoki, sp. nov., M. reidi, sp. nov. and M. schuhi, sp. nov. The following species are redescribed: M. elongata Hacker M. gibber Drake, M. interioris Hacker and M. splendida Drake. Mcateella is defined as a monophyletic group on the basis of clavate guard setae and the stridulatory apparatus. Phylogenetic analysis of the Piesmatidae recognised Eopiesma as the sister-taxon relative to the remaining extant Piesmatidae. The ingroup relationships of Mcateella contained one polytomy. Miespa was found to be the sister taxon to Mcateella, positing a transantarctic relationship. Host plant analysis indicates that Mcateella are not all Acacia specialists as stated in the literature but feed on a wide range of angiosperms.

Corrigendum to: Systematic revision, phylogeny and host plant associations of the Australian endemic genus Mcateella Drake (Hemiptera:Heteroptera:Piesmatidae)

A systematic revision of Mcateella is given including the description of seven new species: M. austera, sp. nov., M. coolgardie, sp. nov., M. esperancensis, sp. nov., M. exocarposa, sp. nov., M. kwoki, sp. nov., M. reidi, sp. nov. and M. schuhi, sp. nov. The following species are redescribed: M. elongata Hacker M. gibber Drake, M. interioris Hacker and M. splendida Drake. Mcateella is defined as a monophyletic group on the basis of clavate guard setae and the stridulatory apparatus. Phylogenetic analysis of the Piesmatidae recognised Eopiesma as the sister-taxon relative to the remaining extant Piesmatidae. The ingroup relationships of Mcateella contained one polytomy. Miespa was found to be the sister taxon to Mcateella, positing a transantarctic relationship. Host plant analysis indicates that Mcateella are not all Acacia specialists as stated in the literature but feed on a wide range of angiosperms.

Extended contact effects as a function of closeness of relationship with ingroup contacts

Using survey data from Catholics and Protestants in Northern Ireland ( N = 428), the authors examined the effects of extended contact via different types of ingroup contacts (neighbors, work colleagues, friends, and family members) and tested whether closeness to ingroup contacts moderated the effects of extended contact on outgroup trust. Results demonstrated that extended contact effects varied as a function of the relationship to ingroup contacts, and that extended contact interacted with closeness ratings in predicting outgroup trust. Consistent with hypotheses, extended contacts via more intimate ingroup relationships (i.e., friends and family) were overall more strongly related to outgroup trust than extended contacts via less intimate ingroup relations (i.e., neighbors and work colleagues). Moreover, within each level of intimacy extended contact was related to outgroup trust only at high, and not at low, levels of rated closeness to ingroup contacts. The theoretical contributions, limitations and practical implications of these findings are discussed.

A taxonomic and phylogenetic re-evaluation of Therizinosauria (Dinosauria: Maniraptora)

Therizinosaurians are among the most poorly understood dinosaurs. Their unusual morphology and fragmentary fossil record has precluded a synthetic understanding of the group since their remains were first discovered over 60 years ago. Although the clade was recently substantiated as a monophyletic group of maniraptoran theropods, little foundational work has been conducted at the species level. A recent plethora of therizinosaurian discoveries – including the most complete primitive and derived members recovered to date – permits an alpha taxonomic and phylogenetic re-evaluation of the clade. The phylogenetic analysis presented is the most comprehensive yet conducted for Therizinosauria, and provides a foundation for scrutinizing previous definitions of Therizinosauria, Therizinosauroidea and Therizinosauridae. Here, support is provided for the maintenance of all three taxa; however, Therizinosauria is redefined and Falcarius is excluded from Therizinosauroidea. In addition, the previously described therizinosauroids, Beipiaosaurus, Enigmosaurus, Suzhousaurus, Segnosaurus and Therizinosaurus, are rediagnosed and photodocumented. In contrast to other analyses, the ingroup topology recovered in this study suggests intermediate (therizinosauroid) status for Neimongosaurus, Erliansaurus and Enigmosaurus (based on relatively primitive pelvic morphology), despite the derived forelimb anatomy evident in the former two taxa. Here, the large-bodied taxa Nothronychus and Nanshiungosaurus brevispinus are recovered as therizinosaurids. This discrepancy indicates a relatively complex pattern of mosaic evolution, which may ultimately be found to correlate with body-size trends in the clade. This work also reviews the chronostratigraphic and biogeographic distribution of therizinosaurian taxa and putatively referred elements and finds no compelling evidence of the clade outside of Asia and North America, nor for the referral of therizinosaurian materials from Kazakhstan to cf. Neimongosaurus. Time calibration of ingroup relationships indicates a pre-Turonian dispersal event is needed to account for the presence of therizinosaurids in the Late Cretaceous of North America and Asia; this conclusion supports previous hypotheses of a Laurasian faunal interchange event during the Albian.

New insights on the phylogenetic relationships of the Proseriata (Platyhelminthes), with proposal of a new genus of the family Coelogynoporidae

Proseriata is a diverse and species-rich taxon of Neoophora (Platyhelminthes). The group is divided into two morphologically well-characterised taxa: Lithophora and Unguiphora. Previous molecular analyses, aimed at ascertaining in-group relationships of the Proseriata, were equivocal in supporting the monophyly of the Lithophora, and of one of its families, the Coelogynoporidae. Here we present a new phylogeny of the Proseriata, based on complete ssrDNA and partial (D1-D6) lsrDNA. The analysis includes 36 proseriate species. 13 new sequences from seven species are used, four of which from species of the Coelogynoporidae and one from a new species of the recently established family Calviriidae. Phylogenetic analysis was performed using Bayesian Inference (BI), maximum likelihood (ML) and maximum parsimony (MP). Clades were considered to have high nodal support if BI posterior probability and ML and MP bootstrap percentages were >90%. The resulting cladogram strongly supports the monophyly of the Lithophora. In addition, the monophyly of the Coelogynoporidae and their inclusion in the Lithophora are corroborated. Within the Lithophora, two sister clades are supported, including Coelogynoporidae+Calviriidae, and Otoplanidae+Archimonocelididae+Monocelididae respectively. In the Coelogynoporidae, there was a poor correspondence between estimates of phylogeny and the monophyly of genera, suggesting the family is in need of systematic revision. Current morphology-based systematic arrangements of Otoplanidae and Monocelididae are not supported by molecular results. The enigmatic taxon Ciliopharyngiella (formerly included in the Rhabdocoela) clusters with the Unguiphora, although with low support. One new species of Coelogynoporidae (Parainvenusta englarorum n. gen n. sp.) is described and discussed in the appendix.

Comparative sperm ultrastructure in Nemertea

Although the monophyly of Nemertea is strongly supported by unique morphological characters and results of molecular phylogenetic studies, their ingroup relationships are largely unresolved. To contribute solving this problem we studied sperm ultrastructure of 12 nemertean species that belong to different subtaxa representing the commonly recognized major monophyletic groups. The study yielded a set of 26 characters with an unexpected variation among species of the same genus (Tubulanus and Procephalothrix species), whereas other species varied in metric values or only one character state (Ramphogordius). In some species, the sperm nucleus has grooves (Zygonemertes virescens, Amphiporus imparispinosus) that may be twisted and give a spiral shape to the sperm head (Paranemertes peregrina, Emplectonema gracile). To make the characters from sperm ultrastructure accessible for further phylogenetic analyses, they were coded in a character matrix. Published data for eight species turned out to be sufficiently detailed to be included. Comparative evaluation of available information on the sperm ultrastructure suggests that subtaxa of Heteronemertea and Hoplonemertea are supported as monophyletic by sperm morphology. However, the data do not provide information on the existing contradictions regarding the internal relationships of "Palaeonemertea." Nevertheless, our study provides evidence that sperm ultrastructure yields numerous potentially informative characters that will be included in upcoming phylogenetic analyses.

Paraphyly in the Bamboo Subtribe Chusqueinae (Poaceae: Bambusoideae) and a Revised Infrageneric Classification for Chusquea

A phylogenetic analysis of five coding and noncoding chloroplast loci, totaling 6.6 kb of aligned nucleotide and indel characters, suggests that the large neotropical bamboo genus Chusquea is embedded within the much smaller Andean genus Neurolepis. Monophyly of each taxon was anticipated due to the unique occurrence of dimorphic multiple buds in Chusquea and the lack of aerial branching in Neurolepis. We tested whether the unexpected placement of Chusquea might be the result of a biased analysis. Both Neurolepis and the outgroup taxa have long branches that could influence rooting and inferred ingroup relationships. A number of methods were employed to test for long-branch attraction and sampling effects in our topology. Alternative hypothesis testing using a conservative form of the Shimodaira-Hasegawa test indicated that paraphyly of Neurolepis is a significantly better explanation of the data than monophyly, even when models of character evolution are changed. Given the robustness of the topology, high support measures for clades on the tree, and the results of the Shimodaira-Hasegawa tests, we conclude that chloroplast genomes indicate probable paraphyly of Neurolepis with respect to Chusquea. The species of Neurolepis are therefore transferred to Chusquea, resulting in the following new combinations and names: Chusquea acuminatissima, C. angusta, C. asymmetrica, C. cylindrica, C. diversiglumis, C. elata, C. fimbriligulata, C. laegaardii, C. magnifolia, C. mollis, C. nana, C. nobilis, C. petiolata, C. rigida, C. silverstonei, C. spectabilis, C. steyermarkii, C. stuebelii, C. tovari and C. villosa. The names Neurolepis elata, N. stuebelii, N. weberbaueri, Planotia ingens, and P. tessellata are lectotypified.

A re-evaluation of Parasaurus geinitzi, the first named pareiasaur (Amniota, Parareptilia)

The historically first named pareiasaur Parasaurus geinitzi from the Upper Permian Kupferschiefer deposits of Germany is re-described in detail, including information from recently discovered specimens, as well as material that was for a long time assumed to be lost. The re-study provides new information on both the cranial and the postcranial anatomy of the taxon, the most notable features being the presence of small spikes in the postero-temporal region of the skull and the apparent lack of osteoderms in the axial skeleton, which is unique among pareiasaurs but requires further investigation and new material. A phylogenetic analysis indicates that Parasaurus groups within the more derived pareiasaurs; however, the overall resolution of the analysis is very poor, most likely because many characters traditionally used in studies on pareiasaurian phylogeny were originally defined only for investigating turtle relationships, but not for deciphering the ingroup relationships of pareiasaurs. A careful revision of the phylogeny of the clade is necessary and should also include a detailed assessment of pareiasaurian anatomy, which is still poorly known.

The skull of the giant predatory pliosaur Rhomaleosaurus cramptoni: implications for plesiosaur phylogenetics

The predatory pliosaurs were among the largest creatures ever to inhabit the oceans, some reaching gigantic proportions greater than 15 m in length. Fossils of this subclade of plesiosaurs are known from sediments all over the world, ranging in age from the Hettangian (approximately 198 Myr) to the Turonian (approximately 92 Myr). However, due to a lack of detailed studies and because only incomplete specimens are usually reported, pliosaur evolution remains poorly understood. In this paper, we describe the three dimensionally preserved skull of the giant Jurassic pliosaur Rhomaleosaurus cramptoni. The first phylogenetic analysis dedicated to in-group relationships of pliosaurs allows us to hypothesise a number of well-supported lineages that correlate with marine biogeography and the palaeoecology of these reptiles. Rhomaleosaurids comprised a short-lived and early diverging lineage within pliosaurs, whose open-water top-predator niche was filled by other pliosaur taxa by the mid-late Jurassic.

A taxonomic revision of the orb-weaving spider genus Acusilas Simon, 1895 (Araneae, Araneidae)

AbstractThe spider genus Acusilas Simon, 1895 (Araneidae) is revised. It includes Acusilas africanus Simon, 1895; Acusilas callidus sp. n.; Acusilas coccineus Simon, 1895; Acusilas dahoneus Barrion & Litsinger, 1995; Acusilas lepidus (Thorell, 1898); Acusilas malaccensis Murphy & Murphy, 1983; Acusilas spiralis sp. n.; Acusilas vei sp. n.; Acusilas vilei sp. n., and one unassigned species of Acusilas from Sulawesi for which only females are available. The status of Acusilas lepidus (Thorell, 1898) is uncertain and the species is here considered incertae sedis. The unknown male of Acusilas africanus Simon, 1895 is described and illustrated for the first time and Acusilas gentingensis (Murphy & Murphy) is a synonym of Acusilas coccineus Simon, 1895. A phylogenetic analysis of the matrix of Scharff & Coddington (1997) with the addition of Acusilas coccineus, suggests sistergroup relationship between Acusilas and cyrthophorines (Cyrthophora and Mecynogea). This knowledge is used to select 6 outgroup taxa for a phylogenetic analysis based on 39 morphological characters scored for the 9 known species of Acusilas. The phylogenetic analysis supports the monophyly of Acusilas, but ingroup relationships are only weakly supported.

Phylogeny of the Grasses (Poaceae) Revisited

The most robust previously published phylogeny for the overall structure of the grass family (Poaceae) shows three early diverging lineages and two major derived clades, the BEP clade and the PACCAD clade (Grass Phylogeny Working Group 2001). A few key taxa were incompletely sampled, however, and support for the BEP clade was moderate at best and relationships among the major lineages within the PACCAD clade remained unresolved. In addition, recent studies indicated that the sister group to Poaceae may be Joinvilleaceae and/or Ecdeiocoleaceae, the latter of which were not previously sampled. In this study, missing structural data were determined and analyzed as well as sequence data for ndhF and rbcL, the two most complete plastid sequence data sets. Sampling was increased with a particular focus on key taxa such as Danthoniopsis, Eriachne, Micraira, and Streptogyna and a representative of the outgroup, Ecdeiocoleaceae. A total of 61 ingroup and two outgroup taxa were analyzed using maximum parsimony for total data, and maximum parsimony, Bayesian inference, and neighbor joining for the molecular data. A strongly supported clade of ((Eriachneae, Isachne) Micraira) was recovered as a sister subfamily to Arundinoideae and excluded from Panicoideae. Arundinaria was strongly united with Bambusoideae. The position of Streptogyna was weakly supported among Ehrhartoideae, and is still unresolved. An outgroup effect on ingroup topology was observed, demonstrating that highly divergent outgroups may unpredictably alter ingroup relationships.

Significance of chaetal arrangement for maldanid systematics (Annelida, Maldanidae)

Maldanids are usually divided into several subfamilies: Euclymeninae, Lumbriclymeninae, Maldaninae, Nicomachinae, Rhodininae, Clymenurinae, Notoproctinae, and Boguinae. The taxonomy of maldanids and the delimination of these taxa are mainly based on head morphology, total number of segments, chaetal structure, shape of the pygidium, and position of the anus. The maldanid ingroup relationships, as well as the monophyly of the proposed subfamilies, have so far not been investigated. Pilgrim (1977) described a shift of the notopodial chaetal rows from a transverse direction in anterior chaetigers to a more longitudinal one in posterior chaetigers in Clymene torquata (Leidy, 1855) and Euclymene oerstedii (Claparede, 1863), both belonging to the Euclymeninae. We investigated several maldanid species to assess the usefulness of this character for maldanid systematics and used 3D-reconstruction techniques to reveal the complete geometry of the chaetal sacs. Our investigation shows that a shift is apparent in Euclymene, Axiothella, Johnstonia (all Euclymeninae) and Clymenura (Clymenurinae), but absent in species like Maldane sarsi (Malmgren, 1865), Metasychis disparidentata (Moore, 1904) (both Maldaninae) and Petaloproctus borealis Ardwisson, 1906 (Nicomachinae). The shift is not typical for sedentary polychaetes and is apomorphic within maldanid polychaetes. It thus argues for a close relationship between Euclymeninae and Clymenurinae. The investigation of further maldanid species of different subfamilies may shed additional light on maldanid systematics.

PHYLOGENETICS OF THE GENUS CHIRONIUS FITZINGER, 1826 (SERPENTES, COLUBRIDAE) BASED ON MORPHOLOGY

I present a phylogeny of the Latin American snake genus Chironius based on 36 external morphological and hemipenal characters. Data from the literature for the 20 recognized species of Chironius were combined with data I collected from 20 outgroup taxa. I assessed the monophyly and in-group relationships of Chironius using both parsimony and methods. Two optimal trees were obtained using parsimony criteria; the genus Chironius is supported as monophyletic with 93% bootstrap support from six unambiguous synapomorphies, including an even number of scale rows at midbody. A Bayesian analysis of the data supports the monophyly of Chironius with a posterior probability of 100%. Both analyses inferred similar topologies; recognized species groups are partially supported, but this analysis suggests that revision of the genus is necessary. I propose elevation to full species status of taxa within the carinatus, fuscus, multiventris, and quadricarinatus species groups on the basis of morphological features unique t...

Using Leader-Member Exchange Theory to Explain Students' Motives to Communicate

The purpose of this investigation was to apply Leader-Member Exchange theory to the student-instructor relationship and explore if students' perceptions of their relational quality with their instructors are reflected in their motives to communicate with their instructors. Participants were 139 undergraduate students who completed the Leader-Member Exchange 7 scale and the Student Communication Motives scale. It was found that students who perceive in-group relationships with their instructors report using the relational, functional, participatory, and sycophantic motives at a higher rate than students who perceive out-group relationships with their instructors. Future researchers should explore the ramifications of student-instructor in-group relationships on classroom climate.

The complete mitochondrial genome of the mantid shrimp Harpiosquilla harpax, and a phylogenetic investigation of the Decapoda using mitochondrial sequences

The complete mitochondrial DNA sequence was determined for the mantid shrimp Harpiosquilla harpax. These data demonstrate that the H. harpax mitochondrial genome is a 15,714 bp circular molecule and encodes the typical 37 metazoan mitochondrial genes (13 protein-coding, 22 tRNA, and two rRNA genes). The gene arrangement of H. harpax is consistent with that of the putative arthropod ancestral gene order as depicted by Limulus polyphemus. H. harpax was employed as an outgroup taxon for a phylogenetic investigation of the Decapoda using sequences from complete mitochondrial genomes. Whilst our results are largely in agreement with current taxonomic treatments, the relationships indicated among the reptantian decapods are novel. Our results provide strong statistical support for a sister-group relationship between the Achelata and the Astacida. These findings not only refute previous phylogenetic hypotheses, but also have serious implications for the interpretation of morphological and developmental evolution in the Decapoda. In addition we also investigated the effects of outgroup selection on the resolution of ingroup relationships. We found outgroup choice to significantly influence tree topology thus reinforcing the importance of appropriate outgroup selection in phylogenetic studies.

The anatomy ofAskeptosaurus italicusfrom the Middle Triassic of Monte San Giorgio and the interrelationships of thalattosaurs (Reptilia, Diapsida)

The anatomy of the thalattosauriform reptile Askeptosaurus italicus from the Middle Triassic of Monte San Giorgio is redescribed. Important anatomical features are the plesiomorphic braincase, the unique fronto-lacrimal contact, the absence of a previously described thyroid fenestra, and significant intraspecific variation in the carpus. An analysis of thalattosaur ingroup relationships reveals that Endennasaurus and the monophyletic Askeptosaurus and Anshunsaurus are the sister group to all other thalattosaurs, whereas the Monte San Giorgio taxa Clarazia and Hescheleria form the sister clade of Thalattosaurus, and the Chinese Xinpusaurus and the Californian Nectosaurus form a monophyletic group. The analysis supports the biogeographic interpretation of trans-Pacific relationships and a re-invasion of Tethyan areas.

In Search of the Origin of Twin Penises: Molecular Phylogeny of Earwigs (Dermaptera: Forficulina) Based on Mitochondrial and Nuclear Ribosomal RNA Genes

Abstract Forficulina, the largest suborder of Dermaptera (earwigs), has eight families. In five families (Pygidicranidae, Diplatyidae, Anisolabididae, Apachyidae, and Labiduridae), the males have two penises, whereas the males of the other three families (Spongiphoridae, Chelisochidae, and Forficulidae) have a single penis. Several cladograms have been proposed for Forficulina. However, those systems are constructed mainly from observations of male genital morphology and outstandingly inconsistent. This study reconstructed an earwig phylogeny with representatives of seven families (excluding Apachyidae) by using partial sequences of the mitochondrial 16S and nuclear 28S rRNA genes, sequences that are independent of genital evolution. To avoid difficulties caused by outgroup-rooting for a deep phylogeny, ingroup relationships were first established as unrooted trees based on the 16S, 28S, and combined data sets. The resulting affinities of the earwig families supported several superfamilies, such as Forfic...

On the cranial anatomy of the polycotylid plesiosaurs, including new material of Polycotylus latipinnis, Cope, from Alabama

ABSTRACT The cranial anatomy of plesiosaurs in the family Polycotylidae (Reptilia: Sauropterygia) has received renewed attention recently because various skull characters are thought to indicate plesiosauroid, rather than pliosauroid, affinities for this family. New data on the cranial anatomy of polycotylid plesiosaurs is presented, and is shown to compare closely to the structure of cryptocleidoid plesiosaurs. The morphology of known polycotylid taxa is reported and discussed, and a preliminary phylogenetic analysis is used to establish ingroup relationships of the Cryptocleidoidea. This study also presents new material referable to Polycotylus latipinnis from the Mooreville Chalk Formation of Alabama. This skeleton is largely complete, and although the skull is fragmentary it does preserve several regions not previously represented. The preserved portions demonstrate that the palate of Polycotylus is similar to that of Trina-cromerum in the possession of robust epipterygoids, and similar to that of all other polycotylids in the possession of a large anterior interpterygoid vacuity. Polycotylus therefore displays the derived palate morphology common to all polycotylids, but also displays some plesiomorphic features. This impression is confirmed by the results of the cladistic analysis.

Phylogenetic Relationships among the Mosses Based on Heterogeneous Bayesian Analysis of Multiple Genes from Multiple Genomic Compartments

Nucleotide sequences from eight nuclear, chloroplast, and mitochondrial genes were obtained from 30 mosses (plus four outgroup liverworts) in order to resolve phylogenetic relationships among the major clades of division Bryophyta. Phylogenetic analyses were conducted using maximum parsimony, maximum likelihood (ML), and Bayesian inference. Inferences were compared from Bayesian analyses using homogeneous and several heterogeneous models. Estimates of clade confidence were based on bootstrap analyses, posterior probabilities (in Bayesian analyses) and novel combined approaches. Most ingroup relationships were congruent among analyses, but support for individual clades depended on the analytical approach. Increasingly parameterized models of nucleotide substitution in the likelihood analyses provided significantly higher goodness-of-fit to the data. The results suggest that 1) the Bryophyta, including Sphagnum and Takakia, are monophyletic, 2) Andreaea and Andreaeobryum form a monophyletic group, 3) Oedipodium griffithianum is sister to all other operculate taxa, 4) mosses with nematodontous peristomes are paraphyletic and basal to arthrodontous mosses, 5) Diphyscium is sister to all other arthrodontous mosses, 6) Encalypta is sister to the Funariaceae, and 6) mosses with diplolepideous-alternate peristomes form a monophyletic group. Implications of the phylogenetic hypothesis for morphological evolution in mosses include 1) a pseudopodium has arisen independently in Sphagnum and Andreaea, 2) the mucilage hairs of Andreaeobryum and Takakia are non-homologous, 3) the stomata found in Sphagnum are not homologous to those of other mosses, and 4) that stomata were absent in the ancestor of all mosses.

Adulthood and phylogenetic analysis in gastropods: character recognition and coding in shells of Lavigeria (Cerithioidea, Thiaridae) from Lake Tanganyika

In the study of gastropod shell morphology, determination of comparable ontogenetic stages is crucial, because all the states that various shell features go through during ontogeny are preserved on the shell. The protoconch/teleoconch transition and marks of episodic growth are among the few ways of defining discrete, comparable, growth stages. In gastropods with determinate growth the attainment of adulthood may provide additional shell markers permitting comparison among individuals and taxa. Adulthood is reflected in shell morphology in ways as diverse as shell deposition covering all the previous whorls and radically changing the shape of the shell through to slight changes in the trajectory of the suture. While the very prominent adulthood-related changes of shell morphology have been used as systematic characters, the more moderate changes have not been studied in detail and their potential systematic value has been ignored. In this paper we give a detailed account of adult modifications of the shell appearing with cessation of growth. Our study group comprises eight closely related species of Lavigeria from Lake Tanganyika. We show that the ways adulthood is manifested are quite diverse. We describe eight characters of the aperture, the suture and the sculptural ornamentation. Character occurrence varies greatly among species. We show that characters appear in suites and that in many cases their appearance is connected to size. We use size as a proxy for adulthood and test whether character occurrence alone or its connection to size can help resolve species relationships. In both cases our characters confirm the monophyly of our ingroup and yield cladograms with various degrees of resolution of ingroup relationships. The coding method that yields the greater character congruence is the one that takes into consideration the connection between appearance of a character and size. This study demonstrates that ontogenetically correlated character transformations may nevertheless be phylogenetically independent. © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 140, 223−240.