The role of chromosome changes in the diversification of Passiflora L. (Passifloraceae)

Passiflora L. has more than 575 species distributed especially in the Neotropics. The chromosome number variation in the genus is highly congruent with its main subgenera, but its basic chromosome number (x) and the underlying events responsible for this variation have remained controversial. Here, we provide a robust and well-resolved time-calibrated phylogeny that includes 102 taxa, and by means of phylogenetic comparative methods (PCM) we tested the relative importance of polyploidy and dysploidy events to Passiflora karyotype evolution and diversification. Passiflora arose 42.9 Mya, with subgenus diversification at the end of the Palaeogene (Eocene-Oligocene). The basic chromosome number of the genus is proposed as x?=?6, and a strong recent diversification found in the Passiflora subgenus (Miocene) correlated to genome size increase and a chromosome change from n?=?6 to n?=?9 by ascending dysploidy. Polyploidy, conversely, appeared restricted to few lineages, such as Astrophea and Deidamioides subgenera, and did not lead to diversification increases. Our findings suggest that ascending dysploidy provided a great advantage for generating long-term persistent lineages and promoting species diversification. Thus, chromosome numbers/genome size changes may have contributed to morphological/ecological traits that explain the pattern of diversification observed in the genus Passiflora.     

Testing for evolutionary trade-offs in a phylogenetic context: ecological diversification and evolution of locomotor performance in emydid turtles

The evolution of ecological trade-offs is an important component of ecological specialization and adaptive radiation. However, the pattern that would show that evolutionary trade-offs have occurred between traits among species has not been clearly defined. In this paper, we propose a phylogeny-based definition of an evolutionary trade-off, and apply it to an analysis of the evolution of trade-offs in locomotor performance in emydid turtles. We quantified aquatic and terrestrial speed and endurance for up to 16 species, including aquatic, semi-terrestrial and terrestrial emydids. Emydid phylogeny was reconstructed from morphological characters and nuclear and mitochondrial DNA sequences. Surprisingly, we find that there have been no trade-offs in aquatic and terrestrial speed among species. Instead, specialization to aquatic and terrestrial habitats seems to have involved trade-offs in speed and endurance. Given that trade-offs between speed and endurance may be widespread, they may underlie specialization to different habitats in many other groups.     

Molecular phylogeny and ecological diversification in a clade of New World songbirds (genus Vireo)

We constructed a molecular phylogeny for a clade of eye-ringed vireos ( Vireo flavifrons and the V . solitarius complex) to examine existing hypotheses of speciation and ecological diversification. Complete sequences of the mtDNA cytochrome b gene were obtained from 47 individuals of this group plus four vireonid outgroups. Mean levels of sequence divergence in the clade varied from 0.29% to 5.7%. Differences were greatest between V . flavifrons and four taxa of ' V . solitarius '. The latter separated into three taxonomic, geographical and ecological groups: V . plumbeus plumbeus, V . cassinii cassinii , and V . solitarius solitarius plus V . solitarius alticola . These differed by an average of 2.6–3.2%. Populations within each group revealed low levels of sequence variation ( x- = 0.20%) and little geographical structuring. The mtDNA data generally corroborate results from allozymes. V. plumbeus shows a loss of yellow-green carotenoid pigmentation from the ancestral condition. The occupancy of relatively dry habitats by this species and V . cassinii represents a derived ecological shift from more-humid environments occupied by other species of vireonids. Ecological divergence in this clade occurred in allopatry and is associated with generic-level stability in morphometrics and foraging styles. Migratory behaviour and seasonal habitat shifts apparently evolved multiple times in vireos breeding in temperate environments. Present geographical and ecological distributions, and low levels of intrataxon genetic divergence, are hypothesized to be the result of postglacial regionalization of climate–plant associations and rapid northward expansion of breeding ranges.     

Losses of female song with changes from tropical to temperate breeding in the New World blackbirds

Birds in which both sexes produce complex songs are thought to be more common in the tropics than in temperate areas, where typically only males sing. Yet the role of phylogeny in this apparent relationship between female song and latitude has never been examined. Here, we reconstruct evolutionary changes in female song and breeding latitude in the New World blackbirds (Icteridae), a family with both temperate and tropical representatives. We provide strong evidence that members of this group have moved repeatedly from tropical to temperate breeding ranges and, furthermore, that these range shifts were associated with losses of female song more often than expected by chance. This historical perspective suggests that male-biased song production in many temperate species is the result not of sexual selection for complex song in males but of selection against such songs in females. Our results provide new insights into the differences we see today between tropical and temperate songbirds, and suggest that the role of sexual selection in the evolution of bird song might not be as simple as we think.     

Molecular evidence for naturalness of genera in the tribe Antirrhineae (Scrophulariaceae) and three independent evolutionary lineages from the New World and the Old

The tribe Antirrhineae consists of 29 genera distributed in the New World and the Old. Phylogenetic analyses of ITS and ndhF sequences served to recognize six main lineages: Anarrhinum group (Anarrhinum, Kickxia); Linaria group (Linaria); Maurandya group (Cymbalaria, Asarina, Maurandella, Rhodochiton, Lophospermum); Schweinfurthia group (Pseudorontium, Schweinfurthia); Antirrhinum group (Antirrhinum, Pseudomisopates, Misopates, Acanthorrhinum, Howeliella, Neogarrhinum, Sairocarpus, Mohavea, Galvezia); Chaenorrhinum group (Chaenorrhinum, Albraunia, Holzneria). Parsimony (cladistics), distance-based (Neighbor-Joining), and Bayesian inference reveal that: (1) the tribe is a natural group; (2) genera such as Linaria, Schweinfurthia, Kickxia, and Antirrhinum also form natural groups; (3) three Antirrhineae lineages containing genera from the New and Old World are the result of three intercontinental disjunctions displaying similar levels of ITS-sequence divergence and differentiation times (Oligocene-Miocene); (4) evolution of flower shapes is not congruent with primitiveness of personate flowers; (5) both polyploidy and dysploidy appear to be responsible for most variation in chromosome number in the six main lineages. Nuclear and chloroplast evidence also supports the split of American and Mediterranean species of Antirrhinum into different genera, a result that should be contemplated in the interest of a more natural (monophyletic) taxonomy. Nucleotide additivity causes poor resolution in the ITS analysis of 22 species of Mediterranean Antirrhinum and lead us to interpret extensive hybridization in the Iberian Peninsula.This research was supported by the Spanish Dirección General de Investigación Científica y Técnica (DGICYT) through the project Flora Iberica (PS91-0070-C03-01). We thank Jorge Martínez and Emilio Cano for his assistance in the lab, Rafael Rubio and Omar Fiz for helping on the Bayesian inferences, the curators/collection managers of BCF, E, JACA, MA, SALA, SALAF, SEV, TARI, VAL for loans and access to specimens, and two anonymous reviewers for improving the quality of this publication.     

Molecular phylogeny, diversification and character evolution in Lupinus (Fabaceae) with special attention to Mediterranean and African lupines

Phylogenetic relationships within the complex genus Lupinus are estimated from internal transcribed spacer (ITS) sequences of the nuclear ribosomal DNA repeat. The molecular data supports Lupinus as a distinct monophyletic group within the tribe Genisteae. Different geographical lineages are revealed within Lupinus, which are each restricted to either the Old or the New World. In the New World, the ITS data support an eastern-western geographic disjunction of the lupines and the recognition of some well-supported clades. In the Old World, almost all the previously recognized taxa are taxonomically well differentiated. The homogeneous African rough-seeded lupines, Scabrispermae, are strongly supported as a monophyletic group, which is distinct from the diverse and heterogeneous circum-Mediterranean smooth-seeded ones. The latter appear to have evolved as two lineages, in which are revealed some intersectional relationships. Also ITS data allow the assessment of the phylogenetic position of the newly discovered species, L. anatolicus (in the Old World) and L. jaimehintoniana (the Mexican tree lupin). The ITS phylogeny suggests a rapid initial radiation of the lupines subsequent to their divergence from a common ancestor. Moreover, the results indicate that the annual and perennial habits have evolved many times in Lupinus and suggest a role for generation time in affecting the evolutionary history of lupines. Data on adaptive processes and character evolution are re-examined and discussed in the light of the ITS phylogeny.These studies were supported by the research unit UMR-CNRS 6553 Ecobio – University of Rennes (France), and by the NSERC of Canada (grant to R.J. Bayer, at the University of Alberta). They are greatfully acknowledged. All the persons that have contributed in different ways to this work on Lupinus, summarized in this presentation, are greatly thanked. Particularly, we would like to mention Malika Aïnouche, Roland Greinwald, André Huon, W.K. Swiecicki, Billie L. Turner and Ludger Witte for their contributions.     

Patterns of cranial shape diversification during the phylogenetic branching process of New World monkeys (Primates: Platyrrhini)

One of the central topics in evolutionary biology is understanding the processes responsible for phenotypic diversification related to ecological factors. New World monkeys are an excellent reference system to investigate processes of diversification at macroevolutionary scales. Here, we investigate the cranial shape diversification related to body size and ecology during the phylogenetic branching process of platyrrhines. To investigate this diversification, we used geometric morphometric techniques, a molecular phylogenetic tree, ecological data and phylogenetic comparative methods. Our statistical analyses demonstrated that the phylogenetic branching process is the most important dimension to understand cranial shape variation among extant platyrrhines and suggested that the main shape divergence among the four principal platyrrhine clades probably occurred during the initial branching process. The phylogenetic conservatism, which is the retention of ancestral traits over time within the four principal platyrrhine clades, could be the most important characteristic of platyrrhine cranial shape diversification. Different factors might have driven early shape divergence and posterior relative conservatism, including genetic drift, stabilizing selection, genetic constraints owing to pleiotropy, developmental or functional constraint, lack of genetic variation, among others. Understanding the processes driving the diversification among platyrrhines will probably require further palaeontological, phylogenetic and comparative studies.     

Exceptional among-lineage variation in diversification rates during the radiation of Australia's most diverse vertebrate clade

The disparity in species richness among groups of organisms is one of the most pervasive features of life on earth. A number of studies have addressed this pattern across higher taxa (e.g. 'beetles'), but we know much less about the generality and causal basis of the variation in diversity within evolutionary radiations at lower taxonomic scales. Here, we address the causes of variation in species richness among major lineages of Australia's most diverse vertebrate radiation, a clade of at least 232 species of scincid lizards. We use new mitochondrial and nuclear intron DNA sequences to test the extent of diversification rate variation in this group. We present an improved likelihood-based method for estimating per-lineage diversification rates from combined phylogenetic and taxonomic (species richness) data, and use the method in a hypothesis-testing framework to localize diversification rate shifts on phylogenetic trees. We soundly reject homogeneity of diversification rates among members of this radiation, and find evidence for a dramatic rate increase in the common ancestor of the genera Ctenotus and Lerista. Our results suggest that the evolution of traits associated with climate tolerance may have had a role in shaping patterns of diversity in this group.     

Evolution of viviparity in horned lizards (Phrynosoma): testing the cold-climate hypothesis

The cold-climate hypothesis for evolution of viviparity in squamates predicts a correlation between reproductive mode, altitude and latitude. I tested this prediction in horned lizards within a phylogenetic context. I first determined whether all viviparous species were monophyletic using Monte Carlo simulations. Secondly, I tested for presence of phylogenetic signal using randomization tests. Thirdly, I analysed relationships between reproductive mode and minimum, midpoint, and maximum altitudes and latitudes by computing conventional correlations and phylogenetically independent contrasts. Viviparous species do not form a monophyletic group suggesting viviparity evolved twice in the genus. Viviparity and altitude showed strong phylogenetic signal based on randomization tests and were significantly correlated, while latitude was not correlated with reproductive mode. This study partially supports the cold-climate model, but also suggests that altitude either may be a better predictor of cold temperatures or may be a surrogate for other selective factors important in the evolution of viviparity.     

Fossils and phylogeny uncover the evolutionary history of a unique antipredator behaviour

Recently, two squirrel species (Spermophilus spp.) were discovered to anoint their bodies with rattlesnake scent as a means of concealing their odour from these chemosensory predators. In this study, we tested multiple species with predator scents (rattlesnake and weasel) to determine the prevalence of scent application across the squirrel phylogeny. We reconstructed the evolutionary history of the behaviour using a phylogenetic analysis and fossil records of historic predator co‐occurrence. Squirrels with historical and current rattlesnake co‐occurrence all applied rattlesnake scent, whereas no relationship existed between weasel scent application and either weasel or rattlesnake co‐occurrence. This was surprising because experimental tests confirmed rattlesnake and weasel scent were both effective at masking prey odour from hunting rattlesnakes (the primary predator of squirrels). Ancestral reconstructions and fossil data suggest predator scent application in squirrels is ancient in origin, arising before co‐occurrences with rattlesnakes or weasels in response to some other, now extinct, chemosensory predator.     

Historical biogeography and evolutionary diversification of Lilium(Liliaceae): New insights from plastome phylogenomicsOA

Here, we infer the historical biogeography and evolutionary diversification of the genus Lilium. For this purpose, we used the complete plastomes of 64 currently accepted species in the genus Lilium(14plastomes were newly sequenced) to recover the phylogenetic backbone of the genus and a timecalibrated phylogenetic framework to estimate biogeographical history scenarios and evolutionary diversification rates of Lilium. Our results suggest that ancient climatic changes and geological tectonic act...     

Monophyletic subgroups of the tribe Millettieae (Leguminosae) as revealed by phytochrome nucleotide sequence data

Phylogenetic analysis of phytochrome (PHY) genes reveals the identity and relationships of four PHY loci among papilionoid Leguminosae. A phylogenetic analysis of loci combined according to species suggests that most of the tribe Millettieae belongs to one of two monophyletic clades: the Derris–Lonchocarpus or the Tephrosia clade. Together these two form a monophyletic group that is sister to a lineage represented by Millettia grandis of Millettia sect. Compresso-gemmatae. Collectively, this large monophyletic group is referred to as the Millettieae-core group, which based on our sampling, includes species of Millettieae that do not accumulate the nonprotein amino acid canavanine and that mostly have pseudoracemose or pseudopaniculate inflorescences. This new phylogenetic framework assists in targeting additional taxa for future sampling. For example, the “American Derris” (Deguelia), which accumulate canavanine, might not be members of the Millettieae core group. Afgekia is also predicted not to be a member because it accumulates canavanine and has an inflorescence of terminal racemes. PHY gene analysis specifically reveals that certain genera traditionally classified in Millettieae are actually distantly related to the Millettieae core group, such as Austrosteensia, Callerya, Craibia, Cyclolobium, Fordia, Platycyamus, Poecilanthe, and Wisteria.     

The geographical pattern of speciation and floral diversification in the neotropics: the tribe sinningieae (gesneriaceae) as a case study

The geographical pattern of speciation and the relationship between floral variation and species ranges were investigated in the tribe Sinningieae (Gesneriaceae), which is found mainly in the Atlantic forests of Brazil. Geographical distribution data recorded on a grid system of 0.5 x 0.5 degree intervals and a near-complete species-level phylogenetic tree of Sinningieae inferred from a simultaneous analysis of seven DNA regions were used to address the role of geographical isolation in speciation. Geographical range overlaps between sister lineages were measured across all nodes in the phylogenetic tree and analyzed in relation to relative ages estimated from branch lengths. Although there are several cases of species sympatry in Sinningieae, patterns of sympatry between sister taxa support the predominance of allopatric speciation. The pattern of sympatry between sister taxa is consistent with range shifts following allopatric speciation, except in one clade, in which the overlapping distribution of recent sister species indicates speciation within a restricted geographical area and involving changes in pollinators and habitats. The relationship between floral divergence and regional sympatry was also examined by analyzing floral contrasts, phenological overlap, and the degree of sympatry between sister clades. Morphological contrast between flowers is not increased in sympatry and phenological divergence is more apparent between allopatric clades than between sympatric clades. Therefore, our results failed to indicate a tendency for sympatric taxa to minimize morphological and phenological overlap (geographic exclusion and/or character displacement hypotheses). Instead, they point toward adaptation in phenology to local conditions and buildup of sympatries at random with respect to flower morphology. Additional studies at a lower geographical scale are needed to identify truely coexisting species and the components of their reproductive isolation.     

Origin and diversification of Thymelaea (Thymelaeaceae): inferences from a phylogenetic study based on ITS (rDNA) sequences

Thymelaea is a Mediterranean genus belonging to a primarily tropical and subtropical family. This genus is here presented as a particular case on which the hypothesis of an in situ evolution of the Mediterranean flora from a Tertiary subtropical stock can be phylogenetically tested. To better understand the evolutionary history of Thymelaea, molecular phylogenies based on ITS (rDNA) sequences are estimated. Phylogenetic analyses strongly support the monophyly of Thymelaea and the sister position of Daphne. Unexpected relationships between sequences of T. granatensis, T. pubescens and T. sanamunda are found and explained as a case of cryptic nuclear introgression. Phylogenetic analyses also reveal the polyphyletic character of Diarthron, sensu Tan, suggesting that Dendrostellera and Stelleropsis should be reinstated as independent genera. Based on the resulting phylogenetic hypothesis and using statistical tests it is suggested that Thymelaea experienced an initial rapid diversification. Time estimates for the origin and radiation of Thymelaea based on independent sources of evidence—paleontological, paleoclimatic and paleogeographic data, on one hand, and ITS substitution rates reported for other angiosperm groups, on the other—give strikingly similar results. The early radiation of Thymelaea is inferred to have occurred in the Upper Miocene, after this lineage diverged from that of Daphne in the Lower or Middle Miocene. Both evolutionary events are related to the overall climatic deterioration (declining temperature and increasing dryness) registered throughout the Miocene in Eurasia. The rapid and extensive diversification of Thymelaea is placed in the Iberian Peninsula or here and in the neighbouring North African territory.     

Phylogeny of the tribe Antirrhineae (Scrophulariaceae) based on morphological andndhF sequence data

Phylogenetic relationships within the tribe Antirrhineae (Scrophulariaceae) are analysed and discussed on the basis of parsimony analyses of morphological andndhF gene sequence data. The results indicate that the tribe Antirrhineae consists of four major groups of genera, theAnarrhinum clade, theGambelia clade, theMaurandya clade, and theAntirrhinum clade. TheAnarrhinum clade, consisting of the Old World bee-pollinated generaAnarrhinum andKickxia, is sister to the rest of the tribe. TheGambelia clade consists of the New World generaGambelia andGalvezia, which are very closely related and pollinated by hummingbirds. TheMaurandya clade consists of one subclade includingMaurandya and a number of related bee- or hummingbird-pollinated New World genera and another subclade with the Old World bee-pollinated generaAsarina andCymbalaria. TheAntirrhinum clade consists mainly of bee-pollinated Old World genera, such asAntirrhinum, Linaria, Chaenorhinum, and their segregates, but also includes the New World generaMohavea andHowelliella, of which the latter is known to be partly pollinated by hummingbirds. It is concluded that hummingbirdpollination has evolved independently within Antirrhineae at least three times from bee-pollinated ancestors.     

Phylogenetic systematics of the tribe Millettieae (Leguminosae) based on chloroplast trnK/matK sequences and its implications for evolutionary patterns in Papilionoideae

Phylogenetic relationships in the tribe Millettieae and allies in the subfamily Papilionoideae (Leguminosae) were reconstructed from chloroplast trnK/matK sequences. Sixty-two accessions representing 57 traditionally recognized genera of Papilionoideae were sampled, including 27 samples from Millettieae. Phylogenies were constructed using maximum parsimony and are well resolved and supported by high bootstrap values. A well-supported "core Millettieae" clade is recognized, comprising the four large genera Millettia, Lonchocarpus, Derris, and Tephrosia. Several other small genera of Millettieae are not in the core Millettieae clade. Platycyamus is grouped with Phaseoleae (in part). Ostryocarpus, Austrosteenisia, and Dalbergiella are neither in the core Millettieae or Phaseoleae clade. These taxa, along with core Millettieae and Phaseoleae, form a monophyletic sister group to Indigofereae. Cyclolobium and Poecilanthe are close to Brongniartieae. Callerya and Wisteria belong to a large clade that includes all the legumes that lack the inverted repeat in their chloroplast genome, which confirms previous rbcL and phytochrome gene family phylogenies. The evolutionary history of four characters was examined in Millettieae and allies: the presence of canavanine, inflorescence types, the dehiscence of pods, and the presence of winged pods. trnK/matK sequence analysis suggests that the presence of a pseudoraceme or pseudopanicle and the accumulation of nonprotein amino acids are phylogenetically informative for Millettieae and allies with only a few exceptions.     

Evolutionary history of two allopatric Terricola species (Arvicolinae, Rodentia) from molecular, morphological, and palaeontological data

To investigate the phylogenetic and phylogeographical relationships of arvicolines, we use several Western European ground voles. More particularly, our study is focused on Microtus ( Terricola ) savii and M. ( T. ) pyrenaicus . These two allopatric species are usually considered as having originated from the same ancestor, possibly M . ( T. ) mariaclaudiae . We propose molecular and morphological approaches: nucleotidic data from the mitochondrial cytochrome b and 12S rRNA genes and global morphological analyses from the first lower molar. Four other Terricola species ( multiplex , lusitanicus , duodecimcostatus , subterraneus ) were added to the data set for both analyses, and two other vole species ( Clethrionomys glareolus and Chionomys nivalis ) as outgroup to the molecular analysis, and five fossil populations to the morphological one. Palaeontological data are also widely taken into account. Both molecular and morphological analyses indicate that intra- Terricola relationships reflect the present-day geographical distribution of our data set species. Our results show that M. ( T. ) savii and M. ( T. ) pyrenaicus are from separate speciation events leading to two different biogeographical groups, respectively the Alpine–Italian group and the French–Iberian group, the latter being much more homogeneous. These speciation events could be related to Quaternary climatic changes, which induced southward migration, leading first to M. ( T. ) savii and second to M. ( T. ) pyrenaicus . The classical hypothesis of a geographical speciation for these two taxa from M. ( T. ) mariaclaudiae is invalid. However, the morphological data suggest a potential phylogenetic relationship between M. ( T. ) mariaclaudiae (ancestor) and M. ( T. ) pyrenaicus (descendant).  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 309–323.     

Integrative taxonomy approach for analysing evolutionary history of the tribe Euseiini Chant & McMurtry (Acari: Phytoseiidae)

Phylogenetic relationships within the mite Family Phytoseiidae are little known. The presently accepted classification is based on the opinion of specialists, but not on cladistics analysis. The present paper focuses on the tribe Euseiini, containing 271 species, three subtribes and 10 genera. It aims to determine phylogenetic relationships between these taxa and test their monophyly. Molecular analysis combining six markers has been carried out for taxa we succeeded in collecting. Morphological, biogeographic and ecological data have been analysed to determine how these factors can explain the evolutionary relationships emphasized on the phylogenetic tree. Those analyses have been carried out for the taxa available for the molecular study, but also for all species of the tribe. The tribe Euseiini and the two subtribes considered are monophyletic (at least considering the available taxa), supporting the present hypothesis on Phytoseiidae classification. However, the genus Iphiseius seems to not be valid and its unique species is included in the genus Euseius. Clades that were observed within the genus Euseius do not match with recent work on species groups within this genus. It seems that some morphological features such as an insemination apparatus shape and seta length on the dorsal shield constitute some elements explaining the clusters within the genus Euseius. Biogeographic and ecological data analysis led us to hypothesize a west Gondwanian origin of the tribe Euseiini (Africa and Neotropical areas) on Rosids plants (especially of the Orders Malpiphiales and Fabales: subclass Fabidae). Further analyses are still required to (i) take into account more taxa (especially rare ones and species from the Ethiopian part), (ii) to consider more accurate morphological features through more powerful microscopic apparatus, and (iii) to associate a phylogenetic and evolutionary scenario to life traits (pollen feeders).     

Evolution of mimicry in the New World Dysdercus (Hemiptera: Pyrrhocoridae)

Evolution of mimicry rings is reconstructed for the New World species of cotton-stainer bugs (Dysdercus, Hemiptera: Pyrrhocoridae). Using a cladistic analysis of 43 species (including D. silaceus Doesburg, 1968 stat. nov . and D. modestus Doesburg, 1968 stat. nov .), evolution of colour patterns and biogeographical history of the clade are hypothesized. Three probable mimicry rings are obtained from the analysis, together with data about their possible model–mimic or comimic–comimic structures: (a) ‘Yellow’ species in north-western South America and in the Andes (several possible models, D. rusticus, D. mimus part., D. imitator part., and D. collaris part. as mimics); (b) ‘Median Yellow–Black’ species with a median-forewing black pattern in north-western South America and Central America (several possible models, D. basialbus as a mimic); (c) ‘Median Red–Black’ species with a median-forewing black pattern in the Caribbean islands (D. andreae, D. discolor, D. neglectus–D. sanguinarius clade, and D. jamaicensis–D. fervidus–D. ocreatus clade as comimics). A few additional, less supported rings are also identified. Two highly polymorphic species (D. obscuratus, D. mimus) seem to enter different mimicry rings in different regions.