Craniodental characters and the relationships of Procyonidae (Mammalia: Carnivora)

Recent phylogenies of Procyonidae based on molecular data differ significantly from previous morphology‐based phylogenies in all generic sister taxon relationships. I have compiled the most comprehensive dataset of craniodental morphology that incorporates previous morphological characters, and with the aid of high‐resolution X‐ray computed tomography, new characters. This expanded craniodental analysis is based on 78 characters and yields new phylogenetic results regarding the ingroup relationships of Procyonidae. These results include Bassariscus astutus as the least derived member of Procyonidae and Ailurus fulgens nested well within the clade. Additionally, there are some similarities to previous morphological analyses of Procyonidae. Although the characters used to unite and diagnose Procyonidae vary depending on the phylogenetic analysis and have ambiguous taxonomic distribution amongst both Procyonidae and Musteloidea, there is significant morphological support for clades within Procyonidae. In addition to the strength of the morphological support within the clade, the disparate topographical regions of the skull from which the characters are derived may indicate that these synapomorphies are indeed the result of homology rather than adaptive convergence, as suggested by analyses based on molecular data. © 2012 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 669–713.     

The Braincase of Eocaecilia micropodia (Lissamphibia,Gymnophiona) and the Origin of Caecilians

The scant fossil record of caecilians has obscured the origin and evolution of this lissamphibian group. Eocaecilia micropodia from the Lower Jurassic of North America remains the only stem-group caecilian with an almost complete skull preserved. However, this taxon has been controversial, engendering re-evaluation of traits considered to be plesiomorphic for extant caecilians. Both the validity of the placement of E. micropodia as a stem caecilian and estimates of the plesiomorphic condition of extant caecilians have been questioned. In order to address these issues, the braincase of E. micropodia was examined via micro-computed tomography. The braincase is considered to be a more reliable phylogenetic indicator than peripheral regions of the skull. These data reveal significant new information, including the possession of an ossified nasal septum, ossified anterior wall of the sphenethmoid, long anterolateral processes on the sphenethmoid, and paired olfactory nerve foramina, which are known only to occur in extant caecilians; the latter are possibly related to the evolution of the tentacle, a caecilian autapomorphy. A phylogenetic analysis that included 64 non-amniote taxa and 308 characters represents the first extensive test of the phylogenetic affinities of E. micropodia. The results place E. micropodia securely on the stem of extant caecilians, representing a clade within Temnospondyli that is the sister taxon to batrachians plus Gerobatrachus. Ancestral character state reconstruction confirms the braincase of E. micropodia to be largely representative of the plesiomorphic condition of extant caecilians. Additionally, the results refine the context within which the evolution of the caecilian form can be evaluated. The robust construction and pattern of the dermal skull of E. micropodia is interpreted as symplesiomorphic with advanced dissorophoid temnospondyls, rather than being autapomorphic in its robust construction. Together these data increase confidence in incorporating E. micropodia into discussions of caecilian evolution.     

Molecules,morphology, missing data and the phylogenetic position of a recently extinct madtom catfish (Actinopterygii: Ictaluridae)

Taxa missing large amounts of data pose challenges that may hinder the recovery of a well‐resolved, accurate phylogeny and leave questions surrounding their phylogenetic position. Systematists commonly have to contend with one or two species in a group for which there is little or no material available suitable for recovering molecular data. It is unclear whether these taxa can be better placed using analyses based on morphological data only, or should be included in broader analyses based on both morphological and molecular data. The extinct madtom catfish Noturus trautmani is known from few specimens for which molecular data are unavailable. We included this taxon in parsimony and Bayesian analyses of relationships of madtom catfishes based on a combination of morphological and molecular data. Results indicate that using a combination of morphological and molecular data does a better job at providing a phylogenetic placement for N. trautmani than morphology alone, even though it is missing all of its molecular characters. We provide a novel hypothesis of relationships among Noturus species and recommendations for classification within the group. © 2009 The Linnean Society of London, Zoological Journal of the Linnean Society, 2009, 155 , 60–75.     

An Expanded Combined Evidence Approach to the Gavialis Problem Using Geometric Morphometric Data from Crocodylian Braincases and Eustachian Systems

The phylogenetic position of the Indian gharial (Gavialis gangeticus) is disputed - morphological characters place Gavialis as the sister to all other extant crocodylians, whereas molecular and combined analyses find Gavialis and the false gharial (Tomistoma schlegelii) to be sister taxa. Geometric morphometric techniques have only begun to be applied to this issue, but most of these studies have focused on the exterior of the skull. The braincase has provided useful phylogenetic information for basal crurotarsans, but has not been explored for the crown group. The Eustachian system is thought to vary phylogenetically in Crocodylia, but has not been analytically tested. To determine if gross morphology of the crocodylian braincase proves informative to the relationships of Gavialis and Tomistoma, we used two- and three-dimensional geometric morphometric approaches. Internal braincase images were obtained using high-resolution computerized tomography scans. A principal components analysis identified that the first component axis was primarily associated with size and did not show groupings that divide the specimens by phylogenetic affinity. Sliding semi-landmarks and a relative warp analysis indicate that a unique Eustachian morphology separates Gavialis from other extant members of Crocodylia. Ontogenetic expansion of the braincase results in a more dorsoventrally elongate median Eustachian canal. Changes in the shape of the Eustachian system do provide phylogenetic distinctions between major crocodylian clades. Each morphometric dataset, consisting of continuous morphological characters, was added independently to a combined cladistic analysis of discrete morphological and molecular characters. The braincase data alone produced a clade that included crocodylids and Gavialis, whereas the Eustachian data resulted in Gavialis being considered a basally divergent lineage. When each morphometric dataset was used in a combined analysis with discrete morphological and molecular characters, it generated a tree that matched the topology of the molecular phylogeny of Crocodylia.     

Testing the phylogenetic utility of morphological character systems,with a revision of Creophilus Leach (Coleoptera: Staphylinidae)

Many studies have examined the phylogenetic utility of different kinds of molecular data, and have often compared these with morphology. However, relatively few studies have phylogenetically evaluated different morphological character systems. Using the Creophilus complex, the phylogenetic utility of external structural characters, male genitalia, female genitalia, and chaetotaxy of adults is examined for the first time in the megadiverse beetle family Staphylinidae. A data set of 121 phylogenetically informative characters of 24 terminal taxa was analysed separately and simultaneously using parsimony and the phylogenetic utility of each partition was compared with widely used statistics and support measures. External structures had the least homoplasy, resolved the most nodes separately, and largely determined the topology of the simultaneous analysis. Male and female genitalia contributed phylogenetic signal mostly congruent with external characters. Despite extensive homoplasy, chaetotaxy contributed the majority of hidden support, and was critical for resolving several terminal nodes in the simultaneous analysis. All character systems were informative throughout the tree, and when combined provided the best‐supported hypothesis. However, there is a need to distinguish hidden support from dispersion of homoplasy when combining data sets of varying quality. The Creophilus complex, Creophilus Leach, Liusus Sharp, and the Creophilus erythrocephalus ( Fabricius ) and C. maxillosus (Linnaeus) species‐groups are each strongly supported monophyletic groups. Hadrotes Mäklin is not monophyletic and Hadrotes wakefieldi Cameron is the sister‐species of the rest of the Creophilus complex, but included within it. Creophilus is revised to include 12 species, including Creophilus galapagensis sp. nov. and C. rekohuensis sp. nov. Creophilus huttoni (Broun) is removed from synonymy with C. oculatus (Fabricius), and C. insularis (Fauvel), C. villipennis Kraatz, and C. violaceus (Fauvel) are synonymized under C. flavipennis (Hope) comb. et stat. nov. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 163 , 723–812.     

The role of character loss in phylogenetic reconstruction as exemplified for the Annelida

Annelid relationships are controversial, and molecular and morphological analyses provide incongruent estimates. Character loss is identified as a major confounding factor for phylogenetic analyses based on morphological data. A direct approach and an indirect approach for the identification of character loss are discussed. Character loss can frequently be found within annelids and examples of the loss of typical annelid characters, like chaetae, nuchal organs, coelomic cavities and other features, are given. A loss of segmentation is suggested for Sipuncula and Echiura; both are supported as annelid ingroups in molecular phylogenetic analyses. Moreover, character loss can be caused by some modes of heterochronic evolution (paedomorphosis) and, as shown for orbiniid and arenicolid polychaetes, paedomorphic taxa might be misplaced in phylogenies derived from morphology. Different approaches for dealing with character loss in cladistic analyses are discussed. Application of asymmetrical character state transformation costs or usage of a dynamic homology framework represents promising approaches. Identifying character loss prior to a phylogenetic analysis will help to refine morphological data matrices and improve phylogenetic analyses of annelid relationships.     

The phylogeny of charadriiform birds (shorebirds and allies) – reassessing the conflict between morphology and molecules

Multiple molecular analyses provide a congruent and well‐supported phylogeny of the charadriiform family‐level taxa, which conflicts with previous hypotheses based on osteological data. In order to revise the latter and to identify new characters of phylogenetic significance, skeletons of most charadriiform family‐level taxa were examined and 49 characters analysed. Tree topology was sensitive to outgroup choice, but the result of the analysis rooted with Columbidae (doves and pigeons) recovered a monophyletic Scolopaci, Charadrii, and nonturnicid Lari. With regard to the inclusion of Alcidae and Glareolidae in the Lari, the results of the present study are also in better concordance with the new molecular phylogenies than previous analyses of morphological data. Furthermore, for the first time an apomorphy of a clade including Thinocoridae, Pedionomidae, Rostratulidae, and Jacanidae was identified. Inclusion of Turnicidae in the Lari could not be supported, but there is no strong morphological evidence for an alternative placement. Pluvianus shares derived osteological features with the Burhinidae, and its position in the molecular analyses likewise cannot be corroborated with morphological data. Based on the topology of the molecular consensus tree, the ancestral state of selected characters is reconstructed. It is finally noted that recent calibrations of molecular analyses, which indicate an origin of extant charadriiform lineages in the Cretaceous, are based on incorrectly identified fossils. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 161 , 916–934.     

Taxonomic classification of the reef coral family Mussidae (Cnidaria: Anthozoa: Scleractinia)

Molecular analyses are transforming our understanding of the evolution of scleractinian corals and conflict with traditional classification, which is based on skeletal morphology. A new classification system, which integrates molecular and morphological data, is essential for documenting patterns of biodiversity and establishing priorities for marine conservation, as well as providing the morphological characters needed for linking present‐day corals with fossil species. The present monograph is the first in a series whose goal is to develop such an integrated system. It addresses the taxonomic relationships of 55 Recent zooxanthellate genera (one new) in seven families (one new), which were previously assigned to the suborder Faviina (eight genera are transferred to incertae sedis). The present monograph has two objectives. First, we introduce the higher‐level classification system for the 46 genera whose relationships are clear. Second, we formally revise the taxonomy of those corals belonging to the newly discovered family‐level clade (restricted today to the western Atlantic and Caribbean regions); this revised family Mussidae consists of ten genera (one of which is new) and 26 species that were previously assigned to the ‘traditional’ families Faviidae and Mussidae. To guide in discovering morphologic characters diagnostic of higher‐level taxa, we mapped a total of 38 morphologic characters [19 macromorphology, eight micromorphology, 11 microstructure] onto a molecular tree consisting of 67 species [22 Indo‐Pacific and seven Atlantic species in the traditional family Faviidae; 13 Indo‐Pacific and ten Atlantic species in the traditional family Mussidae; 13 species in the traditional families Merulinidae (5), Pectiniidae (7), and Trachyphylliidae (1); two Atlantic species of traditional Montastraea], and trace character histories using parsimony. To evaluate the overall effectiveness of morphological data in phylogeny reconstruction, we performed morphology‐based phylogenetic analyses using 27 (80 states) of the 38 characters, and compared morphological trees with molecular trees. The results of the ancestral state reconstructions revealed extensive homoplasy in almost all morphological characters. Family‐ and subfamily‐level molecular clades [previously identified as XVII?XXI] are best distinguished on the basis of the shapes of septal teeth and corresponding microstructure. The newly revised family Mussidae (XXI) has septal teeth with regular pointed tips (a symplesiomorphy) and a stout blocky appearance. It has two subfamilies, Mussinae and Faviinae. The subfamily Mussinae is distinguished by spine‐shaped teeth and widely spaced costoseptal clusters of calcification centres. The subfamily Faviinae is distinguished by blocky, pointed tricorne or paddle‐shaped teeth with elliptical bases, transverse structures such as carinae that cross the septal plane, and well‐developed aligned granules. Defining diagnostic characters for the broader data set is more challenging. In analyses of taxonomic subsets of the data set that were defined by clade, morphological phylogenetic analyses clearly distinguished the families Mussidae (XXI) and Lobophylliidae (XIX), as well as the two subfamilies of Mussidae (Mussinae, Faviinae), with one exception (Homophyllia australis). However, analyses of the entire 67‐species data set distinguished the family Lobophylliidae (XIX), but not the Merulinidae (XVII) and not the newly defined Mussidae (XXI), although the subfamily Mussinae was recovered as monophyletic. Some lower‐level relationships within the Merulinidae (XVII) agree with molecular results, but this particular family is especially problematic and requires additional molecular and morphological study. Future work including fossils will not only allow estimation of divergence times but also facilitate examination of the relationship between these divergences and changes in the environment and biogeography. Published 2012. This article is a U.S. Government work and is in the public domain in the USA. Zoological Journal of the Linnean Society, 2012, 166 , 465–529.     

Phylogeny and evolution of the South African genus Metalasia (Asteraceae–Gnaphalieae) inferred from molecular and morphological data

Metalasia is a genus in tribe Gnaphalieae (Asteraceae), endemic to South Africa and with its main distribution in the Cape Floristic Region. The genus comprises 57 species and, with a number of closely related genera, it constitutes the ‘Metalasia clade’. A species‐level phylogenetic analysis is presented, based on DNA sequences from two nuclear (internal and external transcribed spacer: ITS, ETS) and two plastid (psbA‐trnH, trnL‐trnF) regions together with morphological data. Analyses combining molecular and morphological data attempt not only to resolve species interrelationships, but also to detect patterns in character evolution. Phylogenetic analyses corroborate our earlier study and demonstrate that Metalasia is formed of two equally sized, well‐supported sister groups, one of which is characterized by papillose cypselas. The results differ greatly from earlier hypotheses based on morphology alone, as few morphological characters support the phylogenetic patterns obtained. The two clades of Metalasia do, however, appear to differ in distribution, corresponding to the different rainfall regimes of South Africa. Analyses show a few taxa to be problematic; one example is the widely distributed M. densa which appears to be an intricate species complex. © 2013 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 174 , 173–198.     

Phylogeny of Veneroidea (Mollusca: Bivalvia) based on morphology and molecules

The largest Recent family of Bivalvia, the marine Veneridae with approximately 800 species, comprises one of the least understood and most poorly defined molluscan taxa, despite including some of the most economically important and abundant bivalves, for example quahog, Pismo clams, and Manila clams. A review of previous phylogenetic analyses including the superfamily Veneroidea (Veneridae, Petricolidae, Glauconomidae, Turtoniidae, Neoleptonidae) and within the Veneridae shows minimal taxon sampling leading to weak conclusions and few supported synapomorphies. New phylogenetic analyses on 114 taxa tested the monophyly of Veneroidea, Veneridae, and 17 nominal venerid subfamilies, using morphological (conchological, anatomical) data and molecular sequences from mitochondrial (16S, cytochrome oxidase I) and nuclear (28S, histone 3) genes. Morphological analyses using 45 exemplar taxa and 23 traditional characters were highly homoplastic and failed to reconstruct traditional veneroid classification. Full morphological analyses (31 characters) supported the monophyly of Veneroidea and Veneridae but only when certain taxa were excluded, revealing analytical difficulties caused by a suite of characters associated with neotenous or miniaturized morphology. Molecular analyses resulted in substantially higher clade consistency. The combined molecular data set resulted in significant support for a particular topology. The monophyly of Veneridae was supported only when Petricolidae and Turtoniidae were subsumed, and recognized as members with derived or neotenous morphologies, respectively. Morphological character mapping on molecular trees retained a high level of homoplasy, but revealed synapomorphies for major branch points and supported six subfamily groups (Dosiniinae, Gemminae, Samarangiinae, Sunettinae, Tapetinae, combined Chioninae + Venerinae). Glauconomidae and Neoleptonidae are provisionally maintained in Veneroidea pending further study; Petricolinae and Turtoniinae are placed in Veneridae. © 2006 The Linnean Society of London, Zoological Journal of the Linnean Society, 2006, 148 , 439–521.     

Walk it off: predictive power of appendicular characters toward inference of higher‐level relationships in Laniatores (Arachnida: Opiliones)

Morphological characters are essential for establishing phylogenetic relationships, delimiting higher‐level taxa, and testing phylogenetic relationships inferred from molecular sequence data. In cases where relationships between large clades remain unresolved, it becomes imperative to establish which character systems are sound predictors of phylogenetic signal. In the case of Laniatores, the largest suborder of Opiliones, some superfamilial relationships remain unresolved or unsupported, and traditionally employed phenotypic characters are typically of utility only at the family level. Here we investigated a promising set of morphological characters that can be discretized and scored in all Opiliones: cuticular structures of the distal podomeres (metatarsi and tarsi). We intensively sampled members of all known families of Laniatores, and define here three new, discrete appendicular characters toward refinement of Laniatores superfamilial systematics: metatarsal paired slits (MPS; occurring in all Laniatores except Sandokanidae), proximal tarsomeric gland (PTG; in Icaleptidae, Fissiphalliidae, and Zalmoxidae), and tarsal aggregate pores (TAP; found in Gonyleptoidea, Epedanoidea, and Pyramidopidae). We conducted statistical tests on each character to characterize the strength of phylogenetic signal and assess character independence, based on alternative tree topologies of Laniatores. All three characters had high retention indices and bore significantly strong phylogenetic signal. Excepting one pairwise comparison, morphological characters did not evolve in a correlated manner, indicating that appendicular morphology does not constitute a single character system. Our results demonstrate the predictive power and utility of appendicular characters in Opiliones phylogeny, and proffer a promising source of diagnostic synapomorphies for delimiting superfamilies.     

Inclusive taxon sampling suggests a single,stepwise origin of ectolecithality in Platyhelminthes

Ectolecithality is a form of oogenesis unique within Metazoa but common in Platyhelminthes, in which almost yolkless oocytes and tightly associated yolk cells are deposited together in egg capsules. Despite profound impacts on the embryogenesis and morphology of its beneficiaries, the origins of this developmental phenomenon remain obscure. Traditionally, all ectolecithal flatworms were grouped in a clade called Neoophora. However, there are also morphological arguments for multiple origins of ectolecithality and, to date, Neoophora has seen little support from molecular phylogenetic research, largely as a result of gaps in taxon sampling. Accordingly, we present a molecular phylogeny focused on resolving the deepest divergences among the free‐living Platyhelminthes. Species were chosen to completely span the diversity of all major endo‐ and ectolecithal clades, including several aberrant species of uncertain systematic affinity and, additionally, a thorough sampling of the ‘lecithoepitheliate’ higher taxa Prorhynchida and Gnosonesimida, respectively, under‐ and unrepresented in phylogenies to date. Our analyses validate the monophyly of all classical higher platyhelminth taxa, and also resolve a clade possessing distinct yolk‐cell and oocyte generating organs (which we name Euneoophora new taxon ). Furthermore, implied‐weights parsimony and Bayesian mixture model analyses suggest common ancestry of this clade with the lecithoepitheliates, implying that these taxa may retain a primitive form of ectolecithality. This topology thus corroborates the classical hypothesis of homology between yolk cells and oocytes in all Neoophora, and should serve to guide future evolutionary research on this unique developmental innovation in Platyhelminthes. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 570–588.     

When molecules and morphology clash: reconciling conflicting phylogenies of the Metazoa by considering secondary character loss

Molecular and morphological data sets have yielded conflicting phylogenies for the Metazoa. So far, no general explanation for the existence of this conflict has been suggested. However, I believe that a neglected aspect of metazoan cladistics has introduced a systematic and substantial bias into morphological phylogenetic analyses. Most characters used for metazoan cladistics are coded as binary absence/presence characters. For most of these characters, the absence states are assumed to be uninformative default plesiomorphies, if they are defined at all. This character coding strategy could seriously underestimate the number of informative apomorphic absences or secondary character losses. Because nodes in morphological metazoan phylogenies are typically supported by relatively small numbers of characters each with a potentially strong impact on tree topology, failure to distinguish between primary absence and secondary loss of characters before a cladistic analysis may mislead morphological cladistics. This may falsely suggest conflict with molecular phylogenies, which are not sensitive to this bias. To test the existence of this bias, I compare the phylogenetic placement of a variety of metazoan taxa in molecular and morphological trees. In all instances investigated here, phylogenetic conflict can be resolved by allowing for secondary loss of morphological characters, which were assumed to be primitively absent in cladistic analyses. These findings suggest that we should be cautious in interpreting the results of morphological metazoan cladistic analyses and additionally illustrate the value of a more functional approach to comparative morphology in certain circumstances.     

How useful are the recommended counts and indices in the systematics of the Octopodidae (Mollusca: Cephalopoda)

A morphological dataset based on 14 standard counts and indices was constructed for 68 specimens comprising 12 species of octopuses. This was used to construct distance matrices based on morphological characters. These matrices were compared with genetic distance matrices compiled during molecular phylogenetic analyses of the same 12 species using four mitochondrial and two nuclear genes. Mantel tests showed that there was significant congruence between the phenetic and genetic matrices, suggesting that the genetic signal is reflected in the morphological data set. Matrices of geographical distance were constructed for the 12 species based on the latitude, longitude, and depth of capture of 1726 individuals. These matrices never showed significant congruence with genetic data or with morphological data. Multivariate analysis of the morphological dataset suggests that these counts and indices, traditionally used for discriminating between species in cephalopods, do not show great discrimination at species level, but provide excellent discrimination at the generic level, and, as such, might be useful for resolving the generic placement of some problematic taxa. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 205–218.     

Testing concordance in species boundaries using acoustic,morphological, and molecular data in the field cricket genus Itaropsis (Orthoptera: Grylloidea,Gryllidae: Gryllinae)

In most taxa, species boundaries are inferred based on differences in morphology or DNA sequences revealed by taxonomic or phylogenetic analyses. In crickets, acoustic mating signals or calling songs have species‐specific structures and provide a third data set to infer species boundaries. We examined the concordance in species boundaries obtained using acoustic, morphological, and molecular data sets in the field cricket genus Itaropsis. This genus is currently described by only one valid species, Itaropsis tenella, with a broad distribution in western peninsular India and Sri Lanka. Calling songs of males sampled from four sites in peninsular India exhibited significant differences in a number of call features, suggesting the existence of multiple species. Cluster analysis of the acoustic data, molecular phylogenetic analyses, and phylogenetic analyses combining all data sets suggested the existence of three clades. Whatever the differences in calling signals, no full congruence was obtained between all the data sets, even though the resultant lineages were largely concordant with the acoustic clusters. The genus Itaropsis could thus be represented by three morphologically cryptic incipient species in peninsular India; their distributions are congruent with usual patterns of endemism in the Western Ghats, India. Song evolution is analysed through the divergence in syllable period, syllable and call duration, and dominant frequency. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 285–303.     

Grazers, shredders and filtering carnivores--the evolution of feeding ecology in Drusinae (Trichoptera: Limnephilidae): insights from a molecular phylogeny

We examined the phylogenetic relationships between species and genera within the caddisfly subfamily Drusinae (Trichoptera: Limnephilidae) using sequence data from two mitochondrial loci (cytochrome oxidase 1, large subunit rRNA) and one nuclear gene (wingless). Sequence data were analysed for 28 species from five genera from the subfamily. We analysed individual and combined data sets using a Bayesian Markov Chain Monte Carlo and a maximum parsimony approach and compared the performance of each partition for resolving phylogenetic relationships at this level. In terms of resolution and phylogenetic utility wingless outperformed the two mitochondrial gene partitions. Using both Shimodaira-Hasegawa and expected likelihood weights tests we tested several hypotheses of relationships previously inferred based on adult morphological characters. The data did not support the generic concept, or many previously proposed species groupings, based on adult morphology. In contrast, the molecular data correlated with the morphology and feeding ecology of larvae. Using Bayesian ancestral character state reconstructions we inferred the evolution of feeding ecology and relevant larval morphological characters. Our analyses showed that within the subfamily Drusinae two derived feeding types evolved. One of these--grazing epilithic algae--is otherwise unusual in the Limnephilidae and may have promoted the high degree of diversity in the Drusinae.