Another fly diuretic hormone: tachykinins increase fluid and ion transport by adult Drosophila melanogaster Malpighian 'renal' tubules
Insects such as the model organism Drosophila melanogaster must modulate their internal physiology to withstand changes in temperature and availability of water and food. Regulation of the excretory system by peptidergic hormones is one mechanism by which insects maintain their internal homeostasis. Tachykinins are a family of neuropeptides that have been shown to stimulate fluid secretion from the Malpighian 'renal' tubules (MTs) in some insect species, but it is unclear if that is the case in the fruit fly, D. melanogaster. A central objective of the current study was to examine the physiological role of tachykinin signaling in the MTs of adult D. melanogaster. Using the genetic toolbox available in this model organism along with in vitro and whole-animal bioassays, our results indicate that Drosophila tachykinins (DTKs) function as diuretic hormones by binding to the DTK receptor (DTKR) localized in stellate cells of the MTs. Specifically, DTK activates cation and anion transport across the stimulated MTs, which impairs their survival in response to desiccation because of their inability to conserve water. Thus, besides their previously described roles in neuromodulation of pathways controlling locomotion and food search, olfactory processing, aggression, lipid metabolism and metabolic stress, processing of noxious stimuli and hormone release, DTKs also appear to function as bona fide endocrine factors regulating the excretory system and appear essential for the maintenance of hydromineral balance.
Neogastropod (Mollusca, Gastropoda) phylogeny: A step forward with mitogenomes
The Neogastropoda (Mollusca, Gastropoda) encompass more than 15,000 described species of marine predators, including several model organisms in toxinology, embryology and physiology. However, their phylogenetic relationships remain mostly unresolved and their classification unstable. We took advantage of the many mitogenomes published in GenBank to produce a new molecular phylogeny of the neogastropods. We completed the taxon sampling by using an in-house bioinformatic pipeline to retrieve mitochondrial genes from 13 transcriptomes, corresponding to five families not represented in GenBank, for a final dataset of 113 taxa. Because mitogenomic data are prone to reconstruction artefacts, eight different evolutionary models were applied to reconstruct phylogenetic trees with IQTREE, RAxML and MrBayes. If the over-parametrization of some models produced trees with aberrant internal long branches, the global topology of the trees remained stable over models and softwares, and several relationships were revealed or found supported here for the first time. However, even if our dataset encompasses 60% of the valid families of neogastropods, some key taxa are missing and should be added in the future before proposing a revision of the classification of the neogastropods. Our study also demonstrates that even complex models struggle to satisfactorily handle the evolutionary history of mitogenomes, still leading to long-branch attractions in phylogenetic trees. Other approaches, such as reduced-genome strategies, must be envisaged to fully resolve the neogastropod phylogeny.
Hydroids (Cnidaria, Hydrozoa) from Mauritanian Coral Mounds
Agassiz, L. (1862) Contributions to the natural history of the United States of America. Vol. 4. Little Brown, Boston, 380 pp.Aguirrezabalaga, F., Altuna, A., Borja, A., Feliu, J., GarcíaCarrascosa, A.M., Romero, A., San Vicente, C., Torres-Gómez-de-Cádiz, J.A., Uriz, M.J. Ibánez, M. (1984) Contribución al conocimiento de la fauna marina de la costa Vasca. Π. Lurralde, Investigación y espacio, 1984, pp. 83-133.Alder, J. (1856) A notice of some new genera and species of British hydroid zoophytes. Annals and Magazine of Natural History, Series 2, 18, 353-362. https://doi.org/10.1080/00222935608697652Allman, G.J. (1873) Interim report on the hydroids collected by L.F. de Pourtalès during the Gulf Stream exploration of United States coast survey. Bulletin of the Museum of Comparative Zoology at Harvard College, 3 (7), 185-186.Allman, G.J. (1874) Report on the Hydroida collected during the expeditions of H.M.S. 'Porcupine.' Transactions of the Zoological Society of London, 8, 469-481. https://doi.org/10.1111/j.1096-3642.1874.tb00566.xAllman, G.J. (1877) Report on the Hydroida collected during the exploration of the Gulf Stream by L.F. de Pourtalès, assistant United States Coast Survey. Memoirs of the Museum of Comparative Zoology at Harvard College, 5 (2), 1-66. https://doi.org/10.5962/bhl.
Morphology of obligate ectosymbionts reveals gen. nov.: A new circumtropical genus of marine stilbonematine nematodes
Stilbonematinae are a subfamily of conspicuous marine nematodes, distinguished by a coat of sulphur-oxidizing bacterial ectosymbionts on their cuticle. As most nematodes, the worm hosts have a relatively simple anatomy and few taxonomically informative characters, and this has resulted in numerous taxonomic reassignments and synonymizations. Recent studies using a combination of morphological and molecular traits have helped to improve the taxonomy of Stilbonematinae but also raised questions on the validity of several genera. Here, we describe a new circumtropically distributed genus (Stilbonematinae) with three species: sp. nov., sp. nov. and sp. nov. We used single worm metagenomes to generate host 18S rRNA and cytochrome c oxidase I (COI) as well as symbiont 16S rRNA gene sequences. Intriguingly, COI alignments and primer matching analyses suggest that the COI is not suitable for PCR-based barcoding approaches in Stilbonematinae as the genera have a highly diverse base composition and no conserved primer sites. The phylogenetic analyses of all three gene sets, however, confirm the morphological assignments and support the erection of the new genus as well as corroborate the status of the other stilbonematine genera. most closely resembles the stilbonematine genus in overlapping sets of diagnostic features but can be distinguished from by the morphology of the genus-specific symbiont coat. Our re-analyses of key parameters of the symbiont coat morphology as character for all Stilbonematinae genera show that with amended descriptions, including the coat, highly reliable genus assignments can be obtained.
Molecular phylogeny of the Cyathocotylidae (Digenea, Diplostomoidea) necessitates systematic changes and reveals a history of host and environment switches
The Cyathocotylidae is a globally distributed family of digeneans parasitic as adults in fish, reptiles, birds, and mammals in both freshwater and marine environments. Molecular phylogenetic analysis of interrelationships among cyathocotylids is lacking with only a few species included in previous studies. We used sequences of the nuclear 28S rRNA gene to examine phylogenetic affinities of 11 newly sequenced taxa of cyathocotylids and the closely related family Brauninidae collected from fish, reptiles, birds, and dolphins from Australia, Southeast Asia, Europe, North America and South America. This is the first study to provide sequence data from adult cyathocotylids parasitic in fish and reptiles. Our analyses demonstrated that the members of the genus (family Brauninidae) belong to the Cyathocotylidae, placing the Brauninidae into synonymy with the Cyathocotylidae. In addition, our DNA sequences supported the presence of a second species in the currently monotypic . Our phylogeny revealed that spp. from crocodilians belong to a separate genus (, previously proposed as a subgenus) and subfamily (Suchocyathocotylinae subfam. n.). Morphological study of supported its elevation to species as . The phylogeny did not support within the subfamily Cyathocotylinae; instead, formed a strongly supported clade with members of the subfamily Szidatiinae ( and ). Therefore, we transfer into the Szidatiinae. DNA sequence data revealed the potential presence of cryptic species reported under the name . Our phylogeny indicated at least two major host switching events in the evolutionary history of the subfamily Szidatiinae which likely resulted in the transition of these parasites from birds to fish and snakes. Likewise, the transition to dolphins by represents another major host switching event among the Cyathocotylidae. In addition, our phylogeny revealed more than a single transition between freshwater and marine environments demonstrated in our dataset by and some .
Reappraisal of the Eocene whiptail stingrays (Myliobatiformes, Dasyatidae) of the Bolca Lagerstätte, Italy
The Eocene whiptail stingrays of the family Dasyatidae from the Bolca Lagerstätte, NE Italy, are revised herein in detail. The analysis of the anatomical and morphometric features allows us to identify the species "" (Molin, 1861) as a junior synonym of "" (Volta, 1796), and to assign it to the new genus gen. n. This new taxon exhibits a unique combination of features (e.g., rhombic disc wider than long, elongated tail folds fail to reach the tip of the tail, thorns absent, single serrated tail sting, "caniniform" teeth on upper jaw, tooth crown ornamentation absent, 175-179 vertebrae, 108-117 pectoral radials, 24-27 pelvic radials and other features of clasper anatomy) that clearly support its attribution to the subfamily Neotrygoninae of the stingray family Dasyatidae. The morphological and phylogenetic affinities of gen. n. with the living neotrygonines ( and ) suggest a close association of this taxon with the tropical shallow-water habitats hypothesized for the Bolca palaeoenvironment during the early Eocene. Moreover, the analysis of the fossil occurrences of the neotrygonines provides new insights into the role of the Tethys for the origin and evolutionary history of certain whiptail stingrays.
Form, function and phylogeny: comparative morphometrics of Lake Tanganyika's cichlid tribe Tropheini
Lake Tanganyika's cichlid fishes represent one of the most diverse species assemblages of the world. In this study we focused on the tribe Tropheini which occupies several trophic niches, mostly in rocky habitats. We analysed morphological variation of seventeen closely related species by means of geometric morphometric methods and related these data to ecological characteristics and phylogeny of the study species. It turned out that morphology mostly correlated well with ecological parameters, but not always closely with the degree of the phylogenetic relatedness of the species. Overall, body shapes in the tribe Tropheini are of great evolutionary plasticity, but variation is restricted to particular body parts: the preorbital region once again emerged as a key factor that facilitated their impressive radiation.
Paraphyly and budding speciation in the hairy snail (Pulmonata, Hygromiidae)
Delimitation of species is often complicated by discordance of morphological and genetic data. This may be caused by the existence of cryptic or polymorphic species. The latter case is particularly true for certain snail species showing an exceptionally high intraspecific genetic diversity. The present investigation deals with the complex, which has a complicated taxonomy. Our analyses of the sequence revealed that individuals showing a phenotype are distributed in nine highly differentiated mitochondrial clades (showing p-distances up to 19%). The results of a parallel morphometric investigation did not reveal any differentiation between these clades, although the overall variability is quite high. The phylogenetic analyses based on and sequences show that the s complex is paraphyletic with respect to several other morphologically well-defined species ( and ) which form well-supported monophyletic groups. The nc marker sequence (--) shows only a clear separation of and , and a weakly supported separation of , whereas all other species and the clades of the complex appear within one homogeneous group. The paraphyly of the complex reflects its complicated history, which was probably driven by geographic isolation in different glacial refugia and budding speciation. At our present state of knowledge, it cannot be excluded that several cryptic species are embedded within the complex. However, the lack of morphological differentiation of the mitochondrial clades does not provide any hints in this direction. Thus, we currently do not recommend any taxonomic changes. The results of the current investigation exemplify the limitations of barcoding attempts in highly diverse species such as .
Zebragryllus Desutter-Grandcolas & Cadena-Casteñada, n.gen. a new Gryllinae genus from Eastern and Western Amazonia, South America (Orthoptera, Grylloidea, Gryllidae)
We describe a new genus of grylline cricket, Zebragryllus Desutter-Grandcolas & Cadena-Casteñada n. gen., from the Neotropical Region, using characters of morphology and male genitalia; genitalic characters clearly show that Zebragryllus n. gen. is closely related to Anurogryllus Saussure, 1878. Six species are described as new to science, originating from western (Peru, Colombia) and eastern (French Guiana) Amazonia: Zebragryllus fuscus Desutter-Grandcolas, n. sp., Z. guianensis Desutter-Grandcolas, n. sp., Z. intermedius Desutter-Grandcolas, n. sp., Zebragryllus nauta Desutter-Grandcolas, n. sp., Zebragryllus nouragui Desutter-Grandcolas, n. sp., and Zebragryllus wittoto Desutter-Grandcolas and Cadena-Casteñada, n. sp., type species of the genus. They are characterized by their size, coloration (shining black, most often with white patterns of coloration, hence the genus name), and male and female genitalia. The calling songs of Z. guianensis Desutter-Grandcolas, n. sp., Z. intermedius Desutter-Grandcolas, n. sp., Z. nouragui Desutter-Grandcolas, n. sp., and Z. wittoto Desutter-Grandcolas and Cadena-Casteñada, n. sp. are described. An identification key is proposed for both males and females.
Molecular phylogeny of the Athetini-Lomechusini-Ecitocharini clade of aleocharine rove beetles (Insecta)
Elven, E., Bachmann, L. & Gusarov V. I. (2012) Molecular phylogeny of the Athetini-Lomechusini-Ecitocharini clade of aleocharine rove beetles (Insecta). -Zoologica Scripta, 41, 617-636.It has previously been shown that the Aleocharinae tribes Athetini and Lomechusini form a well-supported clade, which also includes the small Neotropical tribe Ecitocharini. However, neither Athetini nor Lomechusini were recovered as monophyletic. In this study, we addressed the basal phylogenetic relationships among the three tribes using sequence data from (i) a mitochondrial fragment covering the COI, Leu2 and COII genes; (ii) a mitochondrial fragment covering part of the 16S gene, the Leu1 gene and part of the NADH 1 gene; and (iii) a part of the nuclear 18S gene, for 68 Athetini, 33 Lomechusini and 2 Ecitocharini species, plus representatives from 10 other tribes. The athetine subtribe Geostibina was recovered as sister group to the 'true Lomechusini', which included the type genus Lomechusa. The two clades formed a sister group to the main Athetini clade, which also included Ecitocharini and the 'false Lomechusini', a group of New World genera normally placed in Lomechusini. The following changes in classification are proposed: (i) Geostibina Seevers, 1978 is raised to tribal rank, and 13 Athetini genera are placed in Geostibini; (ii) Ecitodonia Seevers, 1965; Ecitopora Wasmann, 1887, and Tetradonia Wasmann, 1894 are moved from Lomechusini to Athetini; (iii) Ecitocharini Seevers, 1965 is placed in synonymy with Athetini; (iv) Discerota Mulsant & Rey, 1874 is tentatively included in Oxypodini; (v) Actocharina Bernhauer, 1907 is placed in synonymy with Hydrosmecta Thomson, 1858.
Genetic divergence and geographic variation in the deep-water Conus orbignyi complex (Mollusca: Conoidea)
Puillandre, N. et al. (2010) Genetic divergence and geographic variation in a deep-water cone lineage: molecular and morphological analyses of the Conus orbignyi complex (Mollusca: Conoidea).The cone snails (family Conidae) are a hyperdiverse lineage of venomous gastropods. Two standard markers, COI and ITS2, were used to define six genetically-divergent groups within a subclade of Conidae that includes Conus orbignyi; each of these was then evaluated based on their shell morphology. We conclude that three forms, previously regarded as subspecies of Conus orbignyi are distinct species, now recognized as Conus orbignyi, Conus elokismenos and Conus coriolisi. In addition, three additional species (Conus pseudorbignyi, Conus joliveti and Conus comatosa) belong to this clade. Some of the proposed species (e.g., Conus elokismenos) are possibly in turn complexes comprising multiple species. Groups such as Conidae illustrate the challenges generally faced in species delimitation in biodiverse lineages. In the case of the Conus orbignyi complex, not only are there definable, genetically divergent lineages, but also considerable geographic variation within each group. Our study suggests that an intensive analysis of multiple specimens within a single locality helps to minimize the confounding effects of geographic variation and can be a useful starting point for circumscribing different species within such a confusing complex.
RNF213, a new nuclear marker for acanthomorph phylogeny
We show that RNF213 is a nuclear gene suitable for investigating large scale acanthomorph teleosteans interrelationships. The gene recovers many clades already found by several independent studies of acanthomorph molecular phylogenetics and considered as reliable. Moreover, we performed phylogenetic analyses of three other independent nuclear markers, first separately and then of all possible combinations (Dettaï, A., Lecointre, G., 2004. In search of nothothenioid (Teleostei) relatives. Antarct. Sci. 16 (1), 71-85. URL http://dx.doi.org/10.1017/S0954102004) of the four genes. This was coupled with an assessment of the reliability of clades using the repetition index of Li and Lecointre (Li, B., Lecointre, G., 2008. Formalizing reliability in the taxonomic congruence approach. Article accepted by Zoologica Scripta. URL http://dx.doi.org/10.1111/j.1463-6409.2008.00361.x). This index was improved here to handle the incomplete taxonomic overlap among datasets. The results lead to the identification of new reliable clades within the 'acanthomorph bush'. Within a clade containing the Atherinomorpha, the Mugiloidei, the Plesiopidae, the Blennioidei, the Gobiesocoidei, the Cichlidae and the Pomacentridae, the Plesiopidae is the sister-group of the Mugiloidei. The Apogonidae are closely related to the Gobioidei. A clade named 'H' grouping a number of families close to stromateids and scombrids (Stromateidae, Scombridae, Trichiuridae, Chiasmodontidae, Nomeidae, Bramidae, Centrolophidae) is related to another clade named 'E' (Aulostomidae, Macrorhamphosidae, Dactylopteridae). The Sciaenidae is closely related to the Haemulidae. Within clade 'X' (Dettaï, A., Lecointre, G., 2004. In search of nothothenioid (Teleostei) relatives. Antarct. Sci. 16 (1), 71-85. URL http://dx.doi.org/10.1017/S0954102004), the Cottoidei, the Zoarcoidei, the Gasterosteidae, the Triglidae, the Scorpaenidae, the Sebastidae, the Synanceiidae, and the Congiopodidae form a clade. Within clade 'L' (Chen, W.-J., Bonillo, C., Lecointre, G., 2003. Repeatability of clades as a criterion of reliability: a case study for molecular phylogeny of Acanthomorpha (Teleostei) with larger number of taxa. Mol. Phylogenet. Evol. 26, 262-288; Dettaï, A., Lecointre, G., 2004. In search of nothothenioid (Teleostei) relatives. Antarct. Sci. 16 (1), 71-85. URL http://dx.doi.org/10.1017/S0954102004) grouping carangoids with flatfishes and other families (Centropomidae, Menidae, Sphyraenidae, Polynemidae, Echeneidae, Toxotidae, Xiphiidae), carangids are the stem-group of echeneids and coryphaenids, and sphyraenids are the sister-group to the Carangoidei. The Howellidae, the Epigonidae and the Lateolabracidae are closely related. We propose names for most of the clades repeatedly found in acanthomorph phylogenetic studies of various teams of the past decade.
Comparative morphology of the venom apparatus in the braconid wasp subfamily Rogadinae (Insecta, Hymenoptera, Braconidae) and related taxa
Zaldivar-Rivern, A., Areekul, B., Shaw, M. R. & Quicke, D. L. J. (2004). Comparative morphology of the venom apparatus in the braconid wasp subfamily Rogadinae (Insecta, Hymenoptera, Braconidae) and related taxa. -, , 223-237. The morphology of the venom apparatus intima in representatives of 38 genera of the problematic braconid wasp subfamily Rogadinae and other cyclostome braconids was investigated and a preliminary phylogenetic analysis for the group was performed with the information obtained. Despite the limited number of characters, the data suggest several relationships at various taxonomic levels. The venom apparatus in the Clinocentrini and the Stiropiini is relatively unmodified and similar to that found in other genera previously placed within a broader concept of the Rogadinae (e.g. genera of Lysitermini, Pentatermini, Tetratermini, Hormiini) and also to that of the Betylobraconinae. The presence of a cone of filaments located inside the secondary venom duct near to its insertion on the venom reservoir/primary venom duct is proposed as a synapomorphy for the tribe Rogadini to the exclusion of Stiropiini, Clinocentrini and Yeliconini. Other features of the secondary venom duct and its insertion on the venom reservoir/primary venom duct support a number of relationships between the genera of the Rogadini and also within the large genus . A clade containing 15 Rogadini genera (, , , , , , , , , , , , , and ) is supported by the presence of a thickened and short secondary venom duct, whereas the different members of (excluding members of the subgenus ) and are distinguished by having a recessed secondary venom duct with well-defined and numerous internal filaments. New World species exhibit a unique venom apparatus and may not be closely related to the Old World ones. Features of the venom apparatus of the enigmatic genus and the exothecine genera and suggest that the Telengainae and Exothecinae are both closely related to the Braconinae, Gnamptodontinae, and possibly to the Opiinae and Alysiinae. An unsculptured venom reservoir was found in one specimen of the type species of , , which is consistent with it occupying either a very basal position within the cyclostome braconids or belonging to a recently recognized 'Gondwanan' clade that also includes the Aphidiinae.
Context of diversification of the viviparous Gyrodactylidae (Platyhelminthes, Monogenoidea)
Using four criteria proposed a decade ago by Brooks & McLennan to identify a case of adaptive radiation indicates that the evolutionary history of the viviparous clade of the Gyrodactylidae is dominated by nonvicariant processes. The viviparous clade, with 446 species, has significantly more species than its sister clade (one species), and high species richness was shown to be an apomorphic trait of only the viviparous gyrodactylids within the Gyrodactylidae. Reconciliation of the phylogenetic tree of the viviparous Gyrodactylidae with that of its hosts showed a low probability for cospeciation suggesting that adaptive modes of speciation and not vicariance were predominant during the historical diversification of the clade. The proposed hypothesis suggests that the Gyrodactylidae originated on the South American continent about 60 Mya after geographical dispersal and host switching of its common ancestor to demersal freshwater catfishes by a marine ancestor. Development of hyperviviparity and the consequent loss of 'sticky' eggs in conjunction with other symplesiomorphic and apomorphic features allowed rapid diversification coupled with high dispersal to new host groups and geographical areas by viviparous members of the Gyrodactylidae.
Discovery of sympatric cryptic species within (Gekkonidae: Squamata) from the Okinawa Islands, Japan, by use of allozyme data
An electrophoretic survey of samples of the gekkonid lizard, , from the East Asian islands demonstrated that two genetically divergent, but morphologically almost identical, entities occur on five islands of the Okinawa Group, Ryukyu Archipelago, Japan. These entities, while sharing all of the external character states diagnostic of , exhibited fixed allele differences at six to eight out of 30 loci examined and great overall genetic distances [Nei's (1978) = 0.489-0.654]. On Kumejima and Tonakijima Islands of the Okinawa Group, the two entities were collected together from identical microhabitats. These results indicate that the two entities represent separate biological species. Genetic comparisons of these two cryptic species from the Okinawa Group with '' from other island groups revealed that one occurs broadly in the insular region of East Asia, whereas the other is restricted to the Okinawa Group. Implications of the present findings for the morphological evolution of '' are also discussed.
Clypeatula cooperensis gen. n., sp. n., a new freshwater sponge (Porifera, Spongillidae) from the Rocky Mountains of Montana, USA
A new genus and species of freshwater sponge, Clypeatula cooperensis, collected from three lakes in the Northern Rocky Mountains of Montana, USA, are described. The sponge grows as a hard, disc-shaped encrustation on the undersides of rocks and logs. It lacks microscleres and has amphioxeal megascleres that often show a slight midregion bulb and are usually covered with short, conical spines except at their tips. The sponge is also non-gemmulating, overwintering in a regressed state in which choanocyte chambers are reduced in number. Phylogenetic analyses of complete 18S rDNA sequences of C. cooperensis, Ephydatia muelleri, Spongilla lacustris and Eunapius fragilis suggest that C. cooperensis is more closely related to Ephydatia muelleri than to Spongilla lacustris or Eunapius fragilis. Our data, nonetheless, do not rule out the possibility that C. cooperensis is more closely related to the non-gemmulating sponges of Lake Baikal (Russia) than it is to Ephydatia muelleri. These phylogenetic analyses support the erection of a new genus, the monophyly of freshwater sponges belonging to the families Spongillidae and Lubomirskiidae, and the monophyly of demosponges.
Monophyly of terrestrial adephagan beetles as indicated by three nuclear genes (Coleoptera: Carabidae and Trachypachidae)
The beetle suborder Adephaga is traditionally divided into two sections on the basis of habitat, terrestrial Geadephaga and aquatic Hydradephaga. Monophyly of both groups is uncertain, and the relationship of the two groups has implications for inferring habitat transitions within Adephaga. Here we examine phylogenetic relationships of these groups using evidence provided by DNA sequences from all four suborders of beetles, including 60 species of Adephaga, four Archostemata, three Myxophaga, and ten Polyphaga. We studied 18S ribosomal DNA and 28S ribosomal DNA, aligned with consideration of secondary structure, as well as the nuclear protein-coding gene wingless. Independent and combined Bayesian, likelihood, and parsimony analyses of all three genes supported placement of Trachypachidae in a monophyletic Geadephaga, although for analyses of 28S rDNA and some parsimony analyses only if Coleoptera is constrained to be monophyletic. Most analyses showed limited support for the monophyly of Hydradephaga. Outside of Adephaga, there is support from the ribosomal genes for a sister group relationship between Adephaga and Polyphaga. Within the small number of sampled Polyphaga, analyses of 18S rDNA, wingless, and the combined matrix supports monophyly of Polyphaga exclusive of Scirtoidea. Unconstrained analyses of the evolution of habitat suggest that Adephaga was ancestrally aquatic with one transition to terrestrial. However, in analyses constrained to disallow changes from aquatic to terrestrial habitat, the phylogenies imply two origins of aquatic habit within Adephaga.