A revision of Afrotropical Agabus Leach (Coleoptera, Dytiscidae), and the evolution of tropicoalpine super specialists

Abstract. The thirteen species of Agabus Leach, 1817, of the Afrotropical Region are revised and classified into four species groups: the dibasic ambulator -group endemic to Ethiopia, ragazzii -group with five species in Ethiopia, raffrayi -group with five species in East and South Africa, and A.discicollis Ancey, 1882, endemic to Ethiopia, is placed in the Nearctic cordatus -group. Agabus perssoni sp.n. and Agabus galamensis sp.n. are described from the highlands of Ethiopia. Agabus ruwenzoricus Guignot, 1936, stat.n. is given specific rank. Agabus ferrugatus Régimbart, 1905, is synonymized with Agabus ragazzii Régimbart, 1887, and Agabus limbicollis Régimbart, 1905, is synonymized with Agabus raffrayi Sharp, 1882, syn.n. Lectotypes are designated for the following nominal species: Agabus discicollis Ancey, 1882, Agabus raffrayi Sharp, 1882, Agabus limbicollis Régimbart, 1905, Agabus pallidus Omer-Cooper, 1931, Agabus ruwenzoricus Guignot, 1936, Agabus sjostedti Régimbart, 1908, Agabus dytiscoides Régimbart, 1908, Agabus ragazzii Régimbart, 1887, Agabus ferrugatus Régimbart, 1905, and Gaurodytes abessinicus Zimmermann, 1928. Reconstructed phylogenies are presented for the raffrayi -and ragazzii -groups. The following adaptations to tropicoalpine habitats are suggested: (1) body and especially pronotum narrow, (2) head broad anteriorly, (3) hindwing small, and (4) hind leg long and slender. Biogeography and evolution of the studied species are discussed. Four different invasions of Holarctic lineages to the East African mountains are suggested. Recurrent periods of cold and dry climate are considered the chief driving force in the evolution of high altitude super specialists.     

Description of a new Homidia species (Collembola,Entomobryidae) from Korea

A new species, Homidia pseudoformosana from Korea is described in the present paper. The new species is characterized by labral papillae, ciliated labial seta L₁ and dorsal chaetotaxy of abdominal segment IV. The detailed differences between similar Homidia species are provided.     

Speciation and mitochondrial DNA diversification of the diving beetles Agabus bipustulatus and A. wollastoni (Coleoptera, Dytiscidae) within Macaronesia

Diversification of populations of two morphologically similar diving beetles within the Agabus tristis group, A. wollastoni and A. bipustulatus, was investigated, with partial mtDNA cytochrome b (Cyt b) sequences, allozymes and landmark‐based morphometrics. The Madeiran endemic A. wollastoni was collected from 11 localities. Population genetic and morphological variation was compared to Scandinavian localities of the widespread west Palearctic A. bipustulatus, recorded also from the Azores. Agabus wollastoni and European A. bipustulatus specimens representing eight and 13 localities respectively, were used in evaluating their phylogenetic relationship. Maximum parsimony analysis of the Cyt b sequences showed that both the A. bipustulatus and A. wollastoni specimens form well‐supported monophyletic groups. Three lines of evidence suggest that Agabus wollastoni has speciated through a few founders: (1) a well‐supported mtDNA line; (2) the mean heterozygosity of A. wollastoni is lower when compared to A. bipustulatus on the mainland; and (3) several uncommon alleles of A. bipustulatus are missing in A. wollastoni. The Azorean A. bipustulatus population was drastically affected by the colonization event, since several loci have become fixed with a resulting lower mean heterozygosity. The colonization was relatively recent, as the mtDNA lineage represented in the Azores is deeply nested within the A. bipustulatus clade. Population structure shows moderate inbreeding of A. wollastoni, and extensive substructuring at all localities with moderate gene‐flow between them. Morphological variation in A. wollastoni showed significant differentiation among several populations. Island colonizations, population structure of A. wollastoni, and an observed pattern of variation of the α‐glycerophosphate dehydrogenase locus are discussed. © 2003 The Linnean Society of London, Biological Journal of the Linnean Society, 2003, 79, 653–666.     

Bees of the genus <Emphasis Type="Italic">Rophites</Emphasis> Spinola (Hymenoptera,Halictidae, Rophitinae) of Russia and adjacent territories

An illustrated key to Rophites species and an annotated list of 9 species from Russia and adjacent territories are given. Rophites clypealis stepposus subsp. n. and the female of R. schoenitzeri Dubitzky, 2004 are described. Lectotypes of Rhophites caucasicus Morawitz, 1876 and Rh. mandibularis Morawitz, 1891 are designated. Rophites algirus is recorded for the first time from Armenia; R. caucasicus, from Azerbaijan; R. clypealis, from Bulgaria, Russia, and Kazakhstan; R. foveolatus, from Georgia; R. quinquespinosus, from Armenia and Uzbekistan; R. schoenitzeri, from Azerbaijan and Iran.     

A mark and recapture study of water beetles (Coleoptera: Dytiscidae) in a group of semi-permanent and temporary ponds

A year mark and recapture study of aquatic Dytiscidae (Coleoptera) in a group of seven semi-permanent and temporary ponds in Cheshire (UK) is described. Pond water levels showed strong seasonal fluctuations with most ponds drying and refilling on several occasions over the study period. Sampling and labelling of aquatic Dytiscidae was carried out at monthly intervals. Beetles were labelled by fixing numbered waterproof paper labels to the elytra with cyanoacrylate `Superglue'. A total of 780 beetles were marked and released of which 11.5% were subsequently recaptured over varying time intervals of one month to eighteen months. The six species marked (Agabus bipustulatus (Linnaeus, 1767), Acilius sulcatus (Linnaeus, 1758), Colymbetes fuscus (Linnaeus, 1758), Dytiscus marginalis (Linnaeus, 1758), Ilybius ater (De Geer, 1774) and Ilybius fuliginosus (Fabricius, 1792)) are all known to be highly flight mobile. Colymbetes fuscus did not move between ponds as a result of ponds drying up in the summer. The behaviour of Dytiscus marginalis and Acilius sulcatus was found to be in broad agreement with previous accounts of dispersal from over-wintering ponds into temporary ponds in the spring with movement back to permanent ponds in the summer as the temporary ponds dry up. In contrast, Agabus bipustulatus did not migrate when ponds dried up in the summer but remained in terrestrial vegetation in damp pond basins for several months after open water had disappeared, only migrating if the basin dried out completely. Both species of Ilybius were captured and marked in very low numbers and no recaptures were made. As more ponds became temporary over the years of the study Agabus bipustulatus persisted while the other five species declined or were lost altogether.     

A new species of shrimp of the genus Anachlorocurtis Hayashi, 1975 from the Red Sea,with range extension of A. commensalis Hayashi, 1975 (Crustacea,Decapoda, Pandalidae)

A new species of pandalid shrimp Anachlorocurtis occidentalis sp. n., associated with antipatharian corals, is described and illustrated from the north-eastern Red Sea. This new species is closely related to Anachlorocurtis commensalis Hayashi, 1975, the only other species in the genus, and can be distinguished by the more slender body and appendages; the carapace with 3 large, and one small, subtriangular lobes in the middorsal line; a flattened dorsal outline of the third abdominal segment; the sixth abdominal segment twice as long as fifth one; propodi of the ambulatory pereiopods bearing only a single posterior spinule; and harbouring 3–5 pairs of dorsolateral spines on the telson. A revised generic diagnosis is provided here to accommodate the present new species. The genetic divergence of mitochondrial gene cytochrome c oxidase subunit I (COI) between Anachlorocurtis occidentalis sp. n., and A. commensalis is 15.2–15.4%. Molecular analysis also confirmed a sister position of the genus Anachlorocurtis to Miropandalus. The present records of A. commensalis from Taiwan constitute an extension of the known range of the species.     

Musculature, nervous system and glands of pregenital abdominal segments of the female of Nomia melanderi Ckll. (Hymenopters, Apoidea)

The gross anatomy of muscles, the topography of nerve tissues, and the histology of the pregenital abdominal glands of Nomia melanderi Ckll. are reported in detail. The movable and fixed points of muscle attachment were utilized in establishing a system of nomenclature for a typical abdominal segment. Names of nervules correspond to those of the tissues they innervate. The points of attachment of muscles of the fifth abdominal segment are essentially the same in both Nomia and Apis, except for the second tergo-sternal muscle which, in Nomia, has shifted its point of movable attachment to the membranous integument in front of the intersegmental membrane gland where it helps in relasing glandular secretion. The general plan of the nerves in the fifth abdominal segment in Nomia is more diffuse than in Apis, but there is no difficulty in establishing homology between the nervules of the two species. A pair of intersegmental stretch organs was found in abdominal segments 3–6. Glands of the sixth intersegmental membrane possess a reservoir with peripheral pouches both of which are absent in those of the fifth. Both types of glands have neither closing nor opening mechanisms, and neither is innervated. Release of glandular secretion is accomplished by the action of the tergo-sternal muscles.     

Embryonic development of a whirligig beetle,Dineutus mellyi,with special reference to external morphology (insecta: Coleoptera,Gyrinidae)

The egg morphology and successive changes of developing embryos of the whirligig beetle, Dineutus mellyi (Adephaga: Gyrinidae) are described from observations based on light and scanning electron microscopy. The egg surface is characterized by minute conical projections covering the entire egg surface, a stalk‐like micropylar projection at the anterior pole of the egg, and a longitudinal split line along which the chorion is cleaved during the middle embryonic stages. The germ band or embryo is formed on the ventral egg surface, and develops on the surface throughout the egg period; thus, the egg is a superficial type, as is the case in most coleopteran species. A pair of lateral tracheal gills (LTGs) of the first abdominal segment originates from appendage‐like projections arising at the lateral side of pleuropodia, and the LTGs of the second to ninth abdominal segments are arranged in a row with that of the first segment. Therefore, LTGs are structures with serial homology. The paired dorsal tracheal gills (DTGs) of the ninth abdominal segment are formed on the regions just latero‐dorsal to the LTGs of this segment. Regarding the pleuropodia as the structures being homologous with thoracic legs, neither the LTGs nor DTGs are homologous with thoracic legs, but originate in the more lateral region corresponding to the future pleura of the thoracic segments. The last (10th) abdominal segment in the larva is formed by the fusion of the embryonic 10th and 11th abdominal segments. Four terminal hooks at the end of the last abdominal segment originate from two pairs of swellings on the posterior end of the embryonic 11th abdominal segment. It is proposed that the terminal hooks possibly correspond to the claws of medially fused cerci of the embryonic 11th abdominal segment. J. Morphol. 2012. © 2012 Wiley Periodicals, Inc.     

A new species of Acentrella Bengtsson (Ephemeroptera: Baetidae) from Great Smoky Mountains National Park,USA

Acentrella barbarae sp. n. is described from eggs, larvae, and imagines collected from Great Smoky Mountains National Park, North Carolina and Tennessee, USA. Larvae were collected from cobble-pebble substrate in stream rapids. Life stages were associated by rearing. Eggs have longitudinal chorionic ridges with small, longitudinal furrows. Larvae are distinguished by having segment 3 of the labial palp apically rounded, abdominal terga with posteromedian projections, and tibiae and tarsi with relatively short setae on the outer margins. Imagines are distinguished by the colouration and posterior elevation of abdominal terga. Other Acentrella larvae occur in the same streams as the new species, including A. parvula, which we report from Tennessee for the first time. We provide an updated key to the larvae of North American Acentrella species.     

A new species of Erebonectes (Copepoda,Calanoida) from marine caves on Caicos Islands,West Indies

The insects of a small and cold aestival stream in northern Sweden were studied from semiquantitative kick-samples taken every ten days throughout the ice-free season in 1987. The stream froze solid at least in some sections in some winters. In addition to dipterans that were only identified to the family level, five ephemeropteran, four plecopteran, 11 coleopteran (Dytiscidae), and four trichopteran (Limnephilidae) species were found. Baetis bundyae Lemkuhl, Heptagenia orbiticola Kluge (Ephemeroptera), Nemoura cinerea (Retzius) (Plecoptera), Hydroporus palustris (L), Agabus setulosus (J. Sahlberg) (Coleoptera), Anabolia concentrica (Zetterstedt), Limnephilus coenosus Curtis, and Micopterna sequax McLachlan (Trichoptera) were all common. Malaise trap data are given for Plecoptera. The two ephemeropteran species Heptagenia orbiticola and Metretopus alter Bengtsson, and the plecopteran species Nemoura viki Lillehammer were recorded for the first time in Sweden. The remarkable species composition was attributed to the harsh abiotic and poorly studied environment. We predicted that the larval composition of this stream was representative for aestival streams in northernmost Fennoscandia.     

A new genus and species of scorpionfly (Mecoptera) from Baltic amber,with an unusually developed postnotal organ

A new genus and species of fossil scorpionflies (Mecoptera) Baltipanorpa damzeni gen. et sp.n. is described from two well‐preserved male specimens in Baltic amber (middle Eocene: Lutetian). The most characteristic feature of the new taxon is an unusually developed postnotal organ on abdominal tergum IV. This is the most extremely developed example of this organ among Mecoptera and the only observation of notal and postnotal organs among fossil scorpionflies. The following combination of characters are provided to distinguish the new genus from other Panorpidae: Sc, short; R₁ and R₂ two‐branched; A₁ joins posterior margin of wing only at same level as fork of vein Rs; unusual shape of abdomen, abdominal segments I–IV strongly reduced, abdominal segment V elongate and widened, segments VII and VIII strongly elongate; notal and postnotal organs present, strongly developed process (postnotal organ) on tergum IV, unknown in all described extant and fossil scorpionflies. Different types of notal organs of Mecoptera are compared and their function and morphology are discussed. Morphological analysis of notal and postnotal organs in extant species permits us to conclude that B. damzeni sp.n. is characterized by the most developed and complex notal organs in all Mecoptera.     

Phylogenetic analysis of Geotrupidae (Coleoptera, Scarabaeoidea) based on larvae

Abstract. Thirty‐eight characters derived from the larvae of Geotrupidae (Scarabaeoidea, Coleoptera) were analysed using parsimony and Bayesian inference. Trees were rooted with two Trogidae species and one species of Pleocomidae as outgroups. The monophyly of Geotrupidae (including Bolboceratinae) is supported by four autapomorphies: abdominal segments 3–7 with two dorsal annulets, chaetoparia and acanthoparia of the epipharynx not prominent, glossa and hypopharynx fused and without sclerome, trochanter and femur without fossorial setae. Bolboceratinae showed notable differences with Pleocomidae, being more related to Geotrupinae than to other groups. Odonteus species (Bolboceratinae s.str.) appear to constitute the closest sister group to Geotrupinae. Polyphyly of Bolboceratinae is implied by the following apomorphic characters observed in the ‘Odonteus lineage’: anterior and posterior epitormae of epipharynx developed, tormae of epipharynx fused, oncyli of hypopharynx developed, tarsal claws reduced or absent, plectrum and pars stridens of legs well developed and apex of antennal segment 2 with a unique sensorium. A ‘Bolbelasmus lineage’ is supported by the autapomorphic presence of various sensoria on the apex of the antennal segment, and the subtriangular labrum (except Eucanthus). This group constituted by Bolbelasmus, Bolbocerosoma and Eucanthus is the first evidence for a close relationship among genera, but more characters should be analysed to test the support for the clade. A preliminary classification at tribe level of Geotrupinae is suggested as follows: Chromogeotrupini (type genus Chromogeotrupes), Lethrini (type genus Lethrus), Taurocerastini (type genus Taurocerastes) and Geotrupini (type genus Geotrupes). Some ecological facts of Geotrupidae evolution could also be explained by the present results, such as those related to diet and nesting behaviour. Both coprophagy and male–female co‐operation in nesting appear as derived traits.     

A new coptoclavid larva (Coleoptera: Adephaga: Dytiscoidea) from the Middle Jurassic of China,and its phylogenetic implication

Daohugounectes primitivus, a new genus and species of coptoclavid beetles, is described from 67 fossil larvae from the Jurassic locality of Daohugou, northeastern China. It differs from other coptoclavids in the combination of the following characters: lateral lobes of nasale present; legs relatively short, with tarsi flat and slightly dilated; abdominal tergite VIII almost circular; derivative of abdominal segment IX present; urogomphi short; and helical thickening of tracheae weak. Most of these features are plesiomorphic for the family Coptoclavidae and the superfamily Dytiscoidea. Among dytiscoids, a derivative of abdominal segment IX is present only in the larvae of the recently discovered relic family Aspidytidae.     

Three new species of Euathlus Ausserer, 1875 (Araneae,Theraphosidae) from Argentina

ABSTRACT

Euathlus

Ausserer, 1875 is a South American genus of spiders of the family Theraphosidae known from Chile and Argentina. Three new species from Argentina: Euathlus mauryi sp. n. (from San Juan province), Euathlus grismadoi sp. n. (from La Rioja province), and Euathlus pampa sp. n. (from Salta province) are described here. New records of the previous species Euathlus diamante and Euathlus tenebrarum are contributed. The distribution of these new species expands the geographical distribution of the genus along the Andean hills, with Salta province as the northernmost record of the genus. A key is provided for identification of Euathlus species as well as a map with all the records in Argentina.     

Megapanorpa,a new genus with a single anal horn in males from Oriental China (Mecoptera: Panorpidae)

A new genus of Panorpidae, Megapanorpa gen. nov. , is erected for five new species, Megapanorpa grandis sp. nov. (type species), M. absens sp. nov. , M. gaokaii sp. nov. , M. jiangorum sp. nov. and M. wanghongjiani sp. nov. This genus is similar to Cerapanorpa Gao et al., 2016 in a single anal horn on the sixth tergum of male, but can be differentiated from the latter by the following characters: the seventh abdominal segment of male is constricted and stalk‐like at base, the subgenital plate of female with a pair of lateral plates, and the medigynium of female with a concealed axis. Keys to genera of Panorpidae and species of Megapanorpa gen. nov. are presented. The morphology and function of anal horns in different genera and species have been discussed.     

Precursor structures and evolution of tympanal organs in Lepidoptera (Insecta, Pterygota)

 The patterns of scolopal organs and their innervation were studied by the methylene blue method in larvae, pupae and adults of an Yponomeuta species (Yponomeutidae) and of tympanate adult representatives of the Noctuoidea, Geometridae, Drepanidae and Pyraloidea. The studies were focused mainly on the mesothorax, the metathorax and some anterior abdominal segments. In the abdominal tympanal organs of Geometridae, Drepanidae and Pyraloidea, the auditory scolopidia are homologous with the lateral scolopal organs of the first abdominal segment; however, the hearing organs as such evolved independently in the three taxa. The studies confirm that the tympanal organ in the Noctuoidea is derived from the caudal dorsolateral region of the metathorax including its dorsal scolopal organ and the B-cell. The adult scolopal organs are present already in the larvae and are maintained nearly unchanged during metamorphosis to the adult. Only in the Noctuoidea are the three sensory cells of the larval scolopal organs, which become part of the tympanal organs, reduced to one (in Notodontidae) or two (in other Noctuoidea) during metamorphosis. A hypothetical scenario of the evolution of the tympanal organs is outlined. Accepted: 12 March 1997     

A NEW SPECIES OF ARRHOPALITES FROM MT. CHANGBAI,CHINA (COLLEMBOLA:SMINTHURIDAE)*

Abstract A new species, Arrhopalites chungbaishanensis is described from Mt. Changbai, northeast China. The present species shares some characters with A. octacanthus, but it differs from the latter by the number of vertical thick spines and manubrial dorsal setae, types of dental setae and the absence of tunica on unguis. In comparison with the other congeneric species, the mast striking character of the present species is its special shapes of the C-series setae besetting on sixth abdominal segment.     

Water beetles (Coleoptera) of coastal areas of the Bolshezemelskaya Tundra,extreme northeastern Europe

The fauna of water beetles in coastal areas of the Bolshezemelskaya Tundra and on the Vaygach and Dolgiy islands, Pechora Sea, comprises a total of 42 recorded species; in local faunas, the number of species decreases northward from 19–26 in southern mainland tundra to nine in typical tundra of the Vaygach Island. The faunal similarity between pairs of areas is 21%–52% (39%–52% between mainland areas). The family Haliplidae is represented by three species, Dytiscidae – 32 species, Helophoridae – four species, Hydrophilidae – two species and Hydraenidae – one species. Most of the species have large ranges both longitudinally and latitudinally. Cryophile species are rather few (five species, 12%). There are clear differences between the beetle assemblages of weakly brackish lakes marshes and waterbodies of the bedrock coast. The strongest preferences for particular habitats have been revealed in species associated with sand substrates in waterbodies or with the floodplain of a large river. The male genitalia and male protarsal claws of Agabus luteaster (Zaitzev, 1906), recently recorded as new to Europe, are illustrated for the first time as well as Ochthebius kaninensis Poppius, 1909.     

The pupa of Calamoceras illiesi Malicky & Kumanski, 1974 (Trichoptera Calamoceratidae)

The previously unknown pupa of Calamoceras illiesi Malicky & Kumanski, 1974 is described and illustrated, based on material collected from northwestern Turkey. It is compared with Calamoceras marsupus Brauer, 1865, the second species of the genus Calamoceras found in western Europe. The pupa of C. illiesi is very different from that of C. marsupus, in particular in having a flattened abdomen, the arrangement of the transversal fringes of the abdominal segments, the shape of the sclerotised hook plate of abdominal segment I, and the shape of the anal processes.     

Stempellina and Zavrelia from China (Diptera,Chironomidae, Tanytarsini)

The first records of the Tanytarsini genera Stempellina and Zavrelia in the Oriental Region based on adult males are presented. Two species, Stempellina clavata sp. n. and Zavrelia clinovolsella sp. n. from southern China, are described, and the generic diagnoses of the two genera are emended to accommodate the new species. A key to the male imagines of the genus Zavrelia is given.