Morphotypes of the lacustrine ostracod <Emphasis Type="Italic">Limnocythere rionegroensis</Emphasis> Cusminsky &amp; Whatley from Patagonia,Argentina, shaped by aquatic environments

This study describes and quantifies morphological valve traits of the ostracod Limnocythere rionegroensis from Patagonian steppe lakes and explores their association with lake water characteristics. Surface ornamentation was examined by scanning electron and transmitted light microscopy, and valve size and shape were analyzed using morphometric techniques. Limnocythere rionegroensis shows remarkable variations in surface ornamentation, based on which three morphotypes (MI, MII, and MIII) were identified. Valves of morphotypes MI and MIII are larger, show slight to moderate external reticulation, and a higher shape variability, whereas MII is characterized by a very conspicuous reticulation, lower shape variability, and smaller valves. Outline analysis yielded a great shape disparity related to the dorsal margin slope. MI was found in sexual populations from euhaline to mesohaline ephemeral lake; MII occurs in parthenogenetic populations from mesohaline to oligohaline permanent or ephemeral lakes; and MIII, from both sexual and asexual populations, inhabits a broad range of environmental conditions in terms of salinity and stability. Limnocythere rionegroensis intraspecific variations may be caused by environmental parameters and genetic factors associated to reproductive strategies. These results contribute to the knowledge of extant L. rionegroensis morphological variability and provide additional clues to improve the environmental interpretation of fossil assemblages.     

The Afromontane Cladocera (Crustacea: Branchiopoda) of the Rwenzori (Uganda–D. R. Congo): taxonomy,ecology and biogeography

The timely characterization of high-altitude freshwater habitats allows an assessment of the diversity of its biota and provides the basis for monitoring community change. In this study, we investigate the Cladocera fauna of 29 water bodies (pools, freshwater lakes, and surrounding swamps sampled at various occasions between 2005 and 2009) in the Rwenzori Mountains (Uganda, D. R. Congo), which are part of the East African Sky Island Complex. All sites except one are located above 3700 m altitude. We include notes on the morphology, taxonomy, distribution, and ecology of each recorded taxon and describe a new species of the Alona rustica-group (Alona sphagnophila n.sp.; Chydoridae). We found 11 species of which seven are restricted to Lake Mahoma, the lowest lake in our study area (2990 m) (Alona affinis barbata, A. intermedia, Alonella exisa, Alonella nana, Daphnia cf. obtusa, Pleuroxus aduncus) and/or Lake Bujuku (Daphnia cf. curvirostris, P. aduncus) (3900 m). Two taxa (Ilyocryptus cf. gouldeni, A. sphagnophila n.sp.) are restricted to Carex/Sphagnum bogs surrounding lakes in the afroalpine zone. Pigmented populations of Chydorus cf. sphaericus occur in all the sites. It is the only cladoceran species surviving the extreme alpine and nival conditions in the Rwenzori. The species is joined by A. guttata at locations at lower altitudes (ca. 3000–4000 m), present in about half of the sites. The Rwenzori Cladocera fauna is characterized by a strong extratropical temperate component and a low level of speciation/endemism. Harboring an impoverished boreal cladoceran community, Lake Mahoma is given closer attention. At 2990 m, the lake is a cold-temperate aquatic island in the tropics and may function as a stepping stone for Palaearctic taxa. We introduce a new term for high-altitude, cold-water habitats in the tropics, which act as climatic islands for extratropical freshwater faunas, Löffler Islands, in honor of Dr Heinz Löffler. In comparison to surveys in 1961, we list five new records in Lake Mahoma, which could indicate cladoceran community changes over the past few decades at ca. 3000 m in the Rwenzori. Since the species distributions correlate to temperature and catchment properties of the lakes, the Rwenzori cladoceran fauna can be expected as sensitive indicators for local changes.     

On the diversity of fishes in mountain lakes of the Amur basin

Results of the study of 11 lakes situated in the mountainous part of the Amur River basin are reported. Six fish species, namely, blunt-snouted lenok Brachymystax tumensis, Baikal-Lena grayling Thymallus baicalolenensis, Siberian stone loach Barbatula toni, Lagowski’s minnow Rhynchocypris lagowskii, Czekanowski’s minnow Rh. czekanowskii, and Amur sculpin Cottus szanaga, were found in five lakes. Four species were found in one lake (Lake Pereval’noe, basin of the Amgun’ River), and two species were found in each of the remaining lakes. Lenok and Amur sculpin were the most common species. The fauna of Lake Okonon (basin of the river Zeya) was the most specific and did not have analogs among the lake faunas; Czekanowski’s minnow and Baikal black grayling were found in this lake. The lenok ecotype that formed in the mountain lakes differed from the river lenok in some regards. The potential pathways of formation and protection of mountain lake ichthyocenoses of the Amur basin are discussed.     

Density dynamics of <Emphasis Type="Italic">Notholca squamula salina</Emphasis> Focke (Rotifera) in Lake Wujka,a freshwater Antarctic lake

The Antarctic Lake Wujka (62°09′28.3″S, 58°27′56.3″W), a shallow water body (Z _m  = 1.38 m), situated at c.15 m from the seashore was sampled at two points (Sp 1 and Sp 2) at 3-day intervals from December 2003 to June 2004. The two sampling points differing in location and depth: Sp 1 (Z _m  = 0.50 m) was the shallowest site, located near the lake outlet, while Sp 2 (Z _m  = 1.38 m) was the deepest spot of the lake. The population density of Notholca squamula salina peaked in June (at 114 ind. l^?1) at Sp 1, while at Sp 2 peaked in January (80 ind. l^?1) and May (150 ind. l^?1). Spearman non-parametric correlations with temperature, salinity, total dissolved solids, conductivity and pH revealed effects that characterize N. squamula salina as a species capable of surviving in a range of aquatic environments, but with a preference for high salinity, food and low temperature. It occurred in highest numbers when the diatom Achnanthes lanceolata var. rostrata (Øestrup) Hust., normally a benthic species, was stirred up into the water during storms that also raised the lake’s salinity to above 20 psu.     

Snake fauna (Reptilia: Serpentes) from the Early/Middle Miocene of Sandelzhausen and Rothenstein 13 (Germany)

The Early/Middle Miocene (European Land Mammal Zone MN5) localities of Sandelzhausen and Rothenstein 13 in southern Germany have yielded remains of about 13 ophidian taxa: Eoanilius sp. (Aniliidae), Bavarioboa ultima (Boidae), “Coluber” sp., ?Telescopus sp., Natrix sp., cf. Natrix sp., cf. “Neonatrix” sp., unidentified “colubrines” and “natricines” (Colubridae), Naja sp., an unidentified elapid (Elapidae), Vipera sp. (“Oriental viper”), Vipera sp. (“aspis complex”) (Viperidae). Both faunas document a transitional phase from those reported from several late Early and Middle Miocene sites of Central and Western Europe. The climates of Sandelzhausen and Rothenstein 13, as indicated by ophidian fossils, were warm, although not tropical or subtropical.     

Malcofaunal Diversity of Chilika Lake,Odisha, India

Investigation during the period of 3 years from 2007 to 2010 on the malacofauna of Chilika lake revealed the occurrence of 126 molluscan taxa belonging to 56 families, 18 orders of three classes in the bottom sediment. Of these 61 species belonged to Bivalvia, 64 species belonged to Gastropoda and one species belonged to Polyplacophora. Maximum Bivalvia and Gastropoda taxa were found in the outer channel region of the lake. The dominating species were Crassostrea cuttackensis, Saccostrea cucullata, Brachidontes undulatus, Meretrix meretrix among bivalves and Cerethideopsilla cingulata, Bullia vittata, Nassarious stolatus, Indothias lacera, Natica tigrina, Turritella attenuata were from the gastropods. Occurrence of a large number of marine taxa is most probably associated with the opening of new lagoon during 1st August 2008.     

Management of <Emphasis Type="Italic">Nymphoides peltatum</Emphasis> using water level fluctuations in freshwater lakes of Kashmir Himalaya

Lakes have an esthetic significance that is particularly important for attracting tourism. In this context, it is often preferable for lakes to have clear water, so many lake managers attempt to achieve clear lake water by various means. However, the lakes of Kashmir Himalaya are undergoing several complex ecological changes due to, for example, increasing tourism, overfishing, and intensive agriculture, which are making these lakes less clear. One such change is the vigorous growth and development of aquatic weeds in the shallow-water areas of Kashmir Himalayan lakes. We thus, investigated the response of Nymphoides peltatum, a rapidly multiplying clonal species, to water depth, in order to determine whether water depth can be used to control the spread of this proliferating macrophyte. Different traits of the given plant species, such as the mean number of ramets, were significantly higher (F = 55.412, p = 0.000) at depth zone D1 (0–100 cm) than at depth zones D2 (101–200 cm) and D3 (201–300 cm). In all of the lakes, mean spacer length—a tool for facilitating plant spread—was observed to be significantly higher (F = 45.890, p = 0.000) at lower water levels (0–100 cm). Also, the reproductive structures (flowers) of N. peltatum showed significant variation with depth (F = 51.909, p = 0.000) and with the lake examined (F = 9.909, p = 0.001). Thus, the results obtained during the present study indicate the importance of water depth in the management of N. peltatum in various Kashmir Himalayan lakes.     

A separate lowstand lake at the northern edge of Lake Tanganyika? Evidence from phylogeographic patterns in the cichlid genus <Emphasis Type="Italic">Tropheus</Emphasis>

In Lake Tanganyika, lake level fluctuations were shown to have had a major impact on the evolution of littoral species. Many species are subdivided into arrays of populations, geographical races and sister species, each colonizing a particular section of the shore. Their often limited dispersal abilities promoted geographic isolation and, on the long run, allopatric speciation. With more than 120 distinct populations, the genus Tropheus represents the most spectacular and best-studied example of this phenomenon. The present study aims at the fine-scale reconstruction of the spread of two mitochondrial Tropheus-lineages in the very north of the lake, where two species, T. sp. ‘black’ and T. brichardi, occur. Using mtDNA sequences and AFLP-data, we analyzed samples from 21 localities and found a highly complex conglomerate of introgressed populations formed by the repeated contact of two lineages. Our data suggest repeated cross-lake dispersal of T. sp. ‘black’ haplotypes along the ridge between the West and East Ubwari Fault, supporting an additional persisting lowstand-lake in the Bujumbura subbasin at the very north of the lake and highlighting once more the impact of lake level fluctuations on the genetic structure and evolution of stenotopic rock-dwelling cichlid species.     

A new species of Townsendia (Compositae) from Utah

An endemic, yellow ray-flowered species ofTownsendia is described asT. aprica from Sevier Co., Utah. It is apparently most closely related toT. montana Nutt. of western Montana and Wyoming south to southern Utah andT. mensana M. E. Jones of northern and western Utah and adjacent Nevada.     

Molecular characterisation of four echinostomes (Digenea: Echinostomatidae) from birds in New Zealand,with descriptions of <Emphasis Type="Italic">Echinostoma novaezealandense</Emphasis> n. sp. and <Emphasis Type="Italic">Echinoparyphium poulini</Emphasis> n. sp.

Morphological and molecular characterisation of echinostome specimens (Digenea: Echinostomatidae) recovered in one Anas platyrhynchos L. and one Cygnus atratus (Latham) (Anseriformes: Anatidae) from New Zealand revealed the presence of two known species, Echinostoma miyagawai Ishii, 1932 and Echinoparyphium ellisi (Johnston & Simpson, 1944) and two species new to science. Comparative morphological and phylogenetic analyses supported the distinct species status of Echinostoma novaezealandense n. sp. ex Branta canadensis (L.), A. platyrhynchos and C. atratus, and Echinoparyphium poulini n. sp. ex C. atratus. Echinostoma novaezealandense n. sp., a species of the “revolutum” species complex characterised by the possession of a head collar armed with 37 spines, keyed down to E. revolutum but was distinguished from the latter in having a much narrower body with almost parallel margins, longer oesophagus, wider cirrus-sac, larger seminal vesicle, much smaller ventral sucker, ovary, Mehlis’ gland and testes, more anteriorly located ovary and testes, and distinctly smaller eggs (81–87 × 42–53 vs 106–136 × 55–70 µm). This new species appears similar to Echinostoma acuticauda Nicoll, 1914 described in Australia but differs in having a longer forebody, more posteriorly located ovary and testes, and much smaller eggs (81–87 × 42–53 vs 112–126 × 63–75 µm). Echinoparyphium poulini n. sp. is differentiated from the four species of Echinoparyphium possessing 37 collar spines considered valid as follows: from E. chinensis Ku, Li & Chu, 1964 in having a much smaller body, four (vs five) angle spines and simple seminal vesicle (vs bipartite); from E. schulzi Matevosyan, 1951 in having a less robust body at a comparable body length, much smaller ventral sucker, ovary and testes, and longer but narrower eggs (87–109 × 50–59 vs 70–85 × 60–84 µm); and from the two smaller forms, E. serratum Howell, 1968 and E. aconiatum Dietz, 1909, in a number of additional metrical features correlated with body size and especially in the possession of much larger collar spines. Partial fragments of the mitochondrial nad1 and 28S rRNA genes were amplified for representative isolates of the four species and analysed together with sequences for Echinostoma spp. and Echinoparyphium spp. available on GenBank. Phylogenetic analyses based on the mitochondrial nad1 gene revealed congruence between the molecular data and species identification/delineation based on morphology; this was corroborated by the 28S rDNA sequence data.     

Abundance,distribution, and territory areas of rock-dwelling Lake Tanganyika cichlid fish species

Lake Tanganyika, the second-oldest and second-deepest lake in the world, harbors an impressive cichlid fish fauna counting about 250 endemic species that are characterized by a great level of ecological, morphological, and behavioral specialization. This study describes and compares cichlid fish communities at two rocky shores with differential human impact in the south of Lake Tanganyika. Species inventories and depth-dependent abundances were elaborated. About 41 and 46 sympatric cichlid species were recorded in the two study sites, respectively. Variabilichromis moorii was the most abundant species (29–60% of total number of fishes), followed by Aulonocranus dewindti (3–19%), Tropheus moorii (12%), Ophthalmotilapia ventralis (4–10%), Eretmodus cyanostictus (6–11%), and Cyathopharynx furcifer (0.01–9%). All other species had abundances below 5%. It further emerged that large cichlids such as Petrochromis species, Cyathopharynx furcifer, and Lobochilotes labiatus were very rare at one location, with frequencies of 0.55% or less. Territorial sizes of three particularly abundant species, Variabilichromis moorii, Aulonocranus dewindti, and Tropheus moorii, were assessed by behavioral observations. We distinguished between territorial core areas and total defended area, yielding average core areas between 0.4 (V. moorii) and 1.6 m² (T. moorii), and total defended areas averaging for each species between 1.6 (V. moorii) and 5.0 m² (A. dewindti) with no significant differences between the two study sites. The data on individual densities are also relevant for evolutionary studies, in that they allow more accurate calculations of effective population sizes.     

Assessment of the ecological status of European surface waters: a work in progress

Medicine Lake is a small (165 ha), relatively shallow (average 7.3 m), intermediate elevation (2,036 m) lake located within the summit caldera of Medicine Lake volcano, Siskiyou County, California, USA. Sediment cores and high-resolution bathymetric and seismic reflection data were collected from the lake during the fall of 1999 and 2000. Sediments were analyzed for diatoms, pollen, density, grain size (sand/mud ratio), total organic carbon (TOC), and micro-scale fabric analysis. Using both ¹⁴C (AMS) dating and tephrochronology, the basal sediments were estimated to have been deposited about 11,400 cal year BP, thus yielding an estimated average sedimentation rate of about 20.66 cm/1,000 year. The lowermost part of the core (11,400–10,300 cal year BP) contains the transition from glacial to interglacial conditions. From about 11,000–5,500 cal year BP, Medicine Lake consisted of two small, steep-sided lakes or one lake with two steep-sided basins connected by a shallow shelf. During this time, both the pollen (Abies/Artemisia ratio) and the diatom (Cyclotella/Navicula ratio) evidences indicate that the effective moisture increased, leading to a deeper lake. Over the past 5,500 years, the pollen record shows that effective moisture continued to increase, and the diatom record indicates fluctuations in the lake level. The change in the lake level pattern from one of the increasing depths prior to about 6,000 cal year BP to one of the variable depths may be related to changes in the morphology of the Medicine Lake caldera associated with the movement of magma and the eruption of the Medicine Lake Glass Flow about 5,120 cal year BP. These changes in basin morphology caused Medicine Lake to flood the shallow shelf which surrounds the deeper part of the lake. During this period, the Cyclotella/Navicula ratio and the percent abundance of Isoetes vary, suggesting that the level of the lake fluctuated, resulting in changes in the shelf area available for colonization by benthic diatoms and Isoetes. These fluctuations are not typical of the small number of low-elevation Holocene lake records in the region, and probably reflect the hydrologic conditions unique to Medicine Lake.     

Rising variance and abrupt shifts of subfossil chironomids due to eutrophication in a deep sub-alpine lake

In response to anthropogenic eutrophication and global warming, deep-water oxygen depletion is expected to have large effects on freshwater lake biogeochemistry and resident communities. In particular, it has been observed that deep-water hypoxia may potentially lead to regime shifts of lake benthic communities. We explored such community shifts by reconstructing a high-resolution subfossil chironomid record from a sediment core collected in the sub-alpine lake Remoray in France. We identified an abrupt shift in chironomid composition triggered by the collapse of the dominant Sergentia coracina-type chironomids around 1980. We found that the collapse of Sergentia coracina type was coupled to a gradual increase in organic matter content in lake sediments caused by eutrophication. We concluded that the most probable cause for the collapse of Sergentia coracina type was a change in oxygen concentrations below the minimal threshold for larval growth. We also analyzed trends in variance and autocorrelation of chironomid dynamics to test whether they can be used as early warnings of the Sergentia collapse. We found that variance rose prior to the collapse, but it was marginally significant (Kendal rank correlation 0.71, p = 0.05), whereas autocorrelation increased but insignificantly and less strongly (Kendal rank correlation 0.23, p = 0.25). By combining reconstructions of ecosystem dynamics and environmental drivers, our approach demonstrates how lake sediments may provide insights into the long-term dynamics of oxygen in lakes and its impact on aquatic fauna.     

Chemical and molecular identification of the invasive termite <Emphasis Type="Italic">Zootermopsis nevadensis</Emphasis> (Isoptera: Archotermopsidae) in Japan

Numerous termite species have been introduced outside their native ranges by human transport, and some have become invasive. The dampwood termite Zootermopsis nevadensis (Hagen), which is native to western North America, has been introduced to and become established in Kawanishi City, Hyogo Prefecture, Japan. Zootermopsis nevadensis is subdivided into two subspecies based on cuticular hydrocarbon (CHC) phenotypes: Z. nevadensis nevadensis and Z. nevadensis nuttingi (Haverty and Thorne). Here, we identified Z. nevadensis in Japan as hybrids between the two subspecies. Chemical analysis showed the presence of 7,15-dimethylhenicosane and 5,17-dimethylhenicosane in the CHCs of Z. nevadensis in Japan, corresponding to the CHC phenotype of Z. n. nevadensis. Conversely, all mitochondrial cytochrome c oxidase subunit I sequences of Z. nevadensis in Japan were identical to sequences from Z. n. nuttingi and hybrids between the two subspecies from a native hybrid zone in California, USA. In addition, phylogenetic analysis showed that Z. nevadensis in Japan formed a clade with Z. n. nuttingi and hybrids between the two subspecies. Our results show discordance between the chemical and genetic features of Z. nevadensis in Japan, indicating that individuals of Z. nevadensis in Japan are hybrids between the two subspecies.     

Environmental predictors of the occurrence of exotic <Emphasis Type="Italic">Hydrilla verticillata</Emphasis> (L.f.) Royle and native <Emphasis Type="Italic">Egeria najas</Emphasis> Planch. in a sub-tropical river floodplain: the Upper River Paraná, Brazil

The Upper River Paraná Floodplain System comprises the rivers Ivinheima, Baía and Paraná, which with their associated waterbodies form three subsystems, each showing individual characteristics. Hydrilla verticillata recently invaded the Upper Paraná Floodplain, while Egeria najas is the native most abundant submersed macrophyte. A large flood-pulse, during January–March 2007, abruptly reduced macrophyte stands in many areas to near-zero and dispersed propagules over the entire floodplain. From April 2007 to April 2008, we conducted three surveys sampling for the presence–absence of H. verticillata and E. najas and environmental variables aiming to answer: (1) How rapid is the colonization–regeneration process for both species? (2) Which habitats seem to be more susceptible to their colonization? (3) Which environmental factors can best predict their occurrence? Neither H. verticillata nor E. najas colonized the Ivinheima subsystem. In the Baía subsystem, E. najas had only two occurrences while H. verticillata was not present. In the Paraná subsystem, E. najas predominantly occurred in river channels, but it was also common in floodplain lakes. In April 2007, it was found in 13% of the sites in the Paraná subsystem, increasing to 30% in November 2007 and reaching 34% in April 2008. H. verticillata did not successfully colonize floodplain lakes of the Paraná subsystem. In channels, it had 34% occurrence in April 2007, increasing to 62% in November 2007 and remaining at 62% in April 2008. The role of environmental variables in predicting species occurrence changed depending upon the scale of the analysis. Considering the whole Upper Paraná floodplain, water transparency followed by electrical conductivity were the strongest predictors for both species. Colonization by submersed plants seems improbable in the Ivinheima subsystem owing to its low water transparency besides frequent localized floods; in the Baía subsystem, it seems inhibited by transparency and low alkalinity. Considering just the Paraná subsystem, the proportion of organic matter in sediment, ten times higher in floodplain lakes than in channels, was the best predictor for H. verticillata occurrence (also related to water pH and transparency), while E. najas was only significantly explained by transparency.     

Zapaliinae,a new subfamily of Tithonian–Berriasian ataxioceratid ammonites

The recent recognition of the earliest Andean Tithonian ammonite fauna (Picunleufuense Zone) has allowed a balanced classification of the ataxioceratid ammonites, reflecting their phylogenetic relationships. The Picunleufuense Zone fauna, first recognized in the Neuquén Basin at the base of the Vaca Muerta Formation, is represented by numerous records of sparse ammonite faunas from Antarctica, the Austral Basin and the Tarapacá Basin (N. Chile and S. Peru). These Southeastern Pacific faunas include the Tithonian genera Indansites, Choicensisphinctes, Zapalia, Krantziceras (Tithonian–lower Berriasian) and Platydiscus. These five genera are grouped in a new subfamily established in this paper: Zapaliinae subfam. nov. (type genus: Zapalia). The origin of Indansites and Choicensisphinctes (possibly via Indanistes) from Lithacoceras is discussed. Zapalia must have been originated from Indanistes in the upper Proximus Zone. Platydiscus could have been originated from Krantziceras. The Tithonian Southeastern Pacific ataxioceratids (Zapaliinae subfam. nov.) form a group well separated from the Tithonian Indo-Madagascan Virgatosphinctinae.     

The carnivoran community from the Miocene of Sandelzhausen (Germany)

From the Bavarian Early/Middle Miocene (MN5) site Sandelzhausen, nine species of carnivoran mammals are identified including the hemicyonine ursid Hemicyon stehlini, the amphicyonids Amphicyon cf. major and Pseudarctos bavaricus, the mustelids Ischyrictis zibethoides and Martes cf. munki, the mephitid Proputorius pusillus, the viverrid Leptoplesictis cf. aurelianensis, the felid Pseudaelurus romieviensis, and finally the recently described barbourofelid Prosansanosmilus eggeri. With these taxa present, Sandelzhausen shows a carnivoran community typical, though deprived, for the Lower to Middle Miocene of Europe, but different from roughly contemporary Mediterranean faunas such as those from Çandir or Pa?alar in Turkey.     

Phenotypic plasticity and taxonomy of <Emphasis Type="Italic">Schloenbachia varians</Emphasis> (J. Sowerby, 1817) (Cretaceous Ammonoidea)

Taxonomic concepts of Early–Middle Cenomanian representatives of Schloenbachia Neumayr, 1875 are evaluated based on well-preserved, abundant faunas from NE Iran (Atamir Formation, Koppeh Dagh) and Germany (Essen Greensand and Baddeckenstedt formations at several localities in northern Germany; Meißen Formation of Saxony). From a single Lower Cenomanian horizon in the Atamir Formation, a complete suite of specimens of Schloenbachia varians (J. Sowerby, 1817), ranging from moderately involute, compressed and finely ornamented (“forma subplana”) to more evolute, depressed and strongly spinose forms (“forma ventriosa”) with all transitional morphologies, is documented. Also in the shallow-water faunas of the “Klippenfazies” of the Essen Greensand Formation at Mülheim-Broich and of the Meißen Formation, all morphotypes co-occur. Thus, these forms cannot be regarded as geographic subspecies or successive chrono-subspecies, but reflect a high degree of variability in shell form and ornament (phenotypic plasticity) in S. varians. Interestingly, strongly tuberculate forms are rare to absent in contemporaneous offshore settings as documented by the Baddeckenstedt Formation. The phenotypic plasticity in Early Cenomanian S. varians populations is explained by ecophenotypic variation along a depth (proximal–distal) gradient: strongly tuberculate, depressed forms reflect comparatively shallow, nearshore environments with higher water energy and predation pressure, whereas compressed, weakly ornamented morphs are forms of open (and deeper) marine waters. The complete range of different forms is also observed in micro- and macroconchs, suggesting that the morphological variability is not controlled by sexual dimorphism. Based on its similarity to S. varians, also the Middle Cenomanian Ammonites coupei Brongniart, 1822 may be placed in the synonymy of the former.     

Delineating species along shifting shorelines: <Emphasis Type="Italic">Tropheus</Emphasis> (Teleostei,Cichlidae) from the southern subbasin of Lake Tanganyika

Background

Species delineation is particularly challenging in taxa with substantial intra-specific variation. In systematic studies of fishes, meristics and linear measurements that describe shape are often used to delineate species. Yet, little is known about the taxonomic value of these two types of morphological characteristics. Here, we used Tropheus (Teleostei, Cichlidae) from the southern subbasin of Lake Tanganyika to test which of these types of characters best matched genetic lineages that could represent species in this group of stenotypic rock-dwelling cichlids. We further investigated intra-population variation in morphology. By linking this to a proxy of a population’s age, we could assess the evolutionary stability of different kinds of morphological markers.

Results

Morphological data was collected from 570 specimens originating from 86 localities. An AFLP approach revealed the presence of five lineages in the southern subbasin: T. moorii, T. brichardi, T. sp. ‘maculatus’, T. sp. ‘Mpimbwe’ and T. sp. ‘red’, which we consider to represent distinct species. Although both types of morphological data supported this classification, a comparison of P_ST-values that describe inter-population morphological differentiation, revealed a better correspondence between the taxon delineation based on AFLP data and the patterns revealed by an analysis of meristics than between the AFLP-based taxon delineation and the patterns revealed by an analysis of shape. However, classifying southern populations of Tropheus was inherently difficult as they contained a large amount of clinal variation, both in genetic and in morphological data, and both within and among species. A scenario is put forward to explain the current-day distribution of the species and colour varieties and the observed clinal variation across the subbasin’s shoreline. Additionally, we observed that variation in shape was larger in populations from shallow shores whereas populations from steep shores were more variable in meristics. This difference is explained in terms of the different timescales at which small and large scale lake level fluctuations affected populations of littoral cichlids at steep and shallow shores.

Conclusions

Our results showed meristics to be more evolutionary stable, and of higher taxonomic value for species delimitation in Tropheus, than linear measurements that describe shape. These results should be taken into account when interpreting morphological differences between populations of highly stenotypic species, such as littoral cichlids from the Great East African Lakes.

     

Chemical defences of native European coccinellid eggs against intraguild predation by the invasive Asian coccinellid, <Emphasis Type="Italic">Harmonia axyridis</Emphasis> (Pallas)

Harmonia axyridis (Pallas) is a coccinellid of Asian origin that has recently invaded substantial parts of Europe and is suspected to affect native coccinellid populations through intraguild predation and competition for food. Previous work has shown that two species from the Calvia genus appeared to be well protected against H. axyridis predation. To deepen our understanding on chemical protection of Calvia spp. and the predation risk by H. axyridis, we tested for susceptibility and palatability of Calvia spp. and H. axyridis eggs against predation by H. axyridis neonate larvae. Results show that eggs of C. quatuordecimguttata were mostly not eaten by H. axyridis, while eggs of the congeneric C. decemguttata were found to be largely unprotected against predation by the invasive coccinellid. We also observed that H. axyridis first instars successfully cannibalized on conspecific eggs. Removing the surface chemicals from C. quatuordecimguttata eggs resulted in significantly reduced protection from being preyed upon by H. axyridis, while applying these extracts onto C. decemguttata and H. axyridis eggs resulted in increased protection against H. axyridis larvae. The importance of surface chemicals in the interactions between H. axyridis and native coccinellids was confirmed by GC–MS analysis, showing a high diversity of hydrocarbons located on the surface of C. quatuordecimguttata eggs, i.e. more than twice as many when compared to C. decemguttata. Survival of H. axyridis larvae feeding on eggs of C. quatuordecimguttata, C. decemguttata or conspecific eggs, from which surface chemicals were removed by washing them with hexane, was not different from survival on unwashed eggs.