Molecular phylogeny confirms Conochilidae as ingroup of Flosculariidae (Rotifera,Gnesiotrocha)

We test the hypothesis that conochilid rotifers represent an independent family‐level taxon within Superorder Gnesiotrocha, by analysing their phylogenetic position based on the 18S rDNA sequence of a large number of representatives of this taxon and its putative relatives. Both Bayesian and maximum likelihood analysis confirm a monophyletic clade of all gnesiotrochans with strong branch supports. Within Gnesiotrocha, Conochilidae form a strongly supported clade with representatives of all but some genera of Flosculariidae. These results refute Conochilidae as separate family‐level taxon within Gnesiotrocha. This finding is also supported by a phylogenetic analysis using morphology, in particular new observations on trophi morphology. Conochilid rotifers are likely specialized Flosculariidae, which evolved to a planktonic lifestyle and reduction of coloniality within the group, in contrast to other Flosculariidae. Furthermore, our analysis reveals that two genera of Flosculariidae, Beachampia and Limnias, form a single, strongly supported clade in a sister‐group relation to a clade consisting of representatives of Order Collothecacea. The present results, both regarding position of the conochild rotifers and of two genera of Flosculariidae, highlight the need for a more extensive analysis of relationships within Gnesiotrocha.     

Phylogeny of the families Pyuridae and Styelidae (Stolidobranchiata, Ascidiacea) inferred from mitochondrial and nuclear DNA sequences

The Order Stolidobranchiata comprises the families Pyuridae, Styelidae and Molgulidae. Early molecular data was consistent with monophyly of the Stolidobranchiata and also the Molgulidae. Internal phylogeny and relationships between Styelidae and Pyuridae were inconclusive however. In order to clarify these points we used mitochondrial and nuclear sequences from 31 species of Styelidae and 25 of Pyuridae. Phylogenetic trees recovered the Pyuridae as a monophyletic clade, and their genera appeared as monophyletic with the exception of Pyura. The Styelidae, on the other hand, appeared as a paraphyletic group split into several clades. One of them was formed by solitary oviparous species, of which the Pyuridae were a sister group. A second clade included the colonial genera Botryllus, Botrylloides and Symplegma. The remaining colonial and solitary genera formed several poorly resolved clades. One of the more species genus, Polycarpa, was shown to be polyphyletic, and the species Styela plicata grouped into two genetically distant clades suggesting the existence of two cryptic species. The internal phylogeny of Styelidae has bearings on the origin of coloniality in this family. We suggest to abandon the traditional division of colonial forms into social and compound species and use instead the categories of aggregated colonies that do not have common vascular systems, and integrated colonies, that do possess such systems. Our molecular results indicate that there have been several independent acquisitions of coloniality in the Styelidae, and that viviparity may be a pre-adaptation for a colonial life-style.     

Does predation select for or against avian coloniality? A comparative analysis

Some studies have supported predation as a selective pressure contributing to the evolution of coloniality. However, evidence also exists that colonies attract predators, selecting against colonial breeding. Using comparative analyses, we tested the reduced predation hypothesis that individuals aggregate into colonies for protection, and the opposite hypothesis, that breeding aggregations increase predation risk. We used locational and physical characteristics of nests to estimate levels of species' vulnerability to predation. We analysed the Ciconiiformes, a large avian order with the highest prevalence of coloniality, using Pagel's general method of comparative analysis for discrete variables. A common requirement of both hypotheses, that there is correlated evolution between coloniality and vulnerability to predation, was fulfilled in our data set of 363 species. The main predictions of the reduced predation hypothesis were not supported, namely that (1) solitary/vulnerable species are more prone to become colonial than solitary/protected species and (2) colonial/protected species are more likely to evolve towards vulnerability than solitary/protected species. In contrast, the main predictions of the increased predation hypothesis were supported, namely that colonial/vulnerable species are more prone (1) to become protected than solitary/vulnerable species and/or (2) to become solitary than colonial/protected species. This suggests that the colonial/vulnerable state is especially exposed to predation as coloniality may often attract predators rather than provide safety.     

A novel mode of colony formation in a hydrozoan through fusion of sexually generated individuals

Coloniality, as displayed by most hydrozoans, is thought to confer a size advantage in substrate-limited benthic marine environments and affects nearly every aspect of a species' ecology and evolution. Hydrozoan colonies normally develop through asexual budding of polyps that remain interconnected by continuous epithelia. The clade Aplanulata is unique in that it comprises mostly solitary species, including the model organism Hydra, with only a few colonial species. We reconstruct a multigene phylogeny to trace the evolution of coloniality in Aplanulata, revealing that the ancestor of Aplanulata was solitary and that coloniality was regained in the genus Ectopleura. Examination of Ectopleura larynx development reveals a unique type of colony formation never before described in Hydrozoa, in that colonies form through sexual reproduction followed by epithelial fusion of offspring polyps to adults. We characterize the expression of manacle, a gene involved in foot development in Hydra, to determine polyp-colony boundaries. Our results suggest that stalks beneath the neck do not have polyp identity and instead are specialized structures that interconnect polyps. Epithelial fusion, brooding behavior, and the presence of a skeleton were all key factors behind the evolution of this novel pathway to coloniality in Ectopleura.     

Mate fidelity and coloniality in waterbirds: a comparative analysis

Increased opportunities for information are one potential benefit of sociality. We apply this idea to the advantages of colonial breeding in bird species that are typically monogamous within a breeding season but often form new pair-bonds in subsequent seasons. Individuals may benefit from nesting in colonies at high density by identifying good-quality potential alternative mates among their neighbours. The opportunities for finding a better mating option are likely to increase with colony size and density. We tested this prediction with a comparative analysis of the association between mate fidelity and coloniality in waterbirds (wading birds and seabirds), where there is wide variation in both the degree of mate retention over consecutive breeding seasons and the degree of coloniality. We used two comparative statistical analyses, one based upon generalized least squares and the other based upon a continuous-time Markov model, to test whether the pattern of association between divorce rate and degree of coloniality was evidence for correlated evolutionary change in the two characters. We found a significant and positive association between divorce rate and the degree of coloniality in waterbirds. The probable ancestral state corresponds to a combination of a high degree of coloniality with no, or weak, mate fidelity. The reconstruction of the historical pattern of character origin and evolution indicates that the transition from a high to a low degree of coloniality occurred before the transition to higher mate fidelity. Received: 5 January 1998 / Accepted: 20 April 1998     

<Emphasis Type="Italic">Gloeocapsopsis aurea</Emphasis>, a new subaerophytic cyanobacterium from maritime Antarctica

The cyanobacterial flora of maritime Antarctica appears to contain many endemic species and only few cosmopolitan or wider-distributed taxa. Several morphospecies that have been erroneously identified in the past following available keys from temperate or tropical zones belong in fact to little-known and poorly described Antarctic cyanobacteria. Here we describe the taxonomy of one such example, the colonial species Gloeocapsopsis aurea . This cyanobacterium produces irregular, packet-like colonies that form black mats, films and crusts. Based on analysis of algal samples from Punta Cierva (Antarctic Peninsula) and King George Island (South Shetland Islands), this taxon is widely distributed in coastal, deglaciated areas of the maritime Antarctic. It is an important, often dominating, ecotype, mostly colonising irrigated rocks but also found in a variety of other aquatic and semi-aquatic habitats under a wide range of conductivities, pH and nutrient regimes.     

Variation of colony types in the antFormica cinerea

Summary Formica cinerea is a rare ant species in northern Europe where it occurs in few isolated populations. Estimates of genetic relatedness among worker nestmates revealed very different colonial structures. Relatedness was g = 0.81 in one population, and g = –0.03 and = 0.01 in two others. These results indicate that some populations of the species have mainly monogynous colonies (perhaps with monandrous queens), whereas others consist of polygynous and possibly polydomous colonies. Genetic differentiation of closely located populations suggests restricted dispersal.     

Coloniality has evolved once in Stolidobranch Ascidians

Ascidians exhibit a rich array of body plans and life history strategies. Colonial species typically consist of zooids embedded in a common test and brood large, fully developed larvae, while solitary species live singly and usually free-spawn eggs that develop into small, undifferentiated larvae. Ascidians in the order Stolidobranchia include both colonial and solitary species, as well as several species with intermediate morphologies. These include social species, which are colonial but do not live completely embedded in a common test, and a few solitary species that brood embryos and larvae until they are competent to metamorphose. We examined how many times coloniality has evolved within the Stolidobranchia, with phylogenetic analyses using full-length 18S rDNA and partial cytochrome oxidase B sequences for taxa in the families Molgulidae, Styelidae, and Pyuridae. Tunicata orders Phlebobranchia and Stolidobranchia are sister groups, and the family Molgulidae is a monophyletic group and should be raised to the subordinal level, as shown previously by analyses from this lab with partial 18S sequences. In contrast to previous studies, styelids and pyurids are separated into monophyletic groups by ML and Bayesian analyses. We show a single clade within the family Styelidae that contains two colonial (compound) botryllid species, a Symplegma (colonial compound), a colonial (social) species Metandrocarpa taylori, as well as four solitary species, thus confirming that the botryllids are a subfamily of the Styelidae. These results suggest that the ancestor of the Stolidobranchia was solitary and that coloniality has evolved only once within this clade of ascidians. Further phylogenetic analyses of aplousobranch and phlebobranch ascidians will be necessary to understand the number of times that coloniality has evolved within the class Ascidiacea.     

Phylogeny and classification of the Conochilidae (Rotifera, Monogononta, Flosculariacea)

To resolve several taxonomic problems within the family Conochilidae (Rotifera, Monogononta, Flosculariacea), we initiated a comparative study of the morphology in this and related taxa using samples collected from widely separated geographical regions. As part of this study, we paid special attention to trophal morphology using scanning electron microscopy. We also constructed and analysed a data matrix comprising 19 morphological characters of 11 taxa using cladistic methods to uncover all most-parsimonious trees. The results indicate that Conochilidae share a body form with Flosculariidae, but they possess a trophal structure which clearly differentiates them from all other Flosculariacea; thus, the diagnosis of the family Conochilidae is amended to incorporate morphological characters of the trophi. The analysis of our data matrix yielded a single, most-parsimonious tree. From the topology of that tree and our scanning electron microscopy observations, we propose the following: (1) the status of Conochilidae as a separate suborder of Flosculariacea is rejected; (2) taxonomic separation of Conochilus and Conochiloides as subgenera of Conochilus is confirmed; and (3) Lacinularia causeyae Vidrine, Mclaughlin & Willis, 1985 is reallocated to a new genus within the family Conochilide, Conochilopsis gen. nov., as Conochilopsis causeyae (Vidrine et al .) comb. nov.     

MORPHOLOGY AND NUTRITION OF PANDORINA UNICOCCA SP. NOV.1,2

Eleven strains of Pandorina unicocca sp. nov. were studied for morphological characterization. The species was delimited by having cells which contain a single basal pyrenoid and by colonies which have cells separate from each other in the colonial envelope. A defined medium was developed for 4 strains of P. unicocca and the nutritional requirements of these strains were examined. All 4 strains were capable of completely autotrophic growth and they could utilize nitrate, ammonium, or urea as nitrogen sources. Optimum growth was nuts attained at pH 8.0.     

Sessile rotifers (Rotifera) exhibit strong seasonality in a shallow,eutrophic Ramsar site in Mexico

To better understand the sessile rotifer community of a shallow, eutrophic lake, we monitored over a 1‐year period the monthly changes in the density and diversity of rotifers on the roots of the common water hyacinth, Eichhornia crassipes, which served as a representative hydrophyte. Selected physical and chemical parameters of the water were also determined. Mean annual density of all sessile species was 630 individuals per ml. Species of Collothecacea (n = 7) were most abundant. Of the 12 species of Flosculariaceae (Flosculariidae), Sinantherina socialis occurred seasonally, while Ptygura beauchampi was most common across all seasons. The Shannon–Wiener diversity index was highest (H′ = 2.8) in April, but decreased in January (H′ = 1.3). Applying the concept of the guild ratio (functional‐based analysis) we found that raptorial species (Collothecacea) were dominant over most of the year (June–February), while microphagous Flosculariidae were dominant during the warm, drier season (March–May). BEST (optimal matching of biota to environment, including stepwise search) analysis and canonical correspondence revealed that temperature, dissolved oxygen, pH, turbidity, and transparency strongly influenced sessile rotifer dynamics. We also found significant correlations between the density of some species and physicochemical variables. These species included Collotheca ambigua, C. coronetta, C. ornata, C. campanulata, C. tenuilobata, and Stephanoceros millsii in the Collothecacea; and Beauchampia crucigere, Limnias melicerta, L. ceratophylli, Ptygura crystallina, P. melicerta, and Sinantherina ariprepes in the Flosculariidae. Our study shows that the communities of sessile rotifers were structured by abiotic factors and that the guild ratio is useful for understanding the relationship between sessile rotifers and their substrata.     

Capsular polysaccharides facilitate enhanced iron acquisition by the colonial cyanobacterium Microcystis sp. isolated from a freshwater lake

Microcystis sp., especially in its colonial form, is a common dominant species during cyanobacterial blooms in many iron‐deficient water bodies. It is still not entirely clear, however, how the colonial forms of Microcystis acclimate to iron‐deficient habitats, and the responses of unicellular and colonial forms to iron‐replete and iron‐deficient conditions were examined here. Growth rates and levels of photosynthetic pigments declined to a greater extent in cultures of unicellular Microcystis than in cultures of the colonial form in response to decreasing iron concentrations, resulting in the impaired photosynthetic performance of unicellular Microcystis as compared to colonial forms as measured by variable fluorescence and photosynthetic oxygen evolution. These results indicate that the light‐harvesting ability and photosynthetic capacity of colonial Microcystis was less affected by iron deficiency than the unicellular form. The carotenoid contents and nonphotochemical quenching of colonial Microcystis were less reduced than those of the unicellular form under decreasing iron concentrations, indicating that the colonial morphology enhanced photoprotection and acclimation to iron‐deficient conditions. Furthermore, large amounts of iron were detected in the capsular polysaccharides (CPS) of the colonies, and more iron was found to be attached to the colonial Microcystis CPS under decreasing iron conditions as compared to unicellular cultures. These results demonstrated that colonial Microcystis can acclimate to iron deficiencies better than the unicellular form, and that CPS plays an important role in their acclimation advantage in iron‐deficient waters.     

Foraging advantages of mixed-species association between solitary and colonial orb-weaving spiders

This study investigated association between solitary orb-weaving spiders and a colonial orb-weaving spider, Metepeira incrassata (Araneae: Araneidae). Spiders were sampled along transects and an index of species association showed that two of the species were associated more frequently than expected based on a null hipothesis of random co-occurrence. The potential advantages of mixed-species association were investigated by comparing prey-capture success of one of these associates, Nephila clavipes (Araneae: Tetragnathidae), when it occurs alone, in single-species groups, and when associated with M. incrassata colonies. Field observations of prey-capture success by all three of these categories of N. clavipes revealed that individuals in M. incrassata colonies captured significantly more prey than solitaries or individuals in single-species groups. The increase in prey capture by N. clavipes in M. incrassata colonies may result from utilization of a foraging niche which intercepts a diffirent spectrum of prey than that available to single-species groups or solitaries. Related to this enhanced prey consumption is greater fecundity of spiders in association with M. incrassata as compared to solitaries or individuals in single-species groups.     

The Protector Species Hypothesis: Do Black Skimmers Find Refuge from Predators in Gull-billed Tern Colonies?

The protector-species hypothesis explains mixed-species coloniality on the basis of benefits individuals of a species may receive by nesting with another species, the ‘protector’ species, that responds aggressively to potential threats. The reactions of nesting individuals to both natural and model predators were observed to determine whether black skimmers (Rhynchops niger) gain an antipredator advantage by nesting with gull-billed terns (Sterna nilotica). Observations of natural predators were gathered from three mixed-species and three single-species (black skimmers) subcolonies. Natural predators most commonly encountered by the colonies were herring gulls (Larus argentatus), laughing gulls (Larus atricilla), and ruddy turnstones (Arenaria interpres). Gull-billed terns responded to the gulls, but not to the turnstones, in higher proportions than did black skimmers. Two decoys, a mink and a gull, were used to simulate predatory encounters, and a duck decoy was used as a control at two mixed-species and one single-species subcolonies. Gull-billed terns responded in significantly higher proportions than did skimmers to all decoy treatments in the mixed-species subcolonies. Mobbing of both natural and model predators by the terns suggests that skimmers may gain a reproductive advantage by nesting with these terns. However, the response of black skimmers to both natural and simulated predators was independent of the presence of gull-billed terns in the colony, indicating that black skimmers may not perceive these objects as threats, or may react differently to predators than do gull-billed terns.     

MORPHOLOGICAL AND ULTRASTRUCTURAL FEATURES OF A STRAIN OF BOTRYOCOCCUS TERRIBILIS (TREBOUXIOPHYCEAE) FROM BRAZIL1

The genus Botryococcus comprises a group of cosmopolitan species of freshwater colonial green algae, some of which synthesize and accumulate an unusually high level (15–76%) of liquid hydrocarbons. This characteristic suggests the possibility of exploiting species from this group as renewable sources for jet fuel. An oil‐rich strain of Botryococcus (Trebouxiophyceae) was isolated from a freshwater pond in the state of Bahia, Brazil, and is presently maintained under standard conditions at the Culture Collection of the Institute of Biology, Federal University of Bahia. The taxonomic classification of the species was based on light microscopy (LM); and TEM and SEM were used to better characterize its features, which have never before been described at this level. The LM characterization included the size of the colonies (35.7–157 μm) and cells (8–10 × 5–9 μm) and their connection in sub‐colonies by mucilaginous strands, as well as the presence of mucilaginous processes on the periphery of some of the colonies, with most of the cells included inside the colony. Reproduction occurred through divisions into two to four autospores. These features characterized the species as Botryococcus terribilis Komárek and Marvan. The TEM study showed, in addition to the presence of starch grains, pyrenoids that are penetrated by thick thylakoids. The pyrenoid bodies appear as electron‐dense protein inclusions located in the chloroplast and surrounded by a starch sheath. These structures, which contain most if not all of the Ribulose‐1,5‐bisphosphate carboxylase oxygenase in several algal species that have been studied closely, are newly discovered for this species.     

Extensive gene flow along the urban–rural gradient in a migratory colonial bird

Urban colonization by wildlife involves a combination of several different mechanisms, including phenotype or genotype sorting, phenotypic plasticity and microevolutionary adaptation. Combination of these processes can produce a rapid phenotypic, but also genetic divergence of urban versus rural populations. Here, we examined the pattern of genetic differentiation between urban and rural populations of a colonial migratory bird, the black‐headed gull Chroicocephalus ridibundus. To this end, we sampled ca 170 individuals from six (two urban and four rural) colonies in northern Poland, and genotyped them at ten microsatellite loci. Our analysis provided evidence for negligible genetic divergence of urban and rural colonies, as assessed with fixation index F_ST and Nei's unbiased genetic distance D (mean pairwise urban‐rural comparisons: F_ST = 0.003 ± 0.001 [SE] and D = 0.012 ± 0.006 [SE]). Bayesian clustering methods provided support for homogeneous genetic structure across all urban and rural populations. Also, we found no support for reduced allelic diversity in urban versus rural colonies. These results stand in a stark contrast to the previous findings on the genetic consequences of urbanization in birds. We hypothesize that this pattern could possibly be attributed to the important life‐history characters of the black‐headed gull, including coloniality, migratoriness, and high dispersal propensity. Our study provides a novel insight into the urban landscape genetics, underlining large variation in the mechanisms of urban colonization and its genetic consequences in wild animal populations.     

Can the presence of curved forms of the diatom Aulacoseira ambigua in the Nile (Egypt) and Vaal (South Africa) Rivers be ascribed to similar water quality conditions?

Spiral colonies of the diatom Aulacoseira ambigua were assigned the rank of forma (Aulacoseira ambigua f. japonica). This spiral-shaped, colonial, centric diatom has limited geographical distribution and is currently reported to occur in only a few countries in the world. In Africa this species was described for the first time from the Vaal River, South Africa, but recent research on the Nile River also revealed its presence in Egypt. Physical and chemical data on water quality in these two river systems were compared to determine whether the presence of this uncommon ‘phenoecodeme’ could be ascribed to similar environmental conditions. Results indicate that the rivers are dissimilar with regard to many variables, but both rivers provide turbid, warm and eutrophic waters with medium to high mineral content and it was concluded that these factors favour and sustain the growth of this curved form.     

The evolution of coloniality: the emergence of new perspectives

The evolution of group living remains an outstanding question in evolutionary ecology. Among the most striking forms of group living are the enormous assemblages of breeders that occur in many colonial marine birds and mammals, with some colonies containing more than a million individuals breeding in close contact. Coloniality is an evolutionary puzzle because individuals pay fitness costs to breed in high densities. Despite numerous potential benefits proposed to overcome these costs, we still lack a general framework to explain coloniality. Several new hypotheses involving breeding habitat and mate selection create promising approaches for studying this enigma.     

Mating strategy and isolation between the two forms, macrogyna and microgyna, of Myrmica ruginodis (Hym. Formicidae)

Abstract.

• 1 It is shown that the size distribution and degree of overlap between individual queens of the two forms of Myrmica ruginodis, microgyna and macrogyna, is exactly the same in southern England as was originally described from Scotland.
• 2 Distinguishing colonies of the two forms is not as easy as distinguishing between individuals, both types can be polygynous and approximately 25% of colonies contain a mixture of queens.
• 3 Males of the two forms can be distinguished solely on the basis of size and, in general, macrogyna colonies produce larger males, workers and gynes than microgyna colonies. There is little correlation between the sizes of males, workers and gynes in colonies within the macrogyna and microgyna groups.
• 4 If the two forms are separate species and mixing is a parasitic association (as suggested elsewhere), then there should be breeding isolation between the forms. This is tested by examining data from nine mating-swarms.
• 5 All the swarms contain a significant proportion of the microgyna form. There is no evidence of assortative mating, although larger males are more likely to get a mate than smaller ones. This behaviour, combined with the possibility that microgynes mate near to the nest, might prevent complete mixing during mating.
• 6 The status of the forms is discussed. Besides the possibility of a‘pre-parasitic’relationship it is suggested that the forms might represent a polymorphism, present in all populations, the balance between them being the result of selection determined by local environmental factors.