First report of Cryptosporidium hominis in a freshwater sponge.

Identification of Cryptosporidium oocyst is essential in ensuring water quality fit for human use, consumption, and recreation. This communication proposes the supplemental analysis of substrate-associated biofilms, in particular, freshwater sponges in improving case finding of waterborne-protozoan pathogens (WBPP) in environmental aquatic samples. In this study, a small portion of a mature freshwater sponge under the Genus Spongilla was subjected to microscopic and molecular analysis to identify the presence of Cryptosporidium. Microscopic screening with modified Kinyoun's staining (MK) and microscopic confirmation using direct antibody fluorescent testing (IFT) returned with Cryptosporidium spp. positive findings. Molecular investigation resulted in the confirmation of Cryptosporidium hominis upon sequencing of PCR products and phylogenetic analysis. This is the first report of a pathogenic protozoan, C. hominis isolated from a freshwater sponge. The results of this study provide evidence of the value of expanding water quality assessment strategies to the analysis of substrate-associated biofilms and sponges in improving case finding of WBPP in natural aquatic environments.

Early sponges and toxic protists: possible sources of cryostane, an age diagnostic biomarker antedating Sturtian Snowball Earth.

The period 800-717 million years (Ma) ago, in the lead-up to the Sturtian Snowball glaciation, saw an increase in the diversity of eukaryotic microfossils. To afford an independent and complementary view of this evolutionary period, this study presents the distribution of eukaryotic biomarkers from three pre-Sturtian successions across the supercontinent Rodinia: the ca. 780 Ma Kanpa Formation of the Western Australian Officer Basin, the ca. 800-740 Ma Visingsö Group of Sweden, and the 740 Ma Chuar Group in Arizona, USA. The distribution of eukaryotic steranes is remarkably similar in the three successions but distinct from all other known younger and older sterane assemblages. Cholestane was the only conventional structure, while indigenous steranes alkylated in position C-24, such as ergostane, stigmastane, dinosterane and isopropylcholestane, and n-propylcholestane, were not observed. This sterane distribution appears to be age diagnostic for the pre-Sturtian Neoproterozoic. It attests to the distinct evolutionary state of pre-Snowball eukaryotes, pointing to a taxonomic disparity that was still lower than in the Ediacaran (635-541 Ma). All three basins also show the presence of a new C28 sterane that was tentatively identified as 26-methylcholestane, here named cryostane. The only known extant organisms that can methylate sterols in the 26-position are demosponges. This assignment is plausible as molecular clocks place the appearance of the earliest animals into the pre-Sturtian Neoproterozoic. The unusual 26-methylsterol may have protected sponges, but also other eukaryotes, against their own membranolytic toxins. Some protists release lytic toxins to deter predators and kill eukaryotic prey. As conventional membrane sterols can be the site of attack for these toxins, sterols with unusual side-chain modification protect the cell. This interpretation of cryostane supports fossil evidence of predation in the Chuar Group and promotes hypotheses about the proliferation of eukaryophagy in the lead-up to the Cryogenian.

A new species of sponge-dwelling Oukuriella (Chironomidae) from Brazil.

Oukuriella Epler, 1986 is an exclusively Neotropical genus that comprises two well supported clades: inhabitants of freshwater sponges and inhabitants of immersed wood (saproxylophilic). In this work, males of a new chironomid species, Oukuriella froehlichi sp. n., are described and illustrated. Specimens were collected near stream in the Brazilian Atlantic Forest. Phylogenetic analyses performed in this study show that the newly discovered species belongs in the clade of freshwater sponge inhabitants.

Comparisons of the fungal and protistan communities among different marine sponge holobionts by pyrosequencing.

To date, the knowledge of eukaryotic communities associated with sponges remains limited compared with prokaryotic communities. In a manner similar to prokaryotes, it could be hypothesized that sponge holobionts have phylogenetically diverse eukaryotic symbionts, and the eukaryotic community structures in different sponge holobionts were probably different. In order to test this hypothesis, the communities of eukaryota associated with 11 species of South China Sea sponges were compared with the V4 region of 18S ribosomal ribonucleic acid gene using 454 pyrosequencing. Consequently, 135 and 721 unique operational taxonomic units (OTUs) of fungi and protists were obtained at 97 % sequence similarity, respectively. These sequences were assigned to 2 phyla of fungi (Ascomycota and Basidiomycota) and 9 phyla of protists including 5 algal phyla (Chlorophyta, Haptophyta, Streptophyta, Rhodophyta, and Stramenopiles) and 4 protozoal phyla (Alveolata, Cercozoa, Haplosporidia, and Radiolaria) including 47 orders (12 fungi, 35 protists). Entorrhizales of fungi and 18 orders of protists were detected in marine sponges for the first time. Particularly, Tilletiales of fungi and Chlorocystidales of protists were detected for the first time in marine habitats. Though Ascomycota, Alveolata, and Radiolaria were detected in all the 11 sponge species, sponge holobionts have different fungi and protistan communities according to OTU comparison and principal component analysis at the order level. This study provided the first insights into the fungal and protistan communities associated with different marine sponge holobionts using pyrosequencing, thus further extending the knowledge on sponge-associated eukaryotic diversity.

Review of Xenochironomus Kieffer, 1921 (Diptera: Chironomidae) with description of six new species.

The genus Xenochironomus Kieffer is reviewed. Five new species are described from Brazil, X. alaidae, X. amazonensis and X. martini as male; X. grini and X. mendesi as male, pupa and larva, and X. ethiopensis from Ethiopia is described as adult male. X. canterburyensis (Freeman) is redescribed as male, pupa and larva; X. trochanteratus (Thompson), X. trisetosus (Kieffer), X. ugandae (Goetghebuer) and X. tuberosus Wang as males; notes on X. xenolabis (Kieffer), X. flaviventris (Kieffer), X. longicrus (Kieffer), X. lacertus Dutta et Chaudhuri and X. ceciliae Roque et Trivinho-Strixino are also given. The species X. nigricaudus Hashimoto is recognized as nomen dubium. Keys to males, pupae and larvae are presented.

Zyzzyzus rubusidaeus (Cnidaria, Hydrozoa, Tubulariidae), a new species of anthoathecate hydroid from the coast of British Columbia, Canada.

Zyzzyzus rubusidaeus, sp. nov., is described from inshore waters near the northern tip of Vancouver Island, British Columbia, Canada. Specimens were collected on rocky bottoms amongst barnacles, sponges, and compound ascidians at a depth of 18 m in Weynton Passage, Broughton Strait, during March, July, and October 2012. Polyps tend to grow in dense aggregations, often covering several square centimetres. Hydroids of Z. rubusidaeus most closely resemble those of Z. robustus Petersen, 1990 from Greenland, but differ in having aboral tentacles that are scattered in a narrow band around the base of the hydranth rather than occurring in a single whorl, thin and transparent instead of thick and stiff perisarc around hydrocaulus and tubers, and gonophores that arise from simple pedicels instead of short, stout branches. Possible embryos were present in female gonophores, although structures recognizable as actinula larvae have not been observed. The cnidome comprises small and large stenoteles, desmonemes, microbasic euryteles, basitrichs, and isorhizas. Polyps are a raspberry colour in life, a hue that has faded but little in our formalin-preserved material. Discovery of this hydroid brings the number of species currently recognized in genus Zyzzyzus Stechow, 1921 to seven.

These squatters are not innocent: the evidence of parasitism in sponge-inhabiting shrimps.

Marine sponges are frequently inhabited by a wide range of associated invertebrates, including caridean shrimps. Symbiotic shrimps are often considered to be commensals; however, in most cases, the relationship with sponge hosts remains unclear. Here we demonstrate that sponge-inhabiting shrimps are often parasites adapted to consumption of sponge tissues. First, we provide detailed examination of morphology and stomach contents of Typton carneus (Decapoda: Palaemonidae: Pontoniinae), a West Atlantic tropical shrimp living in fire sponges of the genus Tedania. Remarkable shear-like claws of T. carneus show evidence of intensive shearing, likely the result of crushing siliceous sponge spicules. Examination of stomach contents revealed that the host sponge tissue is a major source of food for T. carneus. A parasitic mode of life is also reflected in adaptations of mouth appendages, in the reproduction strategy, and in apparent sequestration of host pigments by shrimp. Consistent results were obtained also for congeneric species T. distinctus (Western Atlantic) and T. spongicola (Mediterranean). The distribution of shrimps among sponge hosts (mostly solitary individuals or heterosexual pairs) suggests that Typton shrimps actively prevent colonisation of their sponge by additional conspecifics, thus protecting their resource and reducing the damage to the hosts. We also demonstrate feeding on host tissues by sponge-associated shrimps of the genera Onycocaris, Periclimenaeus, and Thaumastocaris (Pontoniinae) and Synalpheus (Alpheidae). The parasitic mode of life appears to be widely distributed among sponge-inhabiting shrimps. However, it is possible that under some circumstances, the shrimps provide a service to the host sponge by preventing a penetration by potentially more damaging associated animals. The overall nature of interspecific shrimp-sponge relationships thus warrants further investigation.

First fossil record of parasitism in Devonian calcareous sponges (stromatoporoids).

INTRODUCTION: Palaeozoic calcareous sponges (stromatoporoids) are common bio-constructing fossils; they are sometimes found in association with helicoidal structures of unknown biological affinities. The interaction between the tube-forming organisms has usually been classified as commensalism. METHODS: About 260 stromatoporoid skeletons from the Middle Devonian (Givetian) of the Mont d'Haurs section near Givet (Champagne-Ardenne, France) were thin-sectioned and analysed under transmitted light. RESULTS: Approximately 10% of the examined stromatoporoids (mainly belonging to the genera Actinostroma, Stromatopora and Stromatoporella) contain tubes classified as Torquaysalpinx sp. The Torquaysalpinx organisms penetrated the skeletons of stromatoporoids in vivo (as evidenced by skeletal overgrowths); around the infesting organisms, growth bands are bent down. CONCLUSION: Diminished growth rates around the infesting organism demonstrate a negative influence on the host, similar to that seen in the modern demosponge-polychaete association of Verongia-Haplosyllis. This is demonstrated by changes in growth bands. As in the above-mentioned association, the endosymbiont might have been feeding directly upon the tissues of the host. The Torquaysalpinx organisms were gaining habitat and possibly also food resources - for them this interaction was clearly positive. This long-term association can therefore be classified as parasitism. This is the first evidence for parasitism in Palaeozoic sponges.

Exploring the role of microorganisms in the disease-like syndrome affecting the sponge Ianthella basta.

A disease-like syndrome is currently affecting a large percentage of the Ianthella basta populations from the Great Barrier Reef and central Torres Strait. Symptoms of the syndrome include discolored, necrotic spots leading to tissue degradation, exposure of the skeletal fibers, and disruption of the choanocyte chambers. To ascertain the role of microbes in the disease process, a comprehensive comparison of bacteria, viruses, fungi, and other eukaryotes was performed in healthy and diseased sponges using multiple techniques. A low diversity of microbes was observed in both healthy and diseased sponge communities, with all sponges dominated by an Alphaproteobacteria, a Gammaproteobacteria, and a group I crenarchaeota. Bacterial cultivation, community analysis by denaturing gradient gel electrophoresis (Bacteria and Eukarya), sequencing of 16S rRNA clone libraries (Bacteria and Archaea), and direct visual assessment by electron microscopy failed to reveal any putative pathogens. In addition, infection assays could not establish the syndrome in healthy sponges even after direct physical contact with affected tissue. These results suggest that microbes are not responsible for the formation of brown spot lesions and necrosis in I. basta.

The symbiotic lifestyle and its evolutionary consequences: social monogamy and sex allocation in the hermaphroditic shrimp Lysmata pederseni.

Sex allocation theory predicts female-biased sex allocation for simultaneous hermaphrodites with a monogamous mating system. Mating systems theory predicts that monogamy is advantageous in environments where refuges are discrete, scarce, relatively small, and when predation risk is high outside of these refuges. These predictions were tested with the Caribbean shrimp Lysmata pederseni, a simultaneous hermaphrodite which has an early male phase and lives inside tubes of the sponge Callyspongia vaginalis. This host sponge is a scarce resource that, together with the high predation risk typical of tropical environments, should favor monogamy in the shrimp. Field observations demonstrated that shrimps were frequently encountered as pairs within these tube sponges. Pairs were equally likely to comprise two hermaphrodites or one hermaphrodite and one male. Several of these pairs were observed for long periods of time in the field. Experiments demonstrated that hermaphrodites tolerated other hermaphrodites but not males in their host sponge. These results suggest that pairs of hermaphroditic L. pederseni are socially monogamous; they share the same host individual and might reproduce exclusively with their host partners for long periods of time. Nevertheless, males appeared less likely to establish long-term associations with hermaphrodites as indicated by the rate of their disappearance from their hosts (greater than that of hermaphrodites). Sex allocation was female biased in monogamous hermaphrodites. On average, hermaphrodites invested 34 times more to female than to male reproductive structures. Monogamy and female-biased sex allocation seem to be evolutionary consequences of adopting a symbiotic lifestyle in simultaneous hermaphrodites.

Kin structure, ecology and the evolution of social organization in shrimp: a comparative analysis.

Eusocial societies present a Darwinian paradox, yet they have evolved independently in insects, mole-rats and symbiotic shrimp. Historically, eusociality has been thought to arise as a response to ecological challenges, mediated by kin selection, but the role of kin selection has recently been questioned. Here we use phylogenetically independent contrasts to test the association of eusociality with ecological performance and genetic structure (via life history) among 20 species of sponge-dwelling shrimp (Synalpheus) in Belize. Consistent with hypotheses that cooperative groups enjoy an advantage in challenging habitats, we show that eusocial species are more abundant, occupy more sponges and have broader host ranges than non-social sister species, and that these patterns are robust to correction for the generally smaller body sizes of eusocial species. In contrast, body size explains less or no variation after accounting for sociality. Despite strong ecological pressures on most sponge-dwellers, however, eusociality arose only in species with non-dispersing larvae, which form family groups subject to kin selection. Thus, superior ability to hold valuable resources may favour eusociality in shrimp but close genetic relatedness is nevertheless key to its origin, as in other eusocial animals.

Symbiotic interaction between dinoflagellates and the demosponge Lubomirskia baicalensis: aquaporin-mediated glycerol transport.

Lake Baikal is rich in endemic sponge species, among them the arborescently growing species Lubomirskia baicalensis. During winter when the lake is covered by ice, this species reproduces sexually, reflecting a high metabolic activity. Throughout the year, L. baicalensis lives in association with dinoflagellates, which - according to the data presented herein - are symbiotic. The dinoflagellates have been determined on the basis of their rDNA/ITS characteristics and were found to display high sequence similarity to Gymnodinium sanguineum. The dinoflagellates give the sponge its characteristic green color, reflecting the high chlorophyll content (chlorophyll-a content in March and September of 3.2 +/- 0.6 microg/g and 1.9 +/- 0.5 microg/g of protein, respectively). With the in vitro cell culture system for sponges, the primmorphs, it could be demonstrated that [(14)C] glycerol is readily taken up by sponge cells; this process can be inhibited by phloretin, an aquaporin channel blocker. In order to prove the effect of cholesterol on the intermediate metabolism of the sponge cells, molecule probes, cDNAs for key enzymes in gluconeogenesis, glycolysis, and citric acid, have been applied in Northern blot studies. The data revealed that the genes coding for the enzymes citrate synthase and fructose-1,6-bisphosphatase are strongly upregulated after exposure of primmorphs to glycerol. This effect is abolished by phloretin. The genes encoding the phosphoglucose isomerase and pyruvate dehydrogenase do not respond to glycerol supply, suggesting that their expression is not under genetic control in L. baicalensis. To prove the assumption that the aquaporin channel is involved in the influx of glycerol in sponge cells, this cDNA was cloned and applied for in situ hybridization studies. The results obtained show that cells surrounding the dinoflagellates become brightly stained after hybridization with the aquaporin this probe. This demonstrates that L. baicalensis cells respond to glycerol, a metabolite which might be supplied by the dinoflagellates and imported via the aquaporin channel into the sponge cells.

Sponge-dwelling chironomids in the upper Paraná river (Brazil): little known but potentially threatened species.

In this paper, we provide a checklist of chironomids living in freshwater sponges in the upper Paraná River (Brazil) and we call attention to the problem of complete substitution of extensive areas covered by freshwater sponges by banks of the invasive alien species Limnoperna fortunei (golden mussel). We describe a new species of Oukuriella based on male adults and also describe an unusual larva of Chironomidae that probably belongs to the Stenochironomus complex.

Sponge-dwelling chironomids in the upper Paraná river (Brazil): little known but potentially threatened species / Chironomidae habitantes de esponjas de água doce no alto Rio Paraná (Brasil): espécies pouco conhecidas, mas potencialmente ameaçadas

Neste estudo é apresentada uma lista de espécies de Chironomidae habitantes de esponjas de água doce coletados no Alto Rio Paraná e é chamada a atenção para o problema de substituição de grandes áreas cobertas por esponjas de água doce por bancos da espécie invasora Limnoperma fortunei (mexilhão dourado). Também é descrita uma nova espécie de Oukuriella com base em adulto macho e feita uma descrição informal de uma larva de Chironomidae que provavelmente pertence ao complexo Stenochironomus.

In this paper, we provide a checklist of chironomids living in freshwater sponges in the upper Paraná River (Brazil) and we call attention to the problem of complete substitution of extensive areas covered by freshwater sponges by banks of the invasive alien species Limnoperna fortunei (golden mussel). We describe a new species of Oukuriella based on male adults and also describe an unusual larva of Chironomidae that probably belongs to the Stenochironomus complex.

Defense of benthic invertebrates against surface colonization by larvae: a chemical arms race.

Sessile invertebrates evolved in a competitive milieu where space is a limiting resource, setting off an arms race between adults that must maintain clean surfaces and larvae that must locate and attach to a suitable substratum. I review the evidence that invertebrates chemically deter or kill the propagules of fouling animals and protists under natural conditions, and that chemosensory mechanisms may allow larvae to detect and avoid settling on chemically protected organisms. The fouling process is an ecologically complex web of interactions between basibionts, surface-colonizing microbes, and fouling larvae, all mediated by chemical signaling. Host-specific bacterial communities are maintained by many invertebrates, and may inhibit fouling by chemical deterrence of larvae, or by preventing biofilm formation by inductive strains. Larval settlement naturally occurs in a turbulent environment, yet the effects of waterborne versus surface-adsorbed chemical defenses have not been compared in flow, limiting our understanding of how larvae respond to toxic surfaces in the field. The importance of evaluating alternative hypotheses such as mechanical and physical defense is discussed, as is the need for ecologically relevant bioassays that quantify effects on larval behavior and identify compounds likely to play a defensive role in situ.

[Some peculiarities of the relationships between parasitic copepods and their invertebrate hosts]. / Nekotorye osobennosti vzaimootnoshenii paraziticheskykh veslonogikh i ikh bespozvonochnykh-khoziaev.

According to the rule of academican E. N. Pavlovskiy, any organism of host is an environment of inhabit for a parasite (Pavlovskiy, 1934). It was analysed, which "ecological niche" or microbiotop (= microhabitat) is occupied by this or that species of symbiotic (parasitic) copepods in organisms of different groups invertebrate-hosts. The assumption lying in a basis of the given analysis means that each group of hosts may give to cohabitants only certain variants of microbiotopes independently on the general morphological structure and life mode of hosts. Five types of microbiotops offered by various groups of hosts for symbiotic copepods are designated (Ta[symbol: see text] 2). 1. The body surface of benthic invertebrates as a microbiotope is characterized by conditions being little different (concerning any kind of physical and chemical influences on copepods) from those in external environment on any other substrate. Apparently a trophical dependence plays a determining role in this case. There are certain directions in a development of adaptations, which are characteristic in some extent for all water ectoparasitic crustaceans and have one functional task--to help to an ectoparasite to keep itself on a surface of host body. In the first, the maxillules and maxillipeds significantly are developed, they get a form of large claws, with which the dopepods are strongly attached on a surface of host body and have an opportunity to move on it without a danger to be washed off. In the second, the form of the body undergoes a dorso-ventral expression and expansion of prosome, forms a cephalic shield allowing to the symbiont to press itself tightly to the host body surface and to avoid the loss of host (tab. 2). In occasions, some ectoparasites stimulate the formation of galls in skin tissues of the host, that also provides the parasite with constant conditions, without any threat to lose the host. However, this phenomenon has not a wide distribution and is observed in some groups of crustacean and echinoderm hosts. 2. The narrow tubular cavities in the organism of host either they are a part of external environment (as in channel system of spongia) or a part of internal environment of organism (as channels of blood system or thin parts of a digestive system) have always rigidly limited sizes and form. Characteristics of all parasites occupying this microbiotopes are the strong transformations. They are expressed by the reduction of legs or any other appendages (frequently in a significant degree), loss of segmentation to some extent and in eruciform (or vermiform) form of a body (tab. 2). This microbiotope is occupied by an ectoparasite in one case only (Spongicola uncifer from channel system of spongia) and by endoparasites in all other cases. 3. Large cavities connected with external environment. The formations of various genesis, such as mantle cavity of molluscs, gill cavity and marsupium of crustaceans, bursal cavity of ophiuroids and branchial cavity of ascidians, concern this type of microbiotopes. All of them are characterized by the relative difference from the external environment and rather large volume (in comparison to sizes of copepods), that provides the parasites with a sufficient protection from factors of the external environment and constant source of food such as elements of host body or food's particles brought by the water flow. Morphological changes in inhabitants of the microbiotope have two directions. They practically are absent in the overwhelming majority copepods, living in the mantle of cavity of the lamellibranches. On the other hand, the inhabitants of gill cavity and marsupium of crustaceans, bursal cavities of ophiuroids and branchial cavity of ascidians are characterized by the presence of strong transformations. Usually there are expressed in a loss of segmentation to some extent, reduction of appendages and swelling of body, as in species of the genus Sphaeronella (tab. 2). Changes are also observed in the life cycle: the tendency to reduce stages of development (development of nauplii stage, which takes place under the ovarial cover). In this case the copepodid stages hatch from the ova. 4. The internal cavity of organism of host. This type of microbiotopes in different groups of the hosts is represented in a various degree. We recognise it in a coelome of polychaetes, lacunar system of molluscs, mixocoel of crustaceans, coelome of echinoderms and cavity of body in ascidians. Two basic evolutionary directions are observed in copepods occupying this microbiotope. In the first case, the parasite is not exposed to transformations and keeps the initial plan of structure as in ancestral free-living forms. In the second case the parasites are exposed to strong transformations, they either live directly in cavity's liquid, or are surrounded by a cyst (as in Cucumaricolidae). 5. Microbiotope of the last type is most specific. The simultaneous existence in two environments--external environment (environment of the second order) and internal environment (environment of the first order) leads to the complete loss of ancestral type in a structure and level of organisation. At the same time both morphological and functional division of the parasite body into two parts produces a new formation--the ectosome and endosome. In this case we deals with the phenomenon of mesoparasitism.

Multiple origins of eusociality among sponge-dwelling shrimps (Synalpheus).

As the most extreme expression of apparent altruism in nature, eusociality has long posed a central paradox for behavioral and evolutionary ecology. Because eusociality has arisen rarely among animals, understanding the selective pressures important in early stages of its evolution remains elusive. Employing a historical approach to this problem, we used morphology and DNA sequences to reconstruct the phylogeny of 13 species of sponge-dwelling shrimps (Synalpheus) with colony organization ranging from asocial pair-bonding through eusociality. We then used phylogenetically independent contrasts to test whether sociality was associated with evidence of enhanced competitive ability, as suggested by hypotheses invoking an advantage of cooperation in crowded habitats. The molecular, morphological, and combined data each strongly supported three independent origins of monogynous, multigenerational (eusocial) colony organization within this genus. Phylogenetically independent contrasts confirmed that highly social taxa, with strong reproductive skew, have significantly higher relative abundance within the host sponge than do less social taxa, a result that was robust to uncertainty in tree topology and varying models of character change. A similar tendency for highly social species to share their sponge with fewer congener species was suggestive, but not significant. Because unoccupied habitat appears to be limiting for many sponge-dwelling shrimp species, these data are consistent with hypotheses that cooperative social groups enjoy a competitive advantage over less organized groups or individuals, where independent establishment is difficult, and that enemy pressure is of central importance in the evolution of animal sociality.

Parasitic diatoms inside antarctic sponges.

Antarctic sponges may host large populations of planktonic and benthic diatoms. After settling on the sponge, these diatoms enter its body through pinacocytes (1) and form, there, large mono- or pauci-specific assemblages. Yet the total amount of carbohydrates in the invaded sponge tissue is inversely correlated with that of chlorophyll-a. We suggest, therefore, that endobiont diatoms utilize the products of the metabolism of their host as an energy source. This is the first evidence indicating that an endobiotic autotrophic organism may parasitize its animal host. Moreover, this unusual symbiotic behavior could be a successful strategy that allows the diatom to survive in darkness.