Besprechungen

Book reviewed in this article: Blackford Cook , Susan (1969): Experiments on homing in the Limpet Siphonaria normalis (Versuche zum Heimfinden der Napfschnecke S. n.). Varnall , John L. (1969): Aspects of the Behaviour of Octopus cyanea Gray (Zum Verhalten des Tintenfischs O. c.). Szebenyi , Andrew L., S. J. (1969): Cleaning behaviour in Drosophila melanogaster (Putzverhalten bei der Taufliege). Bennet-Clark , H. C., und A. W. Ewing (1969): Pulse interval as a critical parameter in the courtship song of Drosophila melanogaster (Pausenlänge als entscheidendes Merkmal des Balzgesangs der Taufliege). Goodyear , C. Phillip, und Denzel E. Ferguson (1969): Sun compass orientation in the Mosquito Fish, Gambusia affinis (Sonnenkompaßorientierung beim Moskitofisch). Littlejohn , M. J., und A. A. Martin (1969): Acoustic interaction between two species of Leptodactylid Frogs (Rufwettstreit zwischen zwei dünnfingrigen Froscharten). Alcock , John (1969): Observational learning by fork-tailed Flycatchers (Muscivora tyrannus) (Gabelschwänzige Weißbauchtyrannen lernen vom Zusehen). Thompson , William L. (1969): Song recognition by territorial male Buntings (Passerina) (Erkennen des Artgesanges bei Farbfinken im Revier). Sielmann , H. (1968): 1. Scenopoeëtes dentirostris (Ptilonorhynchidae), Verhalten am Blätterteppich (E 1075). - 2. Ptilonorhynchus violaceus (Ptilonorhynchidae); Bauen an der Laube und Anstreichen' (E 1076), Balz und Kopulation (E 1077). - 3. Prionodura newtoniana (Ptilonorhynchidae), Behängen der Reisigtürme (E 1082). - 4. Amblyornis macgregoriae (Ptilonorhynchidae), Behängen des, Mai-baumes' und Balz (E 1083). - 5. A. subalaris (Ptilonorhynchidae), Schmücken der Laube (E 1084). - 6. Chlamydera lauterbachi (Ptilonorhynchidae); Bauen an der Laube (E 1080), Balz (E 1081). - 7. C. nuchalis (Ptilonorhynchidae); Verteilen von Sammlungsstücken an der Laube (E 1078), Balz und Kopulation (E 1079). Rosengren , Anita (1969): Experiments in colour discrimination in dogs (Versuche über Farbensehen von Hunden). Pickenhain , L., und F. Klingberg (1969): Hirnmechanismen und Verhalten: Elektro-physiologische und Verhaltensuntersuchungen an der Ratte. Giersberg , H., und P. Rietschel (1968): Vergleichende Anatomie der Wirbeltiere, Band 2: Ernährungs-, Atmungs-, Kreislauforgane, Leibeshöhlen, Ausscheidungsund Fortpflanzungsorgane. Lattin , Gustaf de (1967) ?: Grundriß der Zoogeographie. McKusick , Victor A. (1968): Humangenetik. Jaynes , Julian (1969): Historical Origins of “Ethology” and “Comparative Psychology” (Zur Geschichte der Begriffe Ethologie und Vergleichende Psychologie). Kalmus , H. (1969): Animal Behaviour and Theories of Games and of Language (Tierverhalten, Spiel- und Sprachtheorien). Eibl-Eibesfeldt , I. (1970): Liebe und Hass, zur Naturgeschichte elementarer Verhaltensweisen. Wickler , Wolfgang (1970): Stammesgeschichte und Ritualisierung. Kolar , Kurt (1965): (1) Kontinent voll Kuriositäten, Tiere, Menschen und Probleme in Australien. (2) Ders. (1967): Von Hund und Äff und Kind und Katz. (3) Ders. (1970): Australien neu entdeckt, ein Verhaltensforscher im fünften Kontinent. Wylie , Philip (1968): The Magic Animal (Das magische Tier). Laver , J. (1970): Die Mode.     

Bower-building: coevolution of display traits in response to the costs of female choice?

Bower-building is peculiar to polygynous species in the family Ptilonorhynchidae (Bowerbirds). Here we summarize past and current theories regarding the evolution of bower-building, evaluate the evidence that can be used to differentiate between them and suggest future areas for research.     

Prevalence and differential host-specificity of two avian blood parasite genera in the Australo-Papuan region

The degree to which widespread avian blood parasites in the genera Plasmodium and Haemoproteus pose a threat to novel hosts depends in part on the degree to which they are constrained to a particular host or host family. We examined the host distribution and host-specificity of these parasites in birds from two relatively understudied and isolated locations: Australia and Papua New Guinea. Using polymerase chain reaction (PCR), we detected infection in 69 of 105 species, representing 44% of individuals surveyed (n = 428). Across host families, prevalence of Haemoproteus ranged from 13% (Acanthizidae) to 56% (Petroicidae) while prevalence of Plasmodium ranged from 3% (Petroicidae) to 47% (Ptilonorhynchidae). We recovered 78 unique mitochondrial lineages from 155 sequences. Related lineages of Haemoproteus were more likely to derive from the same host family than predicted by chance at shallow (average LogDet genetic distance = 0, n = 12, P = 0.001) and greater depths (average distance = 0.014, n = 11, P < 0.001) within the parasite phylogeny. Within two major Haemoproteus subclades identified in a maximum likelihood phylogeny, host-specificity was evident up to parasite genetic distances of 0.029 and 0.007 based on logistic regression. We found no significant host relationship among lineages of Plasmodium by any method of analysis. These results support previous evidence of strong host-family specificity in Haemoproteus and suggest that lineages of Plasmodium are more likely to form evolutionarily-stable associations with novel hosts.     

A ten-year comparative study of the population dynamics and parasitoidism in the native paper wasp Polistes fuscatus and the invasive P. dominulus

The invasive success of Polistes dominulus in North America has been attributed to its greater productivity relative to native Polistes. Liberation from parasites and parasitoids are thought to be major factors contributing to the high productivity of P. dominulus. We analyzed historical records of colony relative abundance and productivity of P. dominulus and the sympatric, native Polistes fuscatus from 1995 to 2010 using historical data from our Michigan Polistes study site. We also analyzed evidence of parasitoids from 294 P. fuscatus and 507 P. dominulus archived combs from 2001 to 2010. Additionally, we examined field and laboratory colonies from outside of our study site for parasites and parasitoids in 2009 and 2010. We documented one parasite and three parasitoids exploiting Polistes in our Michigan study sites. Our historical records document that P. dominulus initially displaced P. fuscatus rapidly, then slowed, and finally the two populations stabilized. Furthermore, the historical pattern of decreasing displacement of P. fuscatus by P. dominulus corresponded temporally with a significant decline in the productivity and a significant increase in Dibrachys cavus infestation of P. dominulus. Our evidence indicates that the parasitoid, D. cavus, is a major factor in stabilizing the populations of the sympatric P. dominulus and P. fuscatus.     

Cloning of the gene encoding the catalytic subunit of casein kinase II from the yeast Yarrowia lipolytica

Casein kinase II from the yeast Yarrowia lipolytica is a heterotetramer of the form αα′β₂. We report on the cloning and sequencing of a partial cDNA and of the complete genomic DNA coding for the catalytic α subunit of the casein kinase II from this yeast species. The sequence of the gene coding for this enzyme has been analyzed. No intron was found in the gene, which is present in a single copy. The deduced amino acid sequence of the gene shows high similarity with those of α subunit described in other species, although, uniquely, Y. lipolytica CKIIα lacks cysteines. We find that the α subunit sequence of Y. lipolytica CKII is shown greater homology with the corresponding protein from S. pombe than with that from S. cerevisiae. We have analyzed CKIIα expression and CKIIα activity. We show that expression of this enzyme is regulated. The catalytic subunit is translated from a single mRNA, and the enzyme is present at a very low level in Y. lipolytica, as in other yeasts.     

Rooting Around the Eutherian Family Tree: the Origin and Relations of the Taeniodonta

Placing early groups into the overall phylogeny of eutherian mammals can be challenging, particularly when the group does not have extant members. We investigated the relationships of the Taeniodonta, an extinct group from the Late Cretaceous through Paleogene of North America. This group has a few purported close relatives, including Cimolestes, Procerberus, and Alveugena, that may form a sequence of ancestors and descendants. The leading hypothesis is that Procerberus gave rise to taeniodonts through Alveugena. We test this hypothesis and analyze relations to known stem and crown Eutheria to determine the place of taeniodonts in eutherian phylogeny. Cladistic analyses were performed using previously published characters and datasets, namely a taeniodont/cimolestid specific dataset and a reanalysis of Wible and colleagues (2009), with added taxa for both. Our studies suggest that taeniodonts arose from Cimolestes through Alveugena, that Procerberus is more distantly related to taeniodonts, and that taeniodonts and their relatives are stem eutherians. We diagnose the Taeniodonta based on these analyses. Other Paleogene groups, especially those allied with Cimolestes such as tillodonts and pantolestans, merit further study. Our findings indicate that stem eutherians such as the Taeniodonta, in addition to crown eutherians, continued to diversify during the Paleogene.     

Comparative proteomic analysis of the saliva of the Rhodnius prolixus,Triatoma lecticularia and Panstrongylus herreri triatomines reveals a high interespecific functional biodiversity

Triatomines are hematophagous arthropods that transmit Trypanosoma cruzi and Trypanosoma rangeli. Feeding behavior and pathogen transmission is known to vary between the different species, and this characteristic is directly or indirectly dependent on the bioactive molecules of the saliva that facilitate the vector–host–parasite interaction. Here, we identify, characterize and compare the sialoproteomic (from the Greek sialo: saliva) repertoire of important species of the main triatomine genera in the Americas (Rhodnius prolixus, Triatoma lecticularia and Panstrongylus herreri) to better explain this interaction through two-dimensional electrophoresis and mass spectrometry. We identified 221 proteins, 69 from R. prolixus, 100 from T. lecticularia and 52 from P. herreri. We identified high abundance molecules with a great potential to modulate host defenses and homeostasis, highlighting Nitrophorin-4 (28.7%), Salivary lipocalin-5 (65.2%) and Putative triabin (20.5%) in R. prolixus, T. lecticularia and P. herreri, respectively. We also observed that only a single hypothetical protein is shared among three species, which was not functionally categorized. This study corroborates previous findings with R. prolixus, increasing the knowledge about this species with relevant proteomic information and comparisons with the other two targets of the study, T. lecticularia and P. herreri, for which no studies are available from a proteomics perspective.     

Sex-specific differences and the role of predation in the interaction between the hermit crab, Pagurus longicarpus, and its epibiont, Hydractinia symbiolongicarpus

The interaction between the hermit crab, Pagurus longicarpus, and the shell epibiont, Hydractinia symbiolongicarpus, varies from mutualism to parasitism based on the environmental context. We tested the hypothesis that this interaction also varies as a function of hermit crab sex. Given that recent work showed a negative effect of Hydractinia on female reproduction, we predicted a greater frequency of males in Hydractinia shells in the field and a stronger preference by males than females for shells with Hydractinia. Field collections documented a significantly greater proportion of males than females occupying shells with Hydractinia, and a significantly greater proportion of ovigerous females than non-ovigerous females in shells with Hydractinia. In laboratory shell-switching experiments, a greater proportion of males than females chose to enter shells with Hydractinia, but there was no difference in the proportions of males and females that vacated shells with Hydractinia.We examined whether the presence of Hydractinia influenced predation rates. Blue crabs fed on more than twice as many hermit crabs in shells with Hydractinia as compared to bare shells, but there was no significant difference for stone crabs. Laboratory experiments showed that the force required to crush shells was significantly greater for shells without Hydractinia. Thus, the lower occupancy and preference exhibited by females than males for shells with Hydractinia appears to result both from the decreased reproduction shown in past studies and an increase in predation risk.     

Relationships of Australo-Papuan songbirds-protein evidence

Allelic frequencies determined by protein electrophoresis were used to assess relationships and familial groupings among the Australian-New Guinean songbirds. Hypotheses of relationships were generated from the data by UPGMA, distance-Wagner and cladistic analyses. The resulting protein-based topologies were compared with recent DNA-DNA hybridization studies which separate the major Australo-Papuan families from Eurasian-centred groups and cluster them in a single endemic radiation. Although both data sets are concordant in may areas, they differ significantly in their alignment of the Australo-Papuan Menuridae, Climacteridae, Ptilonorhynchidae, Maluridae, Eopsaltriidae and Orthonychidae. Possible explanations for these discrepancies are examined. Both data sets otherwise agree in identifying an endemic radiation of Australo-Papuan songbirds in which some 75% of regional species are clustered in two principal assemblages. One includes the honeyeaters Meliphagidae and their allies Epthianuridae, Acanthizidae, Pardalotidae and, arguably, Eopsaltriidae (Australo-Papuan robins) and Orthonychidae (logrunners). The other comprises corvoid birds (Corvoidea), including Pachycephalidae, Monarchidae, Cracticidae, Artamidae, Paradisaeidae, Corvidae, Oriolidae, Campephagidae and possibly Ptilonorhynchidae (bowerbirds) amongst others. Menuridae (lyrebirds), Climacteridae (treecreepers), and Maluridae (fairy wrens) are also members of the radiation but of less certain affinity. The remaining species are grouped in a number of families forming a third assemblage of apparent Eurasian origin, the parvorder Passerida. The concept of a Gondwanan origin for the Passeriformes is canvassed.     

UDP-GlcNAc:Glycoprotein N-acetylglucosamine-1-phosphotransferase mediates the initial step in the formation of the methylphosphomannosyl residues on the high mannose oligosaccharides of Dictyostelium discoideum glycoproteins

The Dictyostelium discoideum gene gpt1 encodes a protein XP_638036 with sequence similarity to the α/β subunits of mammalian UDP-GlcNAc:Glycoprotein N-acetylglucosamine-1-phosphotransferase. We now demonstrate that extracts of D. discoideum clones with mutations in this gene transfer GlcNAc-P from UDP-GlcNAc to mannose residues at less than 5% the wild type value. Further, the lysosomal hydrolases of these mutant clones fail to bind to a cation-independent mannose 6-phosphate receptor affinity column, indicating a lack of methylphosphomannosyl residues on the high mannose oligosaccharides of these proteins. We conclude that the gpt1 gene product catalyzes the initial step in the formation of methylphosphomannosyl residues on D. discoideum lysosomal hydrolases.     

Phenotypic differences,spatial distribution and diversity at the Cytb and BMP4 genes in springbok (Antidorcas marsupialis)

The springbok (Antidorcas marsupialis) is a significant contributor to the economically important game ranching sector in Southern Africa. Phenotypic variation between springbok from the Karoo and Kalahari regions has been reported by several sources, with springbok from the Kalahari regarded as the larger form. There is no consensus on whether the two variants are determined by heredity, environment or a combination of the two. We studied variation in 80 individuals from four springbok populations using both a gene widely used for population studies (Cytb) and a gene that effects growth (BMP4). Results from Cytb haplotypes and BMP4 diploid gene sequences reveal moderate differentiation among springbok sampled from different regions. We also found a CA tandem repeat motive with high variability at the 3′ end of the BMP4 gene region sequenced (the third exon). There is some support for a hypothesis that nominally short and long fragments at this BMP4 repeat are associated with different populations, which may indicate either neutral genetic differentiation between spatially isolated forms, or a relationship between phenotype and BMP4 genotype. We also present new primer sequences to amplify both a partial fragment of the BMP4 gene region and the complete BMP4 tandem repeat motive in springbok.     

Unexpected Rarity of the Pathogen Batrachochytrium dendrobatidis in Appalachian Plethodon Salamanders: 1957–2011

Widespread population declines in terrestrial Plethodon salamanders occurred by the 1980s throughout the Appalachian Mountains, the center of global salamander diversity, with no evident recovery. We tested the hypothesis that the historic introduction and spread of the pathogenic fungus Batrachochytrium dendrobatidis (Bd) into the eastern US was followed by Plethodon population declines. We expected to detect elevated prevalence of Bd prior to population declines as observed for Central American plethodontids. We tested 1,498 Plethodon salamanders of 12 species (892 museum specimens, 606 wild individuals) for the presence of Bd, and tested 94 of those for Batrachochytrium salamandrivorans (Bs) and for ranavirus. Field samples were collected in 2011 from 48 field sites across a 767 km transect. Historic samples from museum specimens were collected at five sites with the greatest number and longest duration of collection (1957–987), four of which were sampled in the field in 2011. None of the museum specimens were positive for Bd, but four P. cinereus from field surveys were positive. The overall Bd prevalence from 1957–2011 for 12 Plethodon species sampled across a 757 km transect was 0.2% (95% CI 0.1–0.7%). All 94 samples were negative for Bs and ranavirus. We conclude that known amphibian pathogens are unlikely causes for declines in these Plethodon populations. Furthermore, these exceptionally low levels of Bd, in a region known to harbor Bd, may indicate that Plethodon specific traits limit Bd infection.     

Computational determination of the pigment binding motif in the chlorosome protein a of green sulfur bacteria

We present a molecular-scale model of Bacteriochlorophyll a (BChl a) binding to the chlorosome protein A (CsmA) of Chlorobaculum tepidum, and the aggregated pigment–protein dimer, as determined from protein–ligand docking and quantum chemistry calculations. Our calculations provide strong evidence that the BChl a molecule is coordinated to the His25 residue of CsmA, with the magnesium center of the bacteriochlorin ring situated <3 Å from the imidazole nitrogen atom of the histidine sidechain, and the phytyl tail aligned along the nonpolar residues of the α-helix of CsmA. We also confirm that the Q _y band in the absorption spectra of BChl a experiences a large (+16 to +43 nm) redshift when aggregated with another BChl a molecule in the CsmA dimer, compared to the BChl a in solvent; this redshift has been previously established by experimental researchers. We propose that our model of the BChl a–CsmA binding motif, where the dimer contains parallel aligned N-terminal regions, serves as the smallest repeating unit in a larger model of the para-crystalline chlorosome baseplate protein.     

Host Species and Environmental Effects on Bacterial Communities Associated with Drosophila in the Laboratory and in the Natural Environment

The fruit fly Drosophila is a classic model organism to study adaptation as well as the relationship between genetic variation and phenotypes. Although associated bacterial communities might be important for many aspects of Drosophila biology, knowledge about their diversity, composition, and factors shaping them is limited. We used 454-based sequencing of a variable region of the bacterial 16S ribosomal RNA gene to characterize the bacterial communities associated with wild and laboratory Drosophila isolates. In order to specifically investigate effects of food source and host species on bacterial communities, we analyzed samples from wild Drosophila melanogaster and D. simulans collected from a variety of natural substrates, as well as from adults and larvae of nine laboratory-reared Drosophila species. We find no evidence for host species effects in lab-reared flies; instead, lab of origin and stochastic effects, which could influence studies of Drosophila phenotypes, are pronounced. In contrast, the natural Drosophila–associated microbiota appears to be predominantly shaped by food substrate with an additional but smaller effect of host species identity. We identify a core member of this natural microbiota that belongs to the genus Gluconobacter and is common to all wild-caught flies in this study, but absent from the laboratory. This makes it a strong candidate for being part of what could be a natural D. melanogaster and D. simulans core microbiome. Furthermore, we were able to identify candidate pathogens in natural fly isolates.     

Isolation and identification of homeobox genes from the human placenta including a novel member of the Distal-less family,DLX4

We have carried out a DNA binding site screen of a 32-week human placental cDNA library using a consensus homeodomain binding site as a probe. This study represents the first library screen carried out to isolate homeobox genes from the human placenta. We have shown that three homeobox genes known to be expressed in the embryo, HB24, GAX and MSX2 are also expressed in the placenta. We have also identified a novel homeobox gene, DLX4, that shows 85% sequence identity with the homeodomain encoded by the Drosophila Distal-less (Dll) gene. DLX4 therefore represents a new member of the Distal-less family of homeobox genes. This is the first evidence that members of the Distal-less family of homeobox genes are expressed in the placenta. Using fluorescence in situ hybridisation (FISH), DLX4 has been assigned to human chromosome 17q21–q22. This places DLX4 in the same region of chromosome 17 as another member of the Distal-less family, DLX3 (Scherer et al., 1995), and the HOX-B homeobox gene cluster (Acampora et al., 1989; Boncinelli et al., 1991). Members of the Distal-less family (DLX1 and DLX2; DLX5 and DLX6) are found as closely linked pairs on human chromosomes (Simeone et al., 1994). We predict that DLX3 and DLX4 are closely linked and have arisen through gene duplication and divergence from a common ancestral precursor.     

Tree-based delimitation of morphologically ambiguous taxa: a study of the lizard malaria parasites on the Caribbean island of Hispaniola

Malaria parasites in the genus Plasmodium have been classified primarily on the basis of differences in morphology. These single-celled organisms often lack distinguishing morphological features, and this can encumber both species delimitation and identification. Six saurian malaria parasites have been described from the Caribbean island of Hispaniola. All six infect lizards in the genus Anolis, but only two of these parasites can be distinguished using morphology. The remaining four species overlap in morphology and geography, and cannot be consistently identified using traditional methods. We compared a morphological approach with a molecular phylogenetic approach for assessing the taxonomy of these parasites. We surveyed for blood parasites from 677 Anolis lizards, representing 26 Anolis spp. from a total of 52 sites across Hispaniola. Fifty-five of these lizards were infected with Plasmodium spp., representing several new host records, but only 24 of these infections could be matched to previously described species using traditional morphological criteria. We then estimated the phylogeny of these parasites using both mitochondrial (cytb and coxI) and nuclear (EF2) genes, and included carefully selected GenBank sequences to confirm identities for certain species. Our molecular results unambiguously corroborated our morphology-based species identifications for only the two species previously judged to be morphologically distinctive. The remaining infections fell into two well-supported and reciprocally monophyletic clades, which contained the morphological variation previously reported for all four of the morphologically ambiguous species. One of these clades was identified as Plasmodium floridense and the other as Plasmodium fairchildi hispaniolae. We elevate the latter to Plasmodium hispaniolae comb. nov. because it is polyphyletic with the mainland species Plasmodium fairchildifairchildi and we contribute additional morphological and molecular characters for future species delimitation. Our phylogenetic hypotheses indicate that two currently recognised taxa, Plasmodium minasense anolisi and Plasmodium tropiduri caribbense, are not valid on Hispaniola. These results illustrate that molecular data can improve taxonomic hypotheses in Plasmodium when reliable morphological characters are lacking.     

Tubby-tagged balancers for the Drosophila X and second chromosomes

We generated FM7a and CyO balancer chromosomes bearing a Tubby¹ (Tb¹) dominant transgene. Flies heterozygous for these FM7a and CyO derivatives exhibit a phenotype undistinguishable from that elicited by the Tb¹ mutation associated with the TM6B balancer. We tested two of these Tb-bearing balancers (FM7-TbA and CyO-TbA) for more than 30 generations and found that the Tb¹ transgene they carry is stable. Thus, these new Tb-tagged balancers are particularly useful for balancing lethal mutations and distinguish homozygous mutant larvae from their heterozygous siblings.     

Multiple components in ink of the sea hare Aplysia californica are aversive to the sea anemone Anthopleura sola

Sea hares of the genus Aplysia rely on an array of behavioral and chemical defenses, including the release of ink and opaline, to protect themselves from predation. While many studies have demonstrated that ink and opaline are repellent to predators, very little is known about which components of these secretions are active against predators. Ink was previously shown to facilitate the escape of Aplysia from predatory anemones (Anthopleura) by eliciting tentacle retraction and/or shriveling, and gastrovascular eversion, but the metabolites mediating this interaction were not identified. We investigated the metabolites in Aplysia californica secretions that were aversive to the anemone Anthopleura sola, as demonstrated by tentacle shriveling and/or retraction. We found that ink elicited tentacle shriveling and/or retraction, while opaline elicited a feeding response. The active components in ink do not appear to be diet-dependent, as ink was aversive regardless of diet (natural seaweed diet vs. Gracilaria ferox). Furthermore, metabolites extracted from G. ferox were not aversive, suggesting that the aversive components are produced by the sea hares. We then examined escapin, a protein in ink with antimicrobial properties. Escapin quickly forms reaction products when mixed with the amino acids l-lysine and l-arginine, which would occur when ink and opaline are released into the sea hare mantle cavity. Neither escapin alone nor escapin mixed with its amino acid substrate l-lysine elicited aversive behaviors either immediately before or 2 min before applying to the tentacles. In addition, escapin mixed with opaline and applied to tentacles after 2 min did not elicit a significant aversive response. Using bioassay-guided fractionation, we attempted to isolate the components in A. californica ink that are aversive to A. sola. We determined that multiple components in ink, including both lipophilic and hydrophilic constituents, elicited aversive responses. We hypothesize that these components may facilitate A. californica's escape from A. sola by eliciting tentacle shriveling and/or retraction, which lead to anemones dropping ensnared sea hares.