The central nervous system of Oweniidae (Annelida) and its implications for the structure of the ancestral annelid brain.

BACKGROUND: Recent phylogenomic analyses congruently reveal a basal clade which consists of Oweniidae and Mageloniidae as sister group to the remaining Annelida. These results indicate that the last common ancestor of Annelida was a tube-dwelling organism. They also challenge traditional evolutionary hypotheses of different organ systems, among them the nervous system. In textbooks the central nervous system is described as consisting of a ganglionic ventral nervous system and a dorsally located brain with different tracts that connect certain parts of the brain to each other. Only limited information on the fine structure, however, is available for Oweniidae, which constitute the sister group (possibly together with Magelonidae) to all remaining annelids. RESULTS: The brain of Oweniidae is ring- shaped and basiepidermal. Ganglia, higher brain centers or complex sensory organs do not exist; instead the central nervous system is medullary. Posterior to the brain the ventral medullary cord arises directly from the ventral region of the brain in Myriowenia sp. while in Owenia fusiformis two medullary cords arise perpendicular to the brain ring, extend caudally and fuse posterior. The central nervous system is composed of a central neuropil and surrounding somata of the neurons. According to ultrastructural and histological data only one type of neuron is present in the central nervous system. CONCLUSION: The central nervous system of Oweniidae is the simplest in terms of enlargement of the dorsal part of the brain and neuron distribution found among Annelida. Our investigation suggests that neither ganglia nor commissures inside the brain neuropil or clusters of polymorphic neurons were present in the annelid stem species. These structures evolved later within Annelida, most likely in the stem lineage of Amphinomidae, Sipuncula and Pleistoannelida. Palps were supposedly present in the last common ancestor of annelids and innervated by two nerves originating in the dorsal part of the brain. A broader comparison with species of each major spiralian clade shows the medullary nervous system to be a common feature and thus possibly representing the ancestral state of the spiralian nervous system. Moreover, ganglia and clusters of polymorphic neurons seemingly evolved independently in the compared taxa of Spiralia and Annelida.

Developmental studies provide new insights into the evolution of sense organs in Sabellariidae (Annelida).

BACKGROUND: Sabellarids, also known as honeycomb or sandcastle worms, when building their tubes, produce chemical signals (free fatty acids) that are responsible for larval settlement and the formation of three-dimensional aggregations. The larval palps and the dorsal hump (becoming the median organ in adults) are presumed to participate in such a substrate selection during settlement. Notably, the sabellariid median organ is an apparently unique organ among annelids that has been attributed with a sensory function and perhaps with some affinities to the nuchal organs of other polychaetes. Nevertheless, detailed investigations of this prominent character complex including ultrastructural examinations are lacking so far. RESULTS: Our comprehensive investigations provide data about the anterior sensory organs in Sabellariidae and inform about their transformation during pelagic larval development. We used a comparative approach including immunostaining with subsequent confocal laser scanning microscopy (clsm), histological sections as well as electron microscopy in a range of larval and adult stages of two sabellariid species. We find that the neuronal innervation as well as the ultrastructure of the sabellariid ciliary structures along the median organ are highly comparable with that of nuchal organs known from other polychaetes. Furthermore, the myoinhibitory protein (MIP) - a protein known to be also involved into chemo-sensation - was detected in the region of the larval median organ. Moreover, we reveal the presence of an unusual type of photoreceptor as part of the median organ in Idanthyrsus australiensis with a corrugated sensory membrane ultrastructure unlike those observed in the segmental ocelli of other polychaetes. CONCLUSIONS: We are describing for the first time the nuchal organ-like structures in different developmental stages of two species of Sabellariidae. The external morphology, neuronal innervation, developmental fate and ultrastructure of the newly-discovered median organ-based ciliary pits are comparable with the characteristics known for annelid nuchal organs and therefore indicate a homology of both sensory complexes. The presence of myoinhibitory peptide (MIP) in the respective region supports such a hypothesis and exhibits the possibility of an involvement of the entire sabellariid median organ complex, and in particular the prominent ciliated pits, in chemo-sensation.

Barcoding and multi-locus phylogeography of the globally distributed calcareous tubeworm genus Hydroides Gunnerus, 1768 (Annelida, Polychaeta, Serpulidae).

Hydroides is a large and diverse group of calcareous tubeworms (Serpulidae, Annelida) recognised by a distinctive but variable two-tiered operculum. Despite considerable research using several species of Hydroides as models in ecological and biofouling studies, phylogenetic and biogeographic relationships within the genus are still poorly understood. Using combined mitochondrial (COI, cytochrome b) and nuclear (18S, 28S and ITS) gene markers for 284 individuals of 45 morphospecies of Hydroides, we investigated the global phylogenetic and biogeographic relationships within the genus. Phylogenetic topologies were well supported and indicated high genetic diversity within Hydroides, revealing potential cryptic species. Present results also include the first COI barcoding data enabling rapid and effective species identification of Hydroides on a global scale. Phylogenetic relationships within Hydroides were more concordant with geographical distributions than morphological similarity of their opercula. Molecular divergence estimates suggested the origin and subsequent diversification in the western Tethys Sea followed by a shift of the historical centre of diversity from the Indo-Mediterranean region to the central Indo-Pacific during the last 50 million years. Further studies on population genetics of species consisting of multiple lineages would provide a better understanding on the status of potential cryptic species. Furthermore, paleogeographic studies based on fossil Hydroides tubes would provide evidence to test this biogeographic hypothesis.

Four new species of Marphysa (Annelida, Eunicida, Eunicidae) from the east coast of Peninsular Malaysia.

Four new species of Marphysa are described from Terengganu state on the east coast of Peninsular Malaysia, using morphological and molecular (cytochrome oxidase subunit I (COI) gene) data. These species belong to different groups of Marphysa: Marphysakertehensis sp. nov. belongs to Group A (Mossambica), Marphysamerchangensis sp. nov. and Marphysasetiuense sp. nov. belong to Group B (Sanguinea) and Marphysaibaiensis sp. nov. belongs to Group E (Gravelyi). Marphysakertehensis sp. nov. is characterised by having only limbate chaetae, absence of subacicular hooks, three types of pectinate chaetae including wide, thick isodont with short and slender inner teeth, and pectinate branchiae with up to nine branchial filaments. Marphysamerchangensis sp. nov. is characterised by the presence of eyes, unidentate subacicular hooks, four types of pectinate chaetae including wide, thick anodont pectinate chaetae with five long and thick inner teeth, and pectinate branchiae with up to six branchial filaments. Marphysasetiuense sp. nov. has mostly unidentate subacicular hooks (bidentate on several posterior chaetigers), four types of pectinate chaetae including wide, thick anodont pectinate chaetae with seven thick and long inner teeth, and pectinate branchiae with up to five branchial filaments. Marphysaibaiensis sp. nov. has bidentate subacicular hooks throughout, five types of pectinate chaetae, including a heterodont with 12 short and slender inner teeth, and pectinate branchiae with up to eight branchial filaments. The designation of these new species based on morphology is fully supported by molecular data. Habitat descriptions of each species are also included.

New species of Terebellides (Polychaeta: Trichobranchidae) from the deep Southern Ocean, with a key to all described species.

The genus Terebellides is, despite its often low abundances, a common and diverse element of benthic, soft sediment communities at all depths. In recent years, careful examination of specimens has resulted in numerous descriptions of new species of Terebellides increasing the number of species in the genus to over forty. For the Southern Ocean currently only two species are considered valid, both recorded for shelf and slope depths. Here, we present findings of eleven new Antarctic species originating from depths between 480 m and 4720 m. Six of these are formally described (T. canopus sp. n., T. crux sp.n., T. mira sp.n., T. rigel sp.n., T. sirius sp.n., and T. toliman, sp.n.). One species, T. crux sp.n., bears two segments with geniculate hooks, a trait already known for the genus but conflicting with the original generic diagnosis. To include this trait the generic diagnosis of Terebellides is amended. An identification key for all species of Terebellides is given, as well as a maximum parsimony phylogeny of the genus based on morphological characters.

The genus Syllis Savigny in Lamarck, 1881 (Annelida: Syllidae: Syllinae) from Australia (Part 3): new species and redescription of previously described species.

Syllis Savigny in Lamarck, 1818, the type genus of Syllidae, is the largest and most diverse within the family. This genus presents many taxonomic, biogeographic, and phylogenetic problems due mainly to the lack of molecular data and morphological synapomorphies for many of the species, but also due to poor or inadequate descriptions. In order to improve the knowledge of the genus, we have undertaken a morphological revision of Australian species, based on type material and material collected over years by the Australian Museum. This is the third paper dealing with this family. Eighteen species are herein described, of which five are new and twelve are redescriptions of already known species, with four new combinations: Syllis imajimai, n. sp.; S. narranuk, n. sp.; S. noolinga, n. sp.; S. similisunzima, n. sp.; S. thylacine, n. sp.; S. busseltonensis (Hartmann-Schröder, 1982) n. comb.; S. patriciae (Hartmann-Schröder, 1981) n. comb.; S. pharobroomensis (Hartmann-Schröder, 1979) n. comb., S. pharynxcircunfusata (Hartmann-Schröder, 1979) n. comb, S. armillaris (O. F. Müller, 1771); S. augeneri Haswell, 1920; S. benelihauae (Campoy & Alquézar, 1982); S. hyalina Grube, 1863; S. cf. nigrescens Grube, 1878; S. profunda Cognetti, 1955; S. truncata Haswell, 1920; and S. variegata Grube, 1860.

Two new species of Marphysa (Annelida, Eunicidae) from southern Australia.

Two new species of Marphysa Quatrefages, 1866 are described from the southeast coast of Australia. With the presence of only compound spinigers and the branchiae present over many chaetigers, Marphysa baudini n. sp. belongs to the Sanguinea-group. This species has ventral cirri with an inflated base and digitiform tip and thick and wide anodont pectinate chaetae, with 3-5 internal long and thick teeth. With the presence of only compound falcigers, Marphysa davidattenboroughi n. sp., belongs to the "Aenea-group". This species is characterised by the presence of a bilobed prostomium, a single pair of pygidial cirri and by the presence of thick, and wide anodont pectinate chaetae with 4-6 long internal and thick teeth.

Research publications of Australia's natural history museums, 1981-2020: Enduring relevance in a changing world.

As a case study of the responses of natural history museums to changing scientific and funding environments, we analysed research publications of Australia's Natural History Museums (ANHMs) 1981-2020. Using Scopus, 9,923 relevant documents 1981-2020 were identified, mainly research papers but with a growing proportion of reviews. The number of documents published increased over tenfold from 39 (1981) to 553 (2020), likely driven by collaborations (rising from 28.5% of documents 1981-1985 to 87.2% of documents 2016-2020), contributions from retired staff, and volunteer support. The mean length of documents (pages) ranged from a low of 15.3 in 2001-2005 to a high of 17.4 in 1991-1995, but this statistically significant result was trivial in practical terms. The sources (i.e., journals, book titles, conference proceedings) in which ANHM authors published changed over time, with growing proportions of publications in journals covering molecular ecology/phylogenetics and biological conservation. We identified the major areas of study canvassed within the corpus of publications by developing structural topic models based on patterns of word use in document titles, abstracts and keyword lists. The topics discovered included study subjects traditional for natural history museums (new taxa, phylogeny, systematics, animal morphology, palaeontology, minerals), new directions (molecular genetics, ecology, biological conservation) and marine biology (probably reflecting Australia's large coastline). Most citations came from Australia, USA and UK, although in 2016-2020 only 27.9% of citing documents included an Australian author. Growth in numbers of documents and collaborations, as well as use of documents internationally over a period of great change in scientific and funding environments, indicate an enduring legacy of ANHM research, grounded on the intrinsic value of the collections.

A new species of giant Eunice (Eunicidae, Polychaeta, Annelida) from the east coast of Australia.

A new giant species is described from New South Wales, Australia. Eunicedharastii sp. nov. differs from described Australian species and is most similar to E.aphroditois (Pallas, 1788), E.flavopicta Izuka, 1912, and E.kinbergi Ehlers, 1868. The unique combination of features that characterizes the new species is irregular articulated prostomial appendages; antennae reaching back beyond chaetiger 4; branchiae starting at chaetiger 10, initially button-shaped and distinctly longer than notopodial cirri where best developed; dorsal fleshy knobs on anterior chaetal lobes; notopodial cirri pendulous, abrupt tapering from inflated bases; bidentate compound falcigerous chaetae with both teeth directed laterally, distal tooth much shorter than proximal tooth in median and posterior chaetigers; and dark bidentate subacicular hooks starting at chaetiger 58, tapering to a small head with both teeth directed distally, and proximal tooth much larger than minute and spur-like distal tooth. This new species lives in sandy sediments in coastal waters 1-8 m deep. It is highly mobile and not easy to collect, which may explain why it was not described before.

Pectinariidae (Annelida, Polychaeta) from the coastal waters of China, with description of new species and new records.

Pectinariidae Quatrefages, 1865 is widely distributed, and easily distinguished from other polychaetes by their characteristic semi-circular operculum with two rows of stout golden paleae and ice-cream cone-shaped tubes made of sand or shell fragments. A total of 387 specimens of pectinariids from the coastal waters of China were examined. Eleven species belonging to three genera of pectinariids were identified and described, including four new species (Amphictene jianqingi n. sp., Pectinaria lizhei n. sp., Pectinaria dayaensis n. sp., Pectinaria xiukaii n. sp.) and one new record (Pectinaria plurihamus Choi, Jung Yoon, 2017). A key to genera of Pectinariidae and recorded species from Chinese waters is given. Based on partial sequences of five genes (COI, 16S rDNA, 18S rDNA, 28S rDNA and H3), the phylogenetic relationship of pectinariids is analyzed. The three new species Amphictene jianqingi n. sp., Pectinaria lizhei n. sp. and Pectinaria dayaensis n. sp. with molecular data are well supported. Three species of Lagis (L. koreni, L. bocki and L. plurihamus), five species of Pectinaria (P. dayaensis, P. dodeka, P. gouldii, P. lizhei and P. torquata), and three species of Amphictene (A. auricoma, A. alata and A. jianqingi) do not form a well supported clade, which requires more detailed analyses.

Three new deep-sea species of Marphysa (Annelida, Eunicida, Eunicidae) from Papua New Guinea (Bismarck and Solomon seas).

Three new species of Marphysa Quatrefages, 1866, Marphysabanana sp. nov., Marphysapapuaensis sp. nov., and Marphysazanolae sp. nov. are described from deep-sea sunken vegetation off Papua New Guinea, using both morphology and molecular data (for two species). With the presence of compound spinigers only and the branchiae present over many chaetigers, Marphysabanana sp. nov. belongs to the group B2. This species is characterised by the presence of eyes, the presence of branchiae starting from chaetiger 20, and by the presence of three types of pectinate chaetae and bidentate subacicular hooks starting from chaetigers 13-52. With the presence of compound falcigers only and the branchiae restricted to a short anterior region, Marphysapapuaensis sp. nov. belongs to the group C1. This species has a bilobed prostomium but no eyes, has branchiae from chaetigers 7 to 14-16 with up to 16 filaments. Marphysapapuaensis sp. nov. is also characterised by the presence of bidentate subacicular hooks from chaetiger 20 and by a single type of pectinate chaetae. Finally, Marphysazanolae sp. nov. belongs to the group C2, with the presence of compound falcigers only and the branchiae present over many chaetigers. This species is characterised by the absence of eyes, by the presence of branchiae with a single long filament starting from chaetiger 31, by unidentate subacicular hooks starting from chaetiger 28 and finally by one type of pectinate chaetae with very long outer teeth.

Phylogeny of Sabellidae (Annelida) and relationships with other taxa inferred from morphology and multiple genes.

The monophyly of Sabellidae, the phylogenetic relationships of its lineages, and the composition of Sabellida have been debated for many decades. Most studies on sabellid phylogeny have focused on morphological features but little DNA work has been published to date. We performed analyses using maximum-parsimony methods that included 36 sabellids and members of previously related taxa. We integrated morphological and DNA sequence data to resolve relationships at different hierarchical levels (135 morphological features, fragments of the nuclear ribosomal RNA genes 18S and 28S, and the mitochondrial gene 16S). The results indicate the monophyly of Sabellida, including Sabellidae and Serpulidae. Monophyly of Fabriciinae and Serpulidae is assessed and the two groups are recovered as sister taxa, but with weak support. There is no significant support for the monophyly of Sabellinae. Relationships between members of the Sabellidae are still partially unresolved due to incongruence between partitions and low support for most clades. The evolution and transformation of certain characters within Sabellidae is explored. © The Willi Hennig Society 2010.

A new species of Petta (Annelida, Pectinariidae), with comments on Pettaassimilis McIntosh, 1885.

The genus Petta Malmgren, 1866 is a small and poorly known genus of the annelid family Pectinariidae Quatrefages, 1866. A previous revision of the genus found that the type material of the species P.assimilis McIntosh, 1885 had been lost. While searching for material from the type locality, we were able to examine material from a similar area but collected in much shallower water from off South Africa which represents another undescribed species of Petta. The new species, Pettabrevis sp. nov., is described and compared to P.assimilis McIntosh, 1885, and a revised key to all species in the genus is provided.

Biodiversity of intertidal marine flatworms (Polycladida, Platyhelminthes) in southeastern Australia.

Flatworms of the Order Polycladida are a group of free-living invertebrates found in a diversity of marine habitats, with over 800 species described worldwide. Marine flatworms are a conspicuous component of Australias marine fauna yet have received little attention. Less than 30 scientific articles have been published on Australian marine flatworms since 1855, of which only nine include species from southeastern Australia. Here, the biodiversity and distribution of species belonging to the Order Polycladida inhabiting intertidal rocky beaches in southeastern Australian waters were identified and analysed. Sampling was conducted at low tide along the coasts of New South Wales and Victoria. Collected samples were serially sectioned for comparative anatomical studies, and tissue was removed from each individual for molecular sequencing and analyses. Both nuclear and mitochondrial DNA sequences were obtained and used as an additional source of evidence for the description of new species as well as providing further insight into the phylogenetic relationships between them. A total of 20 species, six of which are new (e.g., Eulatocestus australis sp. nov.), and a new genus (Parabolia gen. nov.) have been described, as well as two new records for Australia (e.g., Stylochoplana clara Kato, 1937) have been identified increasing our knowledge of this important component of the Australian marine biota.

Revision of the French Terebellidae sensu stricto (Annelida, Terebelliformia), with descriptions of nine new species.

This work is the last of four papers of the Spaghetti Project, aiming to revise the species of terebellids, a.k.a. spaghetti worms, present in the European French waters. In this last paper the Terebellidae, sensu stricto, from French waters are revised based, on material available in the French marine stations, type materials stored in the MNHN collection and newly collected specimens. Nine new species are described using both morphological and molecular tools: Eupolymnia gili n. sp., E. lacazei n. sp., E. meissnerae n. sp., Lanice kellyslateri n. sp., Paramphitrite dragovabeci n. sp., Pista labruneae n. sp., P. miosseci n. sp., P. sauriaui n. sp., and Terebella banksyi n. sp. European species of Eupolymnia are distinguished mainly by the shape of the lateral lobes and the size of the branchial stems. The two species belonging to Lanice genus are distinguished by the fusion of the first ventral shields, the shape of both noto- and neuropodia, and the pigmentation of the upper lip. The two species of Paramphitrite are distinguished by the presence or absence of a medial dorsal gap between the pairs of branchiae, by the shape of the lateral lobes and the presence or absence of a nephridial papilla on segment 4. The different species of Pista are distinguished by the number of pairs of branchiae, the shape of the lateral lobes and uncini. Finally, the two species of Terebella are distinguished by the number of segments with nephridial and genital papillae and the segments on which the branchiae occur. An identification key for European species of Terebellidae sensu stricto is also provided.

Segmented worms (Phylum Annelida): a celebration of twenty years of progress through Zootaxa and call for action on the taxonomic work that remains.

Zootaxa has been the leading journal on invertebrate systematics especially within Annelida. Our current estimates indicate annelids include approximately 20,200 valid species of polychaetes, oligochaetes, leeches, sipunculans and echiurans. We include herein the impact of Zootaxa on the description of new annelid species in the last two decades. Since 2001, there have been over 1,300 new annelid taxa published in about 630 papers. The majority of these are polychaetes (921 new species and 40 new genera) followed by oligochaetes (308 new species and 10 new genera) and leeches (21 new species). The numerous papers dealing with new polychaete species have provided us a clear picture on which polychaete families have had the most taxonomic effort and which authors and countries have been the most prolific of descriptions of new taxa. An estimated additional 10,000+ species remain to be described in the phylum, thus we urge annelid workers to continue their efforts and aid in training a new generation of taxonomists focused on this ecologically important group.

Annelids of the eastern Australian abyss collected by the 2017 RV 'Investigator' voyage.

In Australia, the deep-water (bathyal and abyssal) benthic invertebrate fauna is poorly known in comparison with that of shallow (subtidal and shelf) habitats. Benthic fauna from the deep eastern Australian margin was sampled systematically for the first time during 2017 RV 'Investigator' voyage 'Sampling the Abyss'. Box core, Brenke sledge, and beam trawl samples were collected at one-degree intervals from Tasmania, 42°S, to southern Queensland, 24°S, from 900 to 4800 m depth. Annelids collected were identified by taxonomic experts on individual families around the world. A complete list of all identified species is presented, accompanied with brief morphological diagnoses, taxonomic remarks, and colour images. A total of more than 6000 annelid specimens consisting of 50 families (47 Polychaeta, one Echiura, two Sipuncula) and 214 species were recovered. Twenty-seven species were given valid names, 45 were assigned the qualifier cf., 87 the qualifier sp., and 55 species were considered new to science. Geographical ranges of 16 morphospecies extended along the eastern Australian margin to the Great Australian Bight, South Australia; however, these ranges need to be confirmed with genetic data. This work providing critical baseline biodiversity data on an important group of benthic invertebrates from a virtually unknown region of the world's ocean will act as a springboard for future taxonomic and biogeographic studies in the area.

Two new species of Sabellariidae (Annelida, Polychaeta) from the abyss of eastern Australia.

In May-June 2017 an expedition on board RV 'Investigator' sampled benthic communities along the lower slope and abyss of eastern Australia from off Tasmania to the Coral Sea. Over 200 sabellariid specimens of the genera Phalacrostemma and Gesaia were collected during the voyage and deposited in the Australian Museum. Here we describe two new species Gesaia csiro n. sp. (4414-4436 m) and Phalacrostemma timoharai n. sp. (1013-1093 m). We did not formally describe another species of Phalacrostemma due to poor condition of the single specimen. Gesaia csiro n. sp. is the first record of the genus from Australian waters (only a planktonic larva attributed to the genus has previously been recorded), and it can be distinguished from other congeners by the smooth surface of inner paleae, distal thecae of outer paleae with long, irregular and expanded distal fringe and circled distal margin. Phalacrostemma timoharai n. sp. differs from congeners by the following combination of characters: presence of the buccal flap, absence of tentacular filament, 18-22 pairs of outer paleae, two pairs of neuropodial cirri on first thoracic segment, and only one pair of lateral lobes on second thoracic segment. Morphological descriptions are accompanied by mitochondrial cytochrome c oxidase subunit 1 (COI) and ribosomal (16S, 18S and 28S) sequence data. A key to all Australian species of sabellariids is given.

Eunice sensu lato (Annelida: Eunicidae) from Australia: description of seven new species and comments on previously reported species of the genera Eunice, Leodice and Nicidion.

Species traditionally considered to belong to Eunice are now, also, distributed in two other genera Leodice and Nicidion recently resurrected to reconcile Eunicidae taxonomy with its phylogenetic hypothesis. In Australia, Eunice species have been reported from all seas and sum up to 22 species. In this study, we propose 10 new combinations for traditional Eunice species, which should be moved to Leodice; describe seven new species, four of Eunice and three of Leodice; and comment on species previous recorded from the Australian Coast. Previous records of E. denticulata, E. filamentosa, E. grubei, E. indica, E. longicirris, E. microprion, E. paupera, E. tridentata, E. tubifex and E. vittata from Australia are considered dubious. Eunice tribranchiata, originally described from Australian specimens, is considered here to be an indeterminable species. This study also includes descriptions of Eunice, Leodice and Nicidion as well as key to these genera and their species recorded from Australia.

Revision of the French Polycirridae (Annelida, Terebelliformia), with descriptions of eight new species.

Polycirridae from French waters are revised based on material available in French marine stations and newly collected specimens. This work is the third part of the Spaghetti Project aiming to revise French species of "Spaghetti" terebellid worms. It describes eight new species using both morphological (for all species) and molecular (for some species) tools: Amaeana gremarei n. sp., Polycirrus catalanensis n. sp., P. glasbyi n. sp., P. gujanensis n. sp., P. idex n. sp., P. nogueirai n. sp., P. pennarbedae n. sp. and P. readi n. sp., in addition to a previously described species. An identification key for European species of Polycirridae is also provided.