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Sponge Systematics Facing New Challenges
Abstract and Figures
Systematics is nowadays facing new challenges with the introduction of new concepts and new techniques. Compared to most other phyla, phylogenetic relationships among sponges are still largely unresolved. In the past 10 years, the classical taxonomy has been completely overturned and a review of the state of the art appears necessary. The field of taxonomy remains a prominent discipline of sponge research and studies related to sponge systematics were in greater number in the Eighth World Sponge Conference (Girona, Spain, September 2010) than in any previous world sponge conferences. To understand the state of this rapidly growing field, this chapter proposes to review studies, mainly from the past decade, in sponge taxonomy, nomenclature and phylogeny.
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... As animals with indeterminate growth, where morphological characters may be shaped more by environment than by genetics, sponges can exhibit considerable phenotypic plasticity, (Hill & Hill, 2002;Kaandorp, 1999;Klautau et al., 1999;Loh et al., 2012;López-Legentil et al., 2010;McDonald et al., 2002;Palumbi, 1984Palumbi, , 1986. Conventional reliance on growth forms, skeletal, and spicule characteristics to inform sponge taxonomy suffers from a high potential for confounding phenotypic plasticity (genetically indistinct, phenotypically distinct) (Cárdenas et al., 2012;DeBiasse & Hellberg, 2015), crypsis (genetically distinct and phenotypically indistinct; Andreakis et al., 2012;Swierts et al., 2017) and convergent morphological evolution (Schuster et al., 2015). Resolving the relationship between phylogenetically delimited species (hereafter, 'phylospecies') and morphologically defined subpopulations ('morphospecies') through integrated molecular analyses has become critical as many coral reefs transition to sponge-dominated communities (Cruz- Barraza et al., 2012;Erpenbeck & Wörheide, 2007;Hill et al., 2013;Pankey et al., 2022;Redmond et al., 2013;Wörheide et al., 2012). ...
Fundamental to holobiont biology is recognising how variation in microbial composition and function relates to host phenotypic variation. Sponges often exhibit considerable phenotypic plasticity and also harbour dense microbial communities that function to protect and nourish hosts. One of the most prominent sponge genera on Caribbean coral reefs is Agelas . Using a comprehensive set of morphological (growth form, spicule), chemical and molecular data on 13 recognised species of Agelas in the Caribbean basin, we were able to define only five species (=clades) and found that many morphospecies designations were incongruent with phylogenomic and population genetic analyses. Microbial communities were also strongly differentiated between phylogenetic species, showing little evidence of cryptic divergence and relatively low correlation with morphospecies assignment. Metagenomic analyses also showed strong correspondence to phylogenetic species, and to a lesser extent, geographical and morphological characters. Surprisingly, the variation in secondary metabolites produced by sponge holobionts was explained by geography and morphospecies assignment, in addition to phylogenetic species, and covaried significantly with a subset of microbial symbionts. Spicule characteristics were highly plastic, under greater impact from geographical location than phylogeny. Our results suggest that while phenotypic plasticity is rampant in Agelas , morphological differences within phylogenetic species affect functionally important ecological traits, including the composition of the symbiotic microbial communities and metabolomic profiles.
... Sponge systematics is based mainly on the composition of the skeleton and structure. Traditionally, spicules (morphology and size) and spongin fibers have been useful as taxonomic characters, especially when they have many different types and a complex skeletal organization or structure (Cárdenas et al. 2012). With the development of molecular tools, the systematic description of new species has incorporated DNA barcodes and other molecular markers from the mitochondrial or nuclear genomes to strengthen taxonomic hypotheses on species delimitation; these markers are sequences specifically obtained from individuals used to describe the species Uriz 2008, Cárdenas et al. 2012). ...
Identifying cryptic species is pivotal for understanding marine biodiversity and optimizing strategies for its conservation. A robust understanding of poriferan diversity is a complex endeavour. It has also been extremely hampered by the high phenotypic plasticity and the limited number of diagnostic characters. Mycale (Carmia) cecilia has different body colours, even among individuals living together. We tested whether the colour variation could be due to polymorphism, phenotypic plasticity or cryptic speciation. Phylogenetic reconstructions of nuclear and mitochondrial loci were congruent. Individuals of different body colour did not cluster together and had high levels of genetic divergence. Furthermore, the green morphotype clustered in almost all reconstructions with Mycale (C.) phyllophila, as both showed higher gene similarity at the transcriptomic level (public transcriptome). Morphologically, the green individuals consistently showed discrepancies from the red ones. These results suggest that all individuals with the same body colour, either red or green, correspond to the same species, while individuals with different body colours probably belong to different species. These results reveal high levels of morphologic and genetic diversity, which could have important implications for what is known as M. (C.) cecilia and the Mycalidae systematics.
... Chemotaxonomy is the study of how chemical compounds or groups of chemicals that are produced by living organisms are distributed among different plant species that are related or potentially related [8]. Chemotaxonomy possesses various benefits, such as directing natural product research towards specific plant groups [7], resolving existing taxonomical problems [9], serving as a complementary or essential tool for deciphering classification patterns and identifying potential synapomorphic chemical markers at different taxonomic levels [10], and facilitating evolutionary studies that examine phylogenetic and infra-taxonomic variations. The exclusive reliance on external morphological analysis is inadequate for a comprehensive assessment of a taxon's systematic classification and evolutionary relationships. ...
Chemotaxonomy is a valuable tool for obtaining taxonomic insights, which are most effectively employed in combination with other forms of data to establish a system of classification that closely reflects natural connections. The utilization of plant secondary metabolites possessing diverse therapeutic qualities signifies the growing exploitation of natural products in the medical discipline. The objectives of the current study encompassed the identification of phytochemicals in the extracts of nine species of medicinal plants, the examination of their chemotaxonomic properties, and the assessment of the antibacterial and antioxidant capabilities exhibited by the extracts. GC-MS technology was employed for the identification of phytochemical compounds. The study utilized ClassyFire, an automated chemical classification system that incorporates an extensive and computable classification, to categorize chemicals. The chemical classification of plants was examined by the application of principal component analysis (PCA) and cluster analysis (CA). The bactericidal properties of plants were assessed against four harmful bacterial species using the disc diffusion technique. The antioxidant properties of plant extracts were assessed employing the DPPH free radical scavenging methodology, and the half maximal effective concentration (EC50) was determined using dose response models. The calculator being referred to is the Quest Graph™ EC50 Calculator. In the plant extracts, the analysis disclosed the occurrence of 160 phytochemicals, classified into 36 phytochemical classes. The results of CA and PCA demonstrated the proximity and associations among Asteraceae species, while indicating the divergence of the two Lamiaceae species. Achillea fragrantissima and Ducrosia flabellifolia demonstrated the most diversity in phytochemical classes, while Thymus vulgaris displayed the highest level of dominance. Pulicaria undulata and T. vulgaris had the most notable antibacterial activity. D. flabellifolia and P. incisa demonstrated the highest levels of antioxidant activity. Ethanol exhibited superior antibacterial efficacy compared to other solvents. The remarkable biological activities exhibited by these plant extracts can be ascribed to the copious presence of certain chemicals, predominantly sesquiterpenoids, monoterpenoids, benzene and its derivatives, naphthalenes, fatty acyls, and phenols. The susceptibility of Gram-positive bacterial species to plant extracts was shown to be higher in comparison to Gram-negative bacterial species.
The sponge fauna of the Western Mediterranean stands as one of the most studied in the world. Yet sampling new habitats and a poorly studied region like the Balearic Islands highlights once again our limited knowledge of this group of animals. This work focused on demosponges of the order Tetractinellida collected in several research surveys (2016–2021) on a variety of ecosystems of the Balearic Islands, including shallow caves, seamounts and trawl fishing grounds, in a broad depth range (0–725 m). Tetractinellid material from the North Atlantic and more than twenty type specimens were also examined and, for some, re-described in this work. All species were barcoded with the traditional molecular markers COI (Folmer fragment) and 28S (C1-C2 or C1-D2 fragment). A total of 36 species were identified, mostly belonging to the family Geodiidae (15 species), thereby bringing the number of tetractinellids recorded in the Balearic Islands from 15 to 39. Eight species from this study are new: Stelletta mortarium sp. nov., Penares cavernensis sp. nov., Penares isabellae sp. nov., Geodia bibilonae sp. nov., Geodia microsphaera sp. nov. and Geodia matrix sp. nov. from the Balearic Islands; Geodia phlegraeioides sp. nov. and Caminus xavierae sp. nov. from the North East Atlantic. Stelletta dichoclada and Erylus corsicus are reported for the first time since their description in Corsica in 1983. Pachastrella ovisternata is documented for the first time in the Mediterranean Sea. Finally, after comparisons of type material, we propose new synonymies: Geodia anceps as a junior synonym of Geodia geodina, Erylus cantabricus as a junior synonym of Erylus discophorus and Spongosorites maximus as a junior synonym of Characella pachastrelloides.
Acanthotetilla is a genus of tetillid sponges diagnosed by the presence of medium-sized, heavily spined oxeas, which are known as “megacanthoxeas” (or simply acanthoxeas). Currently, seven species are known worldwide: three from the Western Atlantic (A. gorgonosclera, A. rocasensis and A. walteri), three from the Western Indian (A. hemisphaerica, A. enigmatica and A. seychellensis) and one from the Western Pacific (A. celebensis). A megacanthoxea-bearing specimen was obtained from the Fernando de Noronha Archipelago, Southwest Atlantic, in 2016. It is described here as a new Acanthotetilla, namely, A. noronhensis sp. nov. This new species has two categories of megacanthoxeas, like A. celebensis from Celebes/Sulawesi Sea, Indonesia. However, the former differs from the latter, as well as from other congeners, by the combination of endolithic habit and spiculation. Acanthotetilla noronhensis sp. nov. is known on the basis of a single specimen from a large collection of sponges from Fernando de Noronha. Besides, the species was not found in other sponge collections from different areas of Brazil, including oceanic areas. Thus, the new species has probably a low abundance and restricted distribution, alike all other known Acanthotetilla species, aside A. celebensis, which (regardless its low abundance) occurs in a large sector of the western Pacific. Despite A. noronhensis sp. nov. and A. celebensis being separated by a wide geographic distance, the unique occurrence of two categories of megacanthoxeas might suggest closer phylogenetic relationship and biogeographic history as well. A wider discussion on the diversity, distribution and morphologic characters of Acanthotetilla spp. is supplied in this study. A comparative taxonomic table and a taxonomic identification key to Acanthotetilla spp. is supplied as well.
The demosponge genus Placospongia includes 13 valid species, of which six occur in the Brazilian Exclusive Economic Zone. In the present study, we describe two new species of Placospongia from Northern (off the Amazon River mouth) and Northeastern Brazilian continental shelf and oceanic islands (i.e., Fernando de Noronha Archipelago), viz., Placospongia beatrizae sp. nov. and Placospongia amphinolastra sp. nov., and redescribe the holotype of Placospongia cristata Boury-Esnault, 1973, a poorly known species. In addition, we revise the Brazilian records of the allegedly cosmopolitan species P. carinata, P. melobesioides and P. intermedia, and provide new records of P. colombiensis and P. ruetzleri from Brazil. We demonstrate that P. carinata sensu Coelho & Mello-Leitão and P. intermedia sensu Muricy & Moraes and Jimenez et al. belong in fact to P. ruetzleri, and that P. melobesioides sensu Mothes et al. and González-Farías is a new species, described here as Placospongia beatrizae sp. nov. With the description of the two new species and synonymyzation of previous records, six species of Placospongia are currently known in Brazilian waters, two of which are endemic. A key to Placospongia species is also provided.
The South Orkney Islands (SOI) is a poorly studied Antarctic archipelago located in the Scotia Arc. In this study, we added 25 Demospongiae species to the SOI known sponge richness, which was previously represented by only 16 species. Of these, 22 species represent new records for SOI region, although they were previously recorded from other Antarctic sectors. Samples were collected during two Argentinean Antarctic Research Cruises onboard RV Puerto Deseado. The most frequently collected species were Artemisina apollinis, Iophon gaussi, I. unicorne, Lissodendoryx (E.) ramilobosa, Myxodoryx hanitschi and Tedania (T.) tantula. Data and illustrations of some rare or uncommon species are provided: Iophon gaussi, Lissodendoryx (E.) anacantha, Microxina charcoti, Raspailia (H.) hentscheli, Haliclona spongiosissima, Haliclona (G.) cf. cucurbitiformis. Although the present results increase substantially the previously known information on sponge species, this is still an underestimation of total richness because certain orders (Tetractinellida, Suberitida, Biemnida, Polymastiida) and classes were not studied during the current investigation, although previous records were included.
The online World Porifera Database (WPD), the Porifera part of the World Register of Marine Species (WoRMS), lists virtually all published scientific names of sponges. The names of the WPD (as indeed all names in WoRMS) are guided by the Code of the International Comnission on Zoological Nomenclature (ICZN). The WPD names include all currently accepted as well as original combinations, and a majority of non-accepted non-original combinations. Currently, among the accepted names about 200 original lower taxa combinations were found to be accepted by default, usually varieties or formae, which were given that status in the WPD because there was no sufficient published information to support arguments for or against the accepted status. After 1961, varieties and formae are considered infrasubspecific taxa whose names are not regulated by the Code and the names are unavailable, but prior to that date these trinominal taxa are potential available names. It is the purpose of the present study to evaluate these original default ‘accepted’ combinations and arrive at an argumented judgement on whether they are to be truly accepted or non-accepted. Furthermore, additional lower taxa name violations of the Code are also included. Overall, there are three categories of names of lower taxa treated here, (1) with combinations, which are judged to be accepted with elevated status as (sub)specific taxa, (2) with combinations judged to be junior synonyms, and (3) with combinations, which are violating articles of the Code. Among the last category there are a small number of varietal taxa described after the 1960 cut-off date, which are unavailable but are proposed to have the original name combination retained, but as new names with authorship and year changed to those of the present article. Also, names found to be unavailable for various reasons (four-name combinations, junior homonyms, phylocode names) are in this category. The following 31 new names are proposed (three of which are junior synonyms of senior accepted names, and an additional three are unavailable names made available by employing the same name combination with the present authorship and year): Aaptos hoshinoi nom.nov., Ancorina nanosclera nom.nov., Axinella kurushima nom.nov., Callyspongia (Cladochalina) desqueyrouxfaundezae nom.nov., Cliona carpenteri subsp. hentscheli nom.nov., Callyspongia (Toxochalina) gustavoi nom.nov., Craniella microspira nom.nov., Dictyaulus romani nom.nov., Grantia breitfussi nom.nov., Haliclona alba subsp. albapontica nom.nov., Haliclona aquaeductus subsp. sebastopolensis nom.nov., Haliclona inflata subsp. vladimiri nom.nov., Haliclona informis subsp. voldomaroi nom.nov., Haliclona palmata subsp. pontuseuxiniensis nom.nov., Haliclona (Gellius) arthuri nom.nov., Haliclona (Gellius) godthaabae nom.nov., Haliclona merguiensis nom.nov., Haliclona senjitanitai nom.nov., Iophon hentscheli nom.nov., Leucandra wilsoni nom.nov., Paraleucilla bassensis nom.nov. (= P. saccharata), Pione carpenteri subsp. hentscheli nom.nov., Psammocinia samaaii nom.nov., Protoschmidtia czerniavskyi nom.nov. (= Metschnikowia tuberculata), Reiswiginella nom.nov., Scalarispongia lamarcki nom.nov., Spheciospongia hentscheli nom.nov., Spongia (Spongia) vonlendenfeldi nom.nov. (= S. (S.) lignosa), Suberites austral nom.nov., Suberites dendyi nom.nov., Suberites simae nom.nov., and Timea levii nom.nov.
Oscarella malakhovi sp. nov. is described from the waters of Peter the Great Bay (northwestern Sea of Japan, Russia) at depths between 0.4–4 m from a rocky coast. This is the first record of the genus Oscarella from the North-Western Pacific. The new species is characterized by its pinky-beige to yellow color, undulating appearance, lumpy, microlobate surface, soft, slimy consistency, two particular kinds of cells with inclusions (vacuolar and granular cells), and of two kinds of endobiont bacteria. The new species of Oscarella is clearly different in cell, endobiotic bacteria content, and external morphology from all previously described species of Oscarella.
Tetractinellida Marshall 1876 [Borchiellini et al. 2004] comprises the Orders Spirophorida and Astrophorida. A surveyof their diversity in the Bocas del Toro archipelago (Panama) was conducted. A total of ten species of Tetractinellidawere encountered: Cinachyrella alloclada, Cinachyrella apion, Cinachyrella kuekenthali, Ecionemia megastylifera,Stelletta fibrosa, Stelletta sp., Stryphnus raratriaenus sp. nov., Erylus formosus, Geodia gibberosa and Geodiapapyracea. Five of these species are new to the Atlantic sponge fauna of Panama, one of which is new to science.Stryphnus raratriaenus sp. nov. is very similar to Caribbean Asteropus species but it possesses triaenes. It is the firstspecies of this genus in the Caribbean. The description of Ecionemia megastylifera Wintermann–Kilian & Kilian, 1984 ishere revised and the species fully redescribed. A cytochrome c oxidase subunit I (COI) gene partial fragment and/or a 28S ribosomal gene partial fragment (C1–D2 domains) were sequenced for some of the species collected.
New Hexactinellida from the deep Weddell Sea collected in 2005 during the ANT XXII/3 expedition (ANDEEP III ) are described. This hexactinellid fauna includes 12 species from 8 genera, 2 of which are new to science: Lophocalyx profundum n. sp. and L. topsenti n. sp. It further contains the first record of Caulophacus (Caulodiscus) valdiviae in the Weddell Sea, and the second findings of the species Acoelocalyx brucei and Caulophacus (Oxydiscus) weddelli since their original descriptions. The Southern Ocean deep-sea, including the deep Weddell Sea, still offers a challenging potential for the discovery of new taxa.
Thirteen new species of sponges are described from coral reefs of the Netherlands Antilles and the Colombian Caribbean. Species were collected during quantitative investigations of reef sponges performed by students of the University of Amsterdam in the period between 1984 and 1991. Most of the reported specimens were taken from undersides of coral rubble, crevices or reef caves (sciophilous habitats) and without exception are small encrusting or fistular sponges. The material reported in this paper includes a new genus and species of Placospongiidae, Placospherastra antillensis n. g. n. sp. , the first Caribbean representatives of the genera Triptolemma (Pachastrellidae) and Megaciella (Acarnidae), viz. Triptolemma endolithicum n. sp. and Megaciella incrustans n. sp., a new species of Timeidae, Timea curacaoensis n. sp., a new species of Microcionidae with peculiar colloscleres, Clathria (Thalysias) collosclera n. sp., two new species of Chondropsidae, viz. Batzella fusca n. sp., and Strongylacidon unguiferum n. sp., three new species of Coelosphaeridae, viz. Forcepia (Forcepia) minima n. sp., Forcepia (Forcepia) fistulosa n. sp., and Forcepia (Leptolabis) microlabis n. sp., a new species of Crellidae, Crella (Grayella) beglingerae n. sp., a new species of Hymedesmiidae, Hymedesmia (Hymedesmia) bonairensis n. sp., and a new species of Mycalidae, Mycale (Paresperella) vitellina n. sp. Most species are represented by only small fragments removed from the substrate by scalpel or diving knife, leaving little and often crumbled preserved type material. This study is intended to demonstrate that the small crusts dominating easily accessible shallow water coral rubble habitats in the Caribbean remain understudied.
A common Mediterranean species of Phorbas has been frequently identified under the specific name paupertas, referring to the sponge described as Hymeniacidon paupertas by Bowerbank 1866 from the north coasts of Europe. It is actually different from this species, and a new name, Phorbas topsenti sp.nov., is proposed. A description of the morphology, spicule complement and cytology is given. Another Mediterranean species of Phorbas, Phorbas tailliezi sp. nov. is described.
The Antarctic deep-water fauna of Polymastiidae and Suberitidae is revised using recently collected material from the Weddell Sea. The former family appeared to be more abundant and diverse than the latter family in the studied area. Seven species within five polymastiid genera and three species within three suberitid genera are described. Relatively high sponge abundance at two stations deeper than 4700 m was mainly constituted by a polymastiid species Radiella Antarctica sp. nov. Previously, representatives of Radiella have never been found in the Antarctic. An eurybathic species, Polymastia invaginata, well known from the Antarctic and subantarctic, appeared to be especially abundant at less than 1000 m depth. Another eurybathic polymastiid species, Tentorium cf. semisuberites, known for its bipolar distribution, was the third abundant species at the depths between 1000–2600 m, with the highest density found at the deeper stations. Tentorium papillatum, endemic of the Southern Hemisphere, was registered only at a depth of about 1000 m. Other species studied were less abundant. Astrotylus astrotylus, the representative of the endemic Antarctic genus, was found exclusively deeper than 4500 m, often together with R. antarctica. Acanthopolymastia acanthoxa, the endemic deepwater Antarctic species, was registered at 3000 m. The discovery of suberitid Aaptos robustus sp. nov. at about 2300 m is the first signalization of Aaptos in the Antarctic and at such a considerable depth. The finding of Suberites topsenti deeper than 4700 m is also remarkable. In general the results achieved confirm the high degree of geographical endemism of the Antarctic deep-water sponge fauna and the eurybathic distribution of many Antarctic sponge species.
Discovery of new specimens of Crateromorpha (Neopsacas), which are attributed to new species, from the Pacific, Antarctic and Indian Oceans and the description of new specimen also from the Atlantic Ocean allow us to review the skeletal characters regarding the dermal-hypodermal-atrial-hypoatrial spicule composition in various species of subgenus C. (Neopsacas). A new diagnosis of this taxon is proposed, and several new species and subspecies described: Crateromorpha (Neopsacas) krinovi krinovi sp. nov., ssp. nov., Crateromorpha (Neopsacas) krinovi discoli ssp. nov. and Crateromorpha (Neopsacas) obi sp. nov. The status of the subgenus Neopsacas within the Rossellidae and its close relation to Caulophacus are now well established according to morphological characters. The distribution of this subgenus is wide: Atlantic, Pacific, Indian and Antarctic Oceans. Possible phylogenetic affinities between the genera Crateromorpha and Caulophacus are discussed.
A revision of Brazilian Aplysina was conducted based on morphological study of species alive as well as preserved in the collections of Museu Nacional. In total, 190 specimens were studied in the lab. Identification was achieved by analysis of the external morphology, in combination with skeletal architecture and its components. Our analyses revealed marked within individual variation for most species in characters associated to the spongin fibres, suggesting a low diagnostic strength for this set of characters in Aplysina. External morphology, on the other hand, permitted clearcut diagnoses for most species. Fifteen species were identified and are fully described below. Eight of these are new to science. A neotype for A. fulva is proposed here. The amended list of Aplysina species for the Brazilian coast is: A. alcicornis sp.n., A. caissara, A. cauliformis, A. cristagallus sp.n, A. fistularis, A. fulva, A. insularis, A. lacunosa, A. lactuca sp.n., A. lingua sp.n., A. muricyana sp.n., A. orthoreticulata sp.n., A. pergamentacea, A. pseudolacunosa sp.n. and A. solangeae sp.n. A tentative identification key for these species is provided.
The sponge Tethya leysae sp. nov. (Porifera, Demospongiae, Hadromerida, Tethyidae) is a new species from the Canadian Northeast Pacific. Its type locality is the infralittoral around Ohiat Islet, Barkely Sound, Vancouver Island in Canada. Tethya leysae sp. nov. is a medium-sized spherical sponge with a verrucose surface, orange-yellow to light red alive and white with a greyish core in ethanol. The highly variable oxyspherasters (25-115 µm in size, R/C 0.34-0.69) are densely scattered almost throughout the entire cortex. The micrasters comprise acanthoxyspherasters, acanthostrongylasters, acanthotylasters and small oxyspherasters, which are present at low densities throughout the sponge and form a dense layer associated with the exopinacoderm. The megascleres and the auxiliary megascleres comprise oxeas and strongyloxeas. The new species is clearly distinguishable from the closely related T. californiana by the absence of spherules among the micrasters, the lack of an alveolar cortex and the extremely high density of megasters in the cortex. In addition, the R/C values of the megasters differ between the two species and the oxyspherasters of T. leysae sp. nov. rarely display bent rays. The morphological differences between the species are confirmed by nucleotide and amino acid substitutions within the cytochrome oxidase subunit I (COI) mtDNA gene. Phylogenetic analyses group T. leysae sp. nov. with T. californiana, T. actinia and T. minuta, which together form a sister group to a MediterraneanNorth Atlantic species cluster. Morphological analyses of the skeleton included x-ray microtomography (µCT) and virtual 3D reconstruction, which was used for the first time in conjunction with the description of a new sponge species. Microtomography permitted the visualization and analysis of spicules within the skeletal context or isolated in silico. The method represents a valuable extension to the sponge taxonomist’s toolbox since it allows morphometric measurements in 3D. µCT will thus supplement classical morphological methods such as light and scanning electron microscopy.