Macrobiotus azzunae Marnissi & Cesari & Rebecchi & Bertolani 2021, sp. nov.
Macrobiotus azzunae sp. nov. urn:lsid:zoobank.org:act: 933CCC06-F69D-49E2-AF4F-0C042D8F5C99 Figs 1–4, 5A, C, 7 Etymology The new species is dedicated in honor of Atf Azzouna, professor in the Faculty of Mathematical, Physical and Natural Sciences of Tunis and supervisor of the PhD thesis of Jamila B...
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2021
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Online Access: | https://dx.doi.org/10.5281/zenodo.5092922 https://zenodo.org/record/5092922 |
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Biodiversity Taxonomy Animalia Tardigrada Eutardigrada Parachela Macrobiotidae Macrobiotus Macrobiotus azzunae |
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Biodiversity Taxonomy Animalia Tardigrada Eutardigrada Parachela Macrobiotidae Macrobiotus Macrobiotus azzunae Marnissi, Jamila Ben Cesari, Michele Rebecchi, Lorena Bertolani, Roberto Macrobiotus azzunae Marnissi & Cesari & Rebecchi & Bertolani 2021, sp. nov. |
topic_facet |
Biodiversity Taxonomy Animalia Tardigrada Eutardigrada Parachela Macrobiotidae Macrobiotus Macrobiotus azzunae |
description |
Macrobiotus azzunae sp. nov. urn:lsid:zoobank.org:act: 933CCC06-F69D-49E2-AF4F-0C042D8F5C99 Figs 1–4, 5A, C, 7 Etymology The new species is dedicated in honor of Atf Azzouna, professor in the Faculty of Mathematical, Physical and Natural Sciences of Tunis and supervisor of the PhD thesis of Jamila Ben Marnissi. Type material Holotype TUNISIA • spec. of unidentified sex; North-West Tunisia, Kroumiri Mountains, Ain Soltan forest, Jendouba; 36º31′21.788″ N, 8º19′57.741″ E; 893 m a.s.l.; Apr. 2017; Marnissi leg.; moss on trunk of Quercus canariensis UNIMORE, slide code C4218–S32. Paratypes TUNISIA • 17 specs, sex unidentified; same collection data as for holotype; UNIMORE, slide codes C4218–S2 to C4218–S7, C4218–S9, C4218–S17, C4218–S30, C4218–S31, C4218–S33 to C4218– S35 • 3 eggs; same collection data as for holotype; UNIMORE, slide codes C4218–S10, C4218–S11, C4218–S25. Type depositories The holotype (slide: C 4218– S 32), 17 paratypes (slides C 4218– S 2 to C 4218– S 7, C 4218– S 17, C 4128– S 30, C 4218– S 31, C 4218– S 33 to C 4218- S 35), 3 eggs (slides: C 4218– S 10/11/25) and two vouchers (slides SP 04 and SP 07, corresponding to specimens C 4218 V 4 and C 4218 V 7, respectively) mounted in Faure-Berlese fluid, are deposited in the Bertolani collection at the Department of Life Science, UNIMORE, Modena, Italy. Type locality NW Tunisia, Kroumrie mountains, Ain Soltan forest, Jendouba, 36º31′21.788″ N, 8º19′57.741″ E. Altitude 893 m a.s.l. Description Adult specimens Body white, transparent after mounting in Faure-Berlese, from 162.2 to 410.3 µm in length (Fig. 2A, Table 3; structures measured only in the animals more than 200 µm in length). Eye spots present, even after mounting. Cuticle smooth but with small round or oval pores, 1–1.5 µm in diameter (Fig. 2B), better visible after fixation in Carnoy and orcein staining (Fig. 3C), scattered randomly on the entire cuticle, including the dorsal surface of all legs. With SEM, pores look oval or in the shape of a seed (Fig. 3A, D) with the largest diameter of 0.7–0.8 µm. Weak cuticular granulation also present on the lateral surface of all legs and specially on legs IV (Fig. 2B, arrow). Only with SEM is it possible to define the shape of the granulation on the legs, which looks as a regular disposition of star-shaped protuberances (about 0.3 µm; Fig. 3F). Six buccal sensory lobes around the mouth, well recognizable with SEM. Mouth antero-ventral; buccal-pharyngeal apparatus of the Macrobiotus type (sensu Pilato & Binda 2010), with ventral lamina and ten small peribuccal lamellae (in the holotype, after mounting, separated from the mouth). Buccal armature, corresponding to oral cavity armature, OCA, according to Michalczyk & Kaczmarek (2003), without an anterior band of teeth visible, corresponding to the first band of teeth according to Michalczyk & Kaczmarek (2003), and to the anterior band of the buccal ring according to Guidetti et al . (2012); posterior band of teeth poorly visible, corresponding to second band of teeth, according to Michalczyk & Kaczmarek (2003), followed by three dorsal and three ventral crests, corresponding to third band of teeth according to Michalczyk & Kaczmarek (2003); the dorsal crests (Fig. 2D) are distinct transverse ridges, whereas the ventral crests (Fig. 2E) appear as two separate lateral transverse ridges and a roundish median tooth. The posterior band of teeth and the transverse ridges are part of the buccal tube, according to Guidetti et al . (2012). Buccal tube narrow; pharyngeal bulb spherical with triangular apophyses, two rod-shaped macroplacoids, relatively short, the first longer than the second and evidently but not deeply narrowed at its middle (Fig. 2C), the second with a not particularly evident subterminal constriction. Microplacoid present. Slender claws of the hufelandi type (sensu Pilato & Binda 2010); the external claw longer than the internal one and the posterior longer than the anterior. Primary branches of each claw with distinct accessory points (Fig. 2F), a common tract of medium length (about a third of the total claw length) and an evident stalk connecting the claw to the lunule. Lunules under all claws, smooth, larger on the hind legs (Figs 2G, 3E). Cuticular bars under claws absent. The population is dioecious (gonochoristic). Males were recognized using orcein staining, which revealed that the testis is filled with spermatozoa with a coiled head (Fig. 3G) and spermatids. No morphological secondary sexual dimorphism, such as gibbosities on legs IV in males, was identified. Eggs Eggs are laid freely, and are white, spherical or slightly oval. One egg containing a fully developed embryo showed the shape of the buccal-pharyngeal apparatus (Fig. 4A). Processes of the eggshell are in the shape of inverted goblets (Fig. 4B) with conical trunks and well-defined distal discs as large as the process bases (for measurements see Table 4). Distal discs concave, with a median small protuberance and, using PhC, with border often smooth, or sometimes slightly jagged, or slightly ragged (Fig. 4C), but never clearly jagged, serrated or dentate. Surface among processes of the hufelandi type (sensu Kaczmarek & Michalczyk 2017a), i.e., covered by a very thin grid (Fig. 4D). Meshes around the process bases slightly larger and with slightly thicker wires compared with interbasal meshes. Mesh diameter around 0.5 µm. Comparisons Macrobiotus azzunae sp. nov. has eggs with processes as inverted goblets and a reticulate eggshell between the processes. Consequently, a comparison must be done with the Macrobiotus species listed by Kaczmarek & Michalczyk (2017a) with hufelandi type eggshells, excluding the species with processes that are not like inverted goblets, and adding the species with hufelandi type chorion eggs described after that publication. The species with hufelandi type chorion eggs that do not have processes as inverted goblets are Macrobiotus acadianus (Meyer & Domingue, 2011), M. dariae Pilato & Bertolani, 2004, M. lissostomus Durante Pasa & Maucci, 1979, M. santoroi Pilato & D’Urso, 1976, and M. scoticus Stec, Morek, Gąsiorek, Blagden & Michalczyk, 2017. Moreover, M. azzunae sp. nov. has egg processes with distal discs with a smooth or slightly jagged border, therefore it differs from all the species that have clearly indented, serrated or clearly jagged distal discs, such as: Macrobiotus canaricus Stec, Krzywański & Michalczyk, 2018, M. crustulus Stec, Dudziak & Michalczyk, 2020, M. hannae Nowak & Stec, 2018 (whose egg surface is more cribriform than reticulate), M. hibiscus de Barros, 1942, M. horningi Kaczmarek & Michalczyk, 2017b (which also has very high processes), M. hufelandi C.A.S. Schultze, 1834, M. humilis Binda & Pilato, 2001, M. iharosi Pilato, Binda & Catanzaro 1991, M. joannae Pilato & Binda, 1983, M. julianae (Meyer, 2012), M. kamilae Coughlan & Stec, 2019, M. modestus Pilato & Lisi, 2009, M. noonragi s Coughlan & Stec, 2019, M. papei Stec, Kristensen & Michalczyk, 2018 (with particularly long filaments starting from the distal disc), M. paulinae Stec, Smolak, Kaczmarek & Michalczyk, 2015, M. polypiformis Roszkowska, Ostrowska, Stec, Janko & Kaczmarek, 2017 (even with cog-teeth extended to form a long, thin, hair-like and flexible filament), M. punctillus Pilato, Binda & Azzaro, 1990, M. sapiens Binda & Pilato, 1984, M. sottilei Pilato, Kiosya, Lisi & Sabella, 2012. For the shape of the egg Macrobiotus azzunae sp. nov. differs from M. rawsoni Horning, Schuster & Grigarick, 1978 because this species has only one strip of meshes around each egg process (see Kaczmarek & Michalczyk 2017b); it differs from M. serratus Bertolani, Guidi & Rebecchi, 1996 because in this species the egg surface is porous more than reticulated, with pores small and spaced from each other, and its egg processes have a large, often square, distal disc; it differs from M. seychellensis Biserov, 1994 because the distal disc of the egg processes of this species, even though not dentate, has long and very developed lobes. The remaining nine species of the hufelandi group should be compared singularly. Macrobiotus almadai Fontoura, Pilato & Lisi, 2008 Macrobiotus azzunae sp. nov. differs from M. almadai in having a posterior band of teeth in the buccal cavity visible with LM (not visible in M. almadai ), and distal disc with a jagged margin instead of very small teeth as in M. almadai . Macrobiotus canaricus Stec, Krzywański & Michalczyk, 2018 With LM the margin of the distal disc of M. azzunae sp. nov., never dentate in this species, looks similar to that of M. canaricus , but the SEM images of the eggs of the latter species evidence the presence of an almost dentate disc. Moreover, the peribasal meshes of the eggshell are larger than interbasal ones in the new species while they do not differ from the interbasal ones in M. canaricus regarding the animals there are differences in the buccal armature: in M. azzunae sp. nov. the posterior band of teeth is visible with LM (even though poorly) and the three dorsal crests are distinct transverse ridges, while in M. canaricus the posterior band of teeth is visible only with SEM and with LM the dorsal teeth form a transversal ridge weakly divided into three teeth. Macrobiotus madegassus Maucci, 1993 The new species differs from M. madegassus by the presence of the eye spots (absent in M. madegassus ), pores on the cuticle (absent in M. madegassus ), presence in the buccal armature of posterior band of teeth, even though weak (fully absent in M. madegassus ), buccal tube much larger ( pt of the holotypes 15.9 vs 7), insertion of the stylet supports on the buccal tube much more posterior ( pt of the holotypes 76.1 vs 68), first and second macroplacoid longer ( pt of the holotypes 25.5 and 18.1 vs 21.3 and 12.0), lunules on the hind legs without kerning (crenate in M. madegassus ), eggshell processes with distal disc as large as the base (similar range 3.2–5.2 for both measurements) with respect to that of M. madegassus (disc vs base: 4.3–5.4 vs 2.3–2.6). Macrobiotus martini Bartels , Pilato, Lisi & Nelson, 2009 The cuticular pores in M. azzunae sp. nov. are much smaller than those of M. martini the distal disc of the egg processes in M. azzunae sp. nov. has a diameter similar to that of the process base, while in M. martini the distal disc is much narrower than the base. Macrobiotus nebrodensis Pilato, Sabella, D’Urso & Lisi, 2017 Macrobiotus azzunae sp. nov. differs from M. nebrodensis by the absence of the cuticular bar near the lunules on the first three pairs of legs (a faint bar is present in M. nebrodensis ). The egg processes of M. azzuane sp. nov. are in higher number on the circumference (29–33) with respect to those of M. nebrodensis (18). In the latter species there are some egg processes very high (up to 20.6 µm), while in the new species process height and shape are more uniform. The difference in the eggshell between meshes around the process base and the others is much less evident in M. azzunae sp. nov. than in M. nebrodensis . Macrobiotus personatus Biserov, 1990 The new species differs from M. personatus by the posterior band of the buccal armature less evident, the presence of a clear constriction in the first macroplacoid (Fig. 5A), in the paratype of M. personatus examined by us barely identifiable (Fig. 6A) and, according to Biserov (1990) usually absent in the type material of that species. The pores on the cuticle of M. azzunae sp. nov. are small, approximately 1 µm in diameter, while in M. personatus they are strongly elliptic and about 3 µm in length (Fig. 6B). Lunules on leg IV are always smooth in M. azzunae sp. nov., sometimes indented in M personatus . With respect to the eggs of M. personatus (Fig. 6C–D), the egg processes of M. azzunae sp. nov. (Figs 4C–D, 5C) are clearly shorter, 5.4 ± 0.6 vs 9.5 ± 0.5 (range 4.2–6.4 vs 9–10.5) and with a narrower base and distal disc (both 3.2–5.2 vs 7–10.5 and 7–9 respectively). Males are present in the new species, while in M. personatus only females were found (Biserov 1990), suggesting parthenogenesis in that species. Macrobiotus sandrae Bertolani & Rebecchi, 1993 The new species differs from M. sandrae for the eggshell shape, with thinner wires of the reticulum and meshes around the processes larger than the inter-process meshes in M. azzunae sp. nov. (Fig. 5C), all meshes similar in size in M. sandrae (Fig. 5D). Figure 5C–D also show a difference in the process base diameter, narrower in M. azzunae sp. nov. With regard to the animals, M. azzunae sp. nov. differs from M. sandrae by a constriction of the first macroplacoid much more pronounced (Fig. 5A; it is hardly visible in M. sandrae Fig. 5B). Moreover, in animals of similar size the posterior band of the buccal armature is just less evident in the new species, and lunules on the hind legs are without hint of teeth (but teeth, present in the holotype of M. sandrae , are often difficult to identify in other specimens of that species). Macrobiotus terminalis Bertolani & Rebecchi, 1993 Macrobiotus azzunae sp. nov. differs from M. terminalis for the absence of granulation on the cuticle (noted only in the redescription of M. terminalis see Cesari et al . 2011), for the absence of teeth on the lunules, especially evident on the hind legs of M. terminalis , and for the presence of males, absent in M. terminalis (see redescription by Cesari et al . 2011). Macrobiotus vladimiri Bertolani, Biserov, Rebecchi & Cesari, 2011 With respect to M. vladimiri , animals of M. azzunae sp. nov. reach a shorter length (up to 410.3 µm vs 515.1 µm), in M. azzunae sp. nov. the posterior band of teeth of the buccal armature is less evident and the lunules on the hind legs are not indented. In M. azzunae sp. nov. the egg diameter without processes (64.7–80.6 µm) is less than that of the eggs of M. vladimiri (89.9–92.0 µm); the processes are shorter (4.2–6.4 µm in the new species vs 6.5–8 µm in M. vladimiri ). In the new species the base process diameter is narrower (3.2–5.2 µm) than in M. vladimiri (5.1–7.3 µm), the distal disc is weakly or not jagged (clearly jagged in M. terminalis ). In M. azzunae sp. nov. males are present, while they are absent in M . vladimiri . Genetic distances The ranges of uncorrected genetic p-distances between M. azzunae sp. nov. and the other species of the M. hufelandi group (Supp. file 7, Supp. file 8, Supp. file 9, Supp. file 10), are as follows: 18S 0.1–5.6%, with the most similar being M. sandrae from Germany (present paper) 28S 0.1%, with the only available M. vladimiri from Spain (FJ435751 –5) ITS-2 7.7–32.2%, with the most similar being Macrobiotus vladimiri (MN888347) from Finland COI 6.3–25.6%, with the most similar being Macrobiotus sandrae (HQ876574, HQ876577, HQ876578, HQ876579, HQ876581) from Germany The COI dataset is the most complete and informative for species delimitation investigation. Both phylogenetic reconstructions on the COI dataset resulted in the same topology, and thus the ML tree was utilized for the PTP analysis (Fig. 7, left), which shows 14 putative species clusters: M. crustulus , M.hannae , M. cf. recens , M.canaricus , M.hufelandi , M. cf. hufelandi sp.1, M. terminalis , M. cf. terminalis , M. wandae , M. macrocalix , M . cf. macrocalix , M. vladimiri , M. sandrae and M. azzunae sp. nov. This subdivision is further validated by both the ABGD and the haplotype network analysis (Fig. 7, centre and right). Present molecular data therefore confirms the validity of the erection of M. azzunae sp. nov. : Published as part of Marnissi, Jamila Ben, Cesari, Michele, Rebecchi, Lorena & Bertolani, Roberto, 2021, Integrative description of a new Tunisian tardigrade species, Macrobiotus azzunae sp. nov. 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Macrobiotus Hufelandii animal e crustaceorum classe novum, reviviscendi post diuturnam asphyxia et ariditaten potens, etc. C. Curths, Berlin.", "Pilato G., Binda M. G. & Catanzaro R. 1991. Remarks on some tardigrades of the African fauna with the description of three new species of Macrobiotus Schultze 1834. Tropical Zoology 4: 167 - 178. https: // doi. org / 10.1080 / 03946975.1991.10539487", "Pilato G. & Lisi O. 2009. Tardigrades of the Seychelles Islands, with the description of three new species. Zootaxa 2124: 1 - 20. https: // doi. org / 10.5281 / zenodo. 188178", "Pilato G., Binda M. G. & Azzaro M. 1990. Tardigradi di Terra del Fuoco e Magallanes. III. Macrobiotus punctillus, nuova specie di Macrobiotidae del gruppo hufelandi. Animalia 17: 123 - 129.", "Pilato G., Kiosya Ye., Lisi O. & Sabella G. 2012. New records of Eutardigrada from Belarus with the description of three new species. Zootaxa 3179: 39 - 60. https: // doi. org / 10.11646 / zootaxa. 3179.1.2 Pilato G., Sabella G., D'Urso V. & Lisi O. 2017. Macrobiotus nebrodensis and Adropion vexatum, two new species of Eutardigrada (Tardigrada, Parachela) from Sicily. Zootaxa 4362 (2): 267 - 279. https: // doi. org / 10.11646 / zootaxa. 4362.2.6", "Biserov V. I. 1990. On the revision of the Macrobiotus genus. The subgenus Macrobiotus s. str. is a new taxonomic status of the hufelandi group (Tardigrada, Macrobiotidae). Communication 2. Zoologicheskij Zhurnal 69: 38 - 50. [In Russian.]"]} |
format |
Text |
author |
Marnissi, Jamila Ben Cesari, Michele Rebecchi, Lorena Bertolani, Roberto |
author_facet |
Marnissi, Jamila Ben Cesari, Michele Rebecchi, Lorena Bertolani, Roberto |
author_sort |
Marnissi, Jamila Ben |
title |
Macrobiotus azzunae Marnissi & Cesari & Rebecchi & Bertolani 2021, sp. nov. |
title_short |
Macrobiotus azzunae Marnissi & Cesari & Rebecchi & Bertolani 2021, sp. nov. |
title_full |
Macrobiotus azzunae Marnissi & Cesari & Rebecchi & Bertolani 2021, sp. nov. |
title_fullStr |
Macrobiotus azzunae Marnissi & Cesari & Rebecchi & Bertolani 2021, sp. nov. |
title_full_unstemmed |
Macrobiotus azzunae Marnissi & Cesari & Rebecchi & Bertolani 2021, sp. nov. |
title_sort |
macrobiotus azzunae marnissi & cesari & rebecchi & bertolani 2021, sp. nov. |
publisher |
Zenodo |
publishDate |
2021 |
url |
https://dx.doi.org/10.5281/zenodo.5092922 https://zenodo.org/record/5092922 |
long_lat |
ENVELOPE(-62.933,-62.933,-64.883,-64.883) ENVELOPE(26.733,26.733,67.850,67.850) ENVELOPE(-65.683,-65.683,-65.967,-65.967) ENVELOPE(-159.667,-159.667,-86.333,-86.333) ENVELOPE(-62.167,-62.167,-64.717,-64.717) ENVELOPE(-57.917,-57.917,-61.950,-61.950) ENVELOPE(-68.871,-68.871,-68.093,-68.093) ENVELOPE(168.983,168.983,-77.517,-77.517) ENVELOPE(-57.967,-57.967,-63.550,-63.550) |
geographic |
Greenland Pacific Magallanes Pasa Jagged Kristensen Barros Nowak Faure Tooth The Giovannini |
geographic_facet |
Greenland Pacific Magallanes Pasa Jagged Kristensen Barros Nowak Faure Tooth The Giovannini |
genre |
Greenland Tardigrade |
genre_facet |
Greenland Tardigrade |
op_relation |
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Open Access Creative Commons Zero v1.0 Universal https://creativecommons.org/publicdomain/zero/1.0/legalcode cc0-1.0 info:eu-repo/semantics/openAccess |
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https://doi.org/10.5281/zenodo.5092922 https://doi.org/10.5852/ejt.2021.758.1429 https://doi.org/10.5281/zenodo.5088107 https://doi.org/10.5281/zenodo.5088109 https://doi.org/10.5281/zenodo.5088111 https://doi.org/10.5281/zenodo.5088113 https: |
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ftdatacite:10.5281/zenodo.5092922 2023-05-15T16:30:55+02:00 Macrobiotus azzunae Marnissi & Cesari & Rebecchi & Bertolani 2021, sp. nov. Marnissi, Jamila Ben Cesari, Michele Rebecchi, Lorena Bertolani, Roberto 2021 https://dx.doi.org/10.5281/zenodo.5092922 https://zenodo.org/record/5092922 unknown Zenodo http://zenodo.org/record/5088105 http://publication.plazi.org/id/EF4455697B12FFE5FFF49442FFBEDA0B http://zoobank.org/C4F57584-0AD9-47C9-BA95-02133C434941 https://zenodo.org/communities/biosyslit https://dx.doi.org/10.5852/ejt.2021.758.1429 http://zenodo.org/record/5088105 http://publication.plazi.org/id/EF4455697B12FFE5FFF49442FFBEDA0B https://dx.doi.org/10.5281/zenodo.5088107 https://dx.doi.org/10.5281/zenodo.5088109 https://dx.doi.org/10.5281/zenodo.5088111 https://dx.doi.org/10.5281/zenodo.5088113 https://dx.doi.org/10.5281/zenodo.5088115 https://dx.doi.org/10.5281/zenodo.5088121 https://dx.doi.org/10.5281/zenodo.5088117 http://zoobank.org/C4F57584-0AD9-47C9-BA95-02133C434941 https://dx.doi.org/10.5281/zenodo.5092921 https://zenodo.org/communities/biosyslit Open Access Creative Commons Zero v1.0 Universal https://creativecommons.org/publicdomain/zero/1.0/legalcode cc0-1.0 info:eu-repo/semantics/openAccess CC0 Biodiversity Taxonomy Animalia Tardigrada Eutardigrada Parachela Macrobiotidae Macrobiotus Macrobiotus azzunae Text Taxonomic treatment article-journal ScholarlyArticle 2021 ftdatacite https://doi.org/10.5281/zenodo.5092922 https://doi.org/10.5852/ejt.2021.758.1429 https://doi.org/10.5281/zenodo.5088107 https://doi.org/10.5281/zenodo.5088109 https://doi.org/10.5281/zenodo.5088111 https://doi.org/10.5281/zenodo.5088113 https: 2021-11-05T12:55:41Z Macrobiotus azzunae sp. nov. urn:lsid:zoobank.org:act: 933CCC06-F69D-49E2-AF4F-0C042D8F5C99 Figs 1–4, 5A, C, 7 Etymology The new species is dedicated in honor of Atf Azzouna, professor in the Faculty of Mathematical, Physical and Natural Sciences of Tunis and supervisor of the PhD thesis of Jamila Ben Marnissi. Type material Holotype TUNISIA • spec. of unidentified sex; North-West Tunisia, Kroumiri Mountains, Ain Soltan forest, Jendouba; 36º31′21.788″ N, 8º19′57.741″ E; 893 m a.s.l.; Apr. 2017; Marnissi leg.; moss on trunk of Quercus canariensis UNIMORE, slide code C4218–S32. Paratypes TUNISIA • 17 specs, sex unidentified; same collection data as for holotype; UNIMORE, slide codes C4218–S2 to C4218–S7, C4218–S9, C4218–S17, C4218–S30, C4218–S31, C4218–S33 to C4218– S35 • 3 eggs; same collection data as for holotype; UNIMORE, slide codes C4218–S10, C4218–S11, C4218–S25. Type depositories The holotype (slide: C 4218– S 32), 17 paratypes (slides C 4218– S 2 to C 4218– S 7, C 4218– S 17, C 4128– S 30, C 4218– S 31, C 4218– S 33 to C 4218- S 35), 3 eggs (slides: C 4218– S 10/11/25) and two vouchers (slides SP 04 and SP 07, corresponding to specimens C 4218 V 4 and C 4218 V 7, respectively) mounted in Faure-Berlese fluid, are deposited in the Bertolani collection at the Department of Life Science, UNIMORE, Modena, Italy. Type locality NW Tunisia, Kroumrie mountains, Ain Soltan forest, Jendouba, 36º31′21.788″ N, 8º19′57.741″ E. Altitude 893 m a.s.l. Description Adult specimens Body white, transparent after mounting in Faure-Berlese, from 162.2 to 410.3 µm in length (Fig. 2A, Table 3; structures measured only in the animals more than 200 µm in length). Eye spots present, even after mounting. Cuticle smooth but with small round or oval pores, 1–1.5 µm in diameter (Fig. 2B), better visible after fixation in Carnoy and orcein staining (Fig. 3C), scattered randomly on the entire cuticle, including the dorsal surface of all legs. With SEM, pores look oval or in the shape of a seed (Fig. 3A, D) with the largest diameter of 0.7–0.8 µm. Weak cuticular granulation also present on the lateral surface of all legs and specially on legs IV (Fig. 2B, arrow). Only with SEM is it possible to define the shape of the granulation on the legs, which looks as a regular disposition of star-shaped protuberances (about 0.3 µm; Fig. 3F). Six buccal sensory lobes around the mouth, well recognizable with SEM. Mouth antero-ventral; buccal-pharyngeal apparatus of the Macrobiotus type (sensu Pilato & Binda 2010), with ventral lamina and ten small peribuccal lamellae (in the holotype, after mounting, separated from the mouth). Buccal armature, corresponding to oral cavity armature, OCA, according to Michalczyk & Kaczmarek (2003), without an anterior band of teeth visible, corresponding to the first band of teeth according to Michalczyk & Kaczmarek (2003), and to the anterior band of the buccal ring according to Guidetti et al . (2012); posterior band of teeth poorly visible, corresponding to second band of teeth, according to Michalczyk & Kaczmarek (2003), followed by three dorsal and three ventral crests, corresponding to third band of teeth according to Michalczyk & Kaczmarek (2003); the dorsal crests (Fig. 2D) are distinct transverse ridges, whereas the ventral crests (Fig. 2E) appear as two separate lateral transverse ridges and a roundish median tooth. The posterior band of teeth and the transverse ridges are part of the buccal tube, according to Guidetti et al . (2012). Buccal tube narrow; pharyngeal bulb spherical with triangular apophyses, two rod-shaped macroplacoids, relatively short, the first longer than the second and evidently but not deeply narrowed at its middle (Fig. 2C), the second with a not particularly evident subterminal constriction. Microplacoid present. Slender claws of the hufelandi type (sensu Pilato & Binda 2010); the external claw longer than the internal one and the posterior longer than the anterior. Primary branches of each claw with distinct accessory points (Fig. 2F), a common tract of medium length (about a third of the total claw length) and an evident stalk connecting the claw to the lunule. Lunules under all claws, smooth, larger on the hind legs (Figs 2G, 3E). Cuticular bars under claws absent. The population is dioecious (gonochoristic). Males were recognized using orcein staining, which revealed that the testis is filled with spermatozoa with a coiled head (Fig. 3G) and spermatids. No morphological secondary sexual dimorphism, such as gibbosities on legs IV in males, was identified. Eggs Eggs are laid freely, and are white, spherical or slightly oval. One egg containing a fully developed embryo showed the shape of the buccal-pharyngeal apparatus (Fig. 4A). Processes of the eggshell are in the shape of inverted goblets (Fig. 4B) with conical trunks and well-defined distal discs as large as the process bases (for measurements see Table 4). Distal discs concave, with a median small protuberance and, using PhC, with border often smooth, or sometimes slightly jagged, or slightly ragged (Fig. 4C), but never clearly jagged, serrated or dentate. Surface among processes of the hufelandi type (sensu Kaczmarek & Michalczyk 2017a), i.e., covered by a very thin grid (Fig. 4D). Meshes around the process bases slightly larger and with slightly thicker wires compared with interbasal meshes. Mesh diameter around 0.5 µm. Comparisons Macrobiotus azzunae sp. nov. has eggs with processes as inverted goblets and a reticulate eggshell between the processes. Consequently, a comparison must be done with the Macrobiotus species listed by Kaczmarek & Michalczyk (2017a) with hufelandi type eggshells, excluding the species with processes that are not like inverted goblets, and adding the species with hufelandi type chorion eggs described after that publication. The species with hufelandi type chorion eggs that do not have processes as inverted goblets are Macrobiotus acadianus (Meyer & Domingue, 2011), M. dariae Pilato & Bertolani, 2004, M. lissostomus Durante Pasa & Maucci, 1979, M. santoroi Pilato & D’Urso, 1976, and M. scoticus Stec, Morek, Gąsiorek, Blagden & Michalczyk, 2017. Moreover, M. azzunae sp. nov. has egg processes with distal discs with a smooth or slightly jagged border, therefore it differs from all the species that have clearly indented, serrated or clearly jagged distal discs, such as: Macrobiotus canaricus Stec, Krzywański & Michalczyk, 2018, M. crustulus Stec, Dudziak & Michalczyk, 2020, M. hannae Nowak & Stec, 2018 (whose egg surface is more cribriform than reticulate), M. hibiscus de Barros, 1942, M. horningi Kaczmarek & Michalczyk, 2017b (which also has very high processes), M. hufelandi C.A.S. Schultze, 1834, M. humilis Binda & Pilato, 2001, M. iharosi Pilato, Binda & Catanzaro 1991, M. joannae Pilato & Binda, 1983, M. julianae (Meyer, 2012), M. kamilae Coughlan & Stec, 2019, M. modestus Pilato & Lisi, 2009, M. noonragi s Coughlan & Stec, 2019, M. papei Stec, Kristensen & Michalczyk, 2018 (with particularly long filaments starting from the distal disc), M. paulinae Stec, Smolak, Kaczmarek & Michalczyk, 2015, M. polypiformis Roszkowska, Ostrowska, Stec, Janko & Kaczmarek, 2017 (even with cog-teeth extended to form a long, thin, hair-like and flexible filament), M. punctillus Pilato, Binda & Azzaro, 1990, M. sapiens Binda & Pilato, 1984, M. sottilei Pilato, Kiosya, Lisi & Sabella, 2012. For the shape of the egg Macrobiotus azzunae sp. nov. differs from M. rawsoni Horning, Schuster & Grigarick, 1978 because this species has only one strip of meshes around each egg process (see Kaczmarek & Michalczyk 2017b); it differs from M. serratus Bertolani, Guidi & Rebecchi, 1996 because in this species the egg surface is porous more than reticulated, with pores small and spaced from each other, and its egg processes have a large, often square, distal disc; it differs from M. seychellensis Biserov, 1994 because the distal disc of the egg processes of this species, even though not dentate, has long and very developed lobes. The remaining nine species of the hufelandi group should be compared singularly. Macrobiotus almadai Fontoura, Pilato & Lisi, 2008 Macrobiotus azzunae sp. nov. differs from M. almadai in having a posterior band of teeth in the buccal cavity visible with LM (not visible in M. almadai ), and distal disc with a jagged margin instead of very small teeth as in M. almadai . Macrobiotus canaricus Stec, Krzywański & Michalczyk, 2018 With LM the margin of the distal disc of M. azzunae sp. nov., never dentate in this species, looks similar to that of M. canaricus , but the SEM images of the eggs of the latter species evidence the presence of an almost dentate disc. Moreover, the peribasal meshes of the eggshell are larger than interbasal ones in the new species while they do not differ from the interbasal ones in M. canaricus regarding the animals there are differences in the buccal armature: in M. azzunae sp. nov. the posterior band of teeth is visible with LM (even though poorly) and the three dorsal crests are distinct transverse ridges, while in M. canaricus the posterior band of teeth is visible only with SEM and with LM the dorsal teeth form a transversal ridge weakly divided into three teeth. Macrobiotus madegassus Maucci, 1993 The new species differs from M. madegassus by the presence of the eye spots (absent in M. madegassus ), pores on the cuticle (absent in M. madegassus ), presence in the buccal armature of posterior band of teeth, even though weak (fully absent in M. madegassus ), buccal tube much larger ( pt of the holotypes 15.9 vs 7), insertion of the stylet supports on the buccal tube much more posterior ( pt of the holotypes 76.1 vs 68), first and second macroplacoid longer ( pt of the holotypes 25.5 and 18.1 vs 21.3 and 12.0), lunules on the hind legs without kerning (crenate in M. madegassus ), eggshell processes with distal disc as large as the base (similar range 3.2–5.2 for both measurements) with respect to that of M. madegassus (disc vs base: 4.3–5.4 vs 2.3–2.6). Macrobiotus martini Bartels , Pilato, Lisi & Nelson, 2009 The cuticular pores in M. azzunae sp. nov. are much smaller than those of M. martini the distal disc of the egg processes in M. azzunae sp. nov. has a diameter similar to that of the process base, while in M. martini the distal disc is much narrower than the base. Macrobiotus nebrodensis Pilato, Sabella, D’Urso & Lisi, 2017 Macrobiotus azzunae sp. nov. differs from M. nebrodensis by the absence of the cuticular bar near the lunules on the first three pairs of legs (a faint bar is present in M. nebrodensis ). The egg processes of M. azzuane sp. nov. are in higher number on the circumference (29–33) with respect to those of M. nebrodensis (18). In the latter species there are some egg processes very high (up to 20.6 µm), while in the new species process height and shape are more uniform. The difference in the eggshell between meshes around the process base and the others is much less evident in M. azzunae sp. nov. than in M. nebrodensis . Macrobiotus personatus Biserov, 1990 The new species differs from M. personatus by the posterior band of the buccal armature less evident, the presence of a clear constriction in the first macroplacoid (Fig. 5A), in the paratype of M. personatus examined by us barely identifiable (Fig. 6A) and, according to Biserov (1990) usually absent in the type material of that species. The pores on the cuticle of M. azzunae sp. nov. are small, approximately 1 µm in diameter, while in M. personatus they are strongly elliptic and about 3 µm in length (Fig. 6B). Lunules on leg IV are always smooth in M. azzunae sp. nov., sometimes indented in M personatus . With respect to the eggs of M. personatus (Fig. 6C–D), the egg processes of M. azzunae sp. nov. (Figs 4C–D, 5C) are clearly shorter, 5.4 ± 0.6 vs 9.5 ± 0.5 (range 4.2–6.4 vs 9–10.5) and with a narrower base and distal disc (both 3.2–5.2 vs 7–10.5 and 7–9 respectively). Males are present in the new species, while in M. personatus only females were found (Biserov 1990), suggesting parthenogenesis in that species. Macrobiotus sandrae Bertolani & Rebecchi, 1993 The new species differs from M. sandrae for the eggshell shape, with thinner wires of the reticulum and meshes around the processes larger than the inter-process meshes in M. azzunae sp. nov. (Fig. 5C), all meshes similar in size in M. sandrae (Fig. 5D). Figure 5C–D also show a difference in the process base diameter, narrower in M. azzunae sp. nov. With regard to the animals, M. azzunae sp. nov. differs from M. sandrae by a constriction of the first macroplacoid much more pronounced (Fig. 5A; it is hardly visible in M. sandrae Fig. 5B). Moreover, in animals of similar size the posterior band of the buccal armature is just less evident in the new species, and lunules on the hind legs are without hint of teeth (but teeth, present in the holotype of M. sandrae , are often difficult to identify in other specimens of that species). Macrobiotus terminalis Bertolani & Rebecchi, 1993 Macrobiotus azzunae sp. nov. differs from M. terminalis for the absence of granulation on the cuticle (noted only in the redescription of M. terminalis see Cesari et al . 2011), for the absence of teeth on the lunules, especially evident on the hind legs of M. terminalis , and for the presence of males, absent in M. terminalis (see redescription by Cesari et al . 2011). Macrobiotus vladimiri Bertolani, Biserov, Rebecchi & Cesari, 2011 With respect to M. vladimiri , animals of M. azzunae sp. nov. reach a shorter length (up to 410.3 µm vs 515.1 µm), in M. azzunae sp. nov. the posterior band of teeth of the buccal armature is less evident and the lunules on the hind legs are not indented. In M. azzunae sp. nov. the egg diameter without processes (64.7–80.6 µm) is less than that of the eggs of M. vladimiri (89.9–92.0 µm); the processes are shorter (4.2–6.4 µm in the new species vs 6.5–8 µm in M. vladimiri ). In the new species the base process diameter is narrower (3.2–5.2 µm) than in M. vladimiri (5.1–7.3 µm), the distal disc is weakly or not jagged (clearly jagged in M. terminalis ). In M. azzunae sp. nov. males are present, while they are absent in M . vladimiri . Genetic distances The ranges of uncorrected genetic p-distances between M. azzunae sp. nov. and the other species of the M. hufelandi group (Supp. file 7, Supp. file 8, Supp. file 9, Supp. file 10), are as follows: 18S 0.1–5.6%, with the most similar being M. sandrae from Germany (present paper) 28S 0.1%, with the only available M. vladimiri from Spain (FJ435751 –5) ITS-2 7.7–32.2%, with the most similar being Macrobiotus vladimiri (MN888347) from Finland COI 6.3–25.6%, with the most similar being Macrobiotus sandrae (HQ876574, HQ876577, HQ876578, HQ876579, HQ876581) from Germany The COI dataset is the most complete and informative for species delimitation investigation. Both phylogenetic reconstructions on the COI dataset resulted in the same topology, and thus the ML tree was utilized for the PTP analysis (Fig. 7, left), which shows 14 putative species clusters: M. crustulus , M.hannae , M. cf. recens , M.canaricus , M.hufelandi , M. cf. hufelandi sp.1, M. terminalis , M. cf. terminalis , M. wandae , M. macrocalix , M . cf. macrocalix , M. vladimiri , M. sandrae and M. azzunae sp. nov. This subdivision is further validated by both the ABGD and the haplotype network analysis (Fig. 7, centre and right). Present molecular data therefore confirms the validity of the erection of M. azzunae sp. nov. : Published as part of Marnissi, Jamila Ben, Cesari, Michele, Rebecchi, Lorena & Bertolani, Roberto, 2021, Integrative description of a new Tunisian tardigrade species, Macrobiotus azzunae sp. nov. 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[In Russian.]"]} Text Greenland Tardigrade DataCite Metadata Store (German National Library of Science and Technology) Greenland Pacific Magallanes ENVELOPE(-62.933,-62.933,-64.883,-64.883) Pasa ENVELOPE(26.733,26.733,67.850,67.850) Jagged ENVELOPE(-65.683,-65.683,-65.967,-65.967) Kristensen ENVELOPE(-159.667,-159.667,-86.333,-86.333) Barros ENVELOPE(-62.167,-62.167,-64.717,-64.717) Nowak ENVELOPE(-57.917,-57.917,-61.950,-61.950) Faure ENVELOPE(-68.871,-68.871,-68.093,-68.093) Tooth The ENVELOPE(168.983,168.983,-77.517,-77.517) Giovannini ENVELOPE(-57.967,-57.967,-63.550,-63.550) |