Claparedrilus Klinth, Rota & Erséus, 2017b

Claparedrilus Klinth, Rota & Erséus, 2017b View in CoL

Type species. Claparedrilus semifuscoides Klinth, Rota & Erséus, 2017b View in CoL .

Other species. Claparedrilus semifuscus (Claparède, 1861) View in CoL .

Claparedrilus torquatus Klinth & Rota View in CoL sp. nov. Figure 16 View Figure 16

urn:lsid:zoobank.org:act:F346D5D2-E051-43FE-B089-A7D26FC7AE82

Holotype. ZMBN 110970 View Materials ( CE23134 ), Figs 16A,C–E View Figure 16 , an amputated mature specimen stained in paracarmine and mounted on a slide. COI barcode: BOLD NOENC418-18 .

Type locality. Rotsundselv, Nordreisa, Troms, Norway. In sand at the upper intertidal. Coll. C. Erséus, 14 August 2014 .

Paratype. ZMBN 110972 View Materials ( CE23153 ) one immature specimen from the type locality. For details on collection site and GenBank accession numbers see Table S1.

Etymology. The name refers to the large ectal glands forming a collar at the ectal end of the spermathecae (Latin torquatus , meaning collar-bearing).

Diagnosis. This new species can be distinguished from C. semifuscus (Claparède, 1861) which has huge penial bulbs, much larger than the sperm funnels, whereas the penial bulbs of C. torquatus sp. nov. are about as large as the sperm funnels. It can be distinguished from C. semifuscoides by having an ectal spermathecal gland mass that is larger than the spermathecal ampulla, whereas C. semifuscoides has a gland mass that is about the same size or smaller than the ampulla.

Description. Length of first 30–35 segments 4.0–5.0 mm (fixed, amputated specimens); first 15 segments 2.0– 2.2 mm long; width at clitellum 0.44–0.54 mm. Chaetae sigmoid. Dorsolateral bundles with 3–5 chaetae anterior to clitellum, and 3–6 chaetae in postclitellar segments. Ventral bundles with 4–6 chaetae anterior to clitellum, and 4–7 chaetae in postclitellar segments. Each worm’s longest measured chaetae 75–80 µm long, about 5 µm wide. Clitellum extending over XII–1/2XIII. Head pore at 0/1.

Coelomocytes numerous, about 15–20 µm long, round or oval, granulated with distinct nucleus. Paired pharyngeal glands present in IV, V, VI and VII; dorsal connection uncertain ( Figure 16A View Figure 16 ). Dorsal vessel originating in XIII. Nephridia observed in 7/8–8/9 and postclitellar segments, about 135 µm long in segment VIII. Anteseptale small, consisting of funnel and a thin neck. Postseptale oval, tapering into thin posteroventral efferent duct ( Figure 16B View Figure 16 ). Brain with posterior incision.

Male genitalia paired ( Figure 16C View Figure 16 ). Testes originating in XI, with some developing sperm seemingly held together by testis sacs, forming irregular lobes in IX–XI, other cysts of developing sperm floating free in VIII–XI. Sperm funnels in XI, 145 µm long, 90 µm wide, making them about 1.5 times longer than wide, funnels tapering towards vasa deferentia. Large parts of vasa irregularly coiled around ovaries in XII, vasa 10 µm wide. Penial bulbs compact, 140 µm in diameter. No fully mature eggs observed.

Spermathecae ( Figures 16D–E View Figure 16 ) in V, club-shaped, with ectal duct suddenly widening into ampulla. Ectal duct lined by layer of musculature. Ampulla round, connected to oesophagus via tapering ental duct. Sperm filling lumen of ectal duct and ampulla. Spermathecae 120 µm long, 35 µm wide at the ectal duct, 55 µm wide at widest part of ampulla. Ectal pore at lateral line, surrounded by gland cells forming a large collar-like, somewhat lobed compact mass, up to 115 µm wide. No midventral subneural glands observed.

Geographical distribution including BOLD data. Only known from the type locality in Northern Norway (Troms) .

Remarks. This species is supported as a member of Claparedrilus both genetically ( Figure 2 View Figure 2 ) and morphologically by the following characters: four pairs of pharyngeal glands, irregularly lobed testis sacs, and a thin anteseptale neck on the nephridium. It is similar to both C. semifuscus and C. semifuscoides in chaetal pattern and size, but has smaller penial bulbs than the former and larger spermathecal ectal glands than the latter. The quality of the original description of C. semifuscus was questioned by Klinth et al. (2017b) and its relationship to both C. semifuscoides and C. torquatus sp. nov. may be revaluated after future sampling. DISCUSSION

come from the combined collecting effort of many years with

This paper is the culmination of work started in 2017 (Klinth et different sources of funding, mainly from the Norwegian and Swedish al. 2017a; 2017b) that became the basis for a revision of Lumbricillus , Taxonomy Initiatives, and the Swedish Environmental Protection with a focus on the North East Atlantic species. The addition of Agency.

specimens from the Southern Hemisphere, which also included material of what we consider to be Marionina s. str., provided another important piece of the puzzle ( Klinth et al. 2022). Now, with the SUPPLEMENTARY INFORMATION

establishment of Paralumbricillus gen. nov., we can finally exclude the arenarius group from Lumbricillus . This has led to a more clearly Figure S1 View Figure 1 . Majority-rule consensus tree for 16S estimated with defined Lumbricillus in which most species have the characteristic Bayesian inference. Specimens marked in green are new to this study. lobed testis sacs arranged in a fan shape. Exceptions remain in the Support values are posterior probabilities. Scale bar represents the buelowi group, which constitutes the sister clade to the remaining estimated number of substitutions per site.

Lumbricillus species, where the included species have testes with only a single lobe. The members of the buelowi group resemble those of Figure S2 View Figure 2 . Majority-rule consensus tree for ITS2 estimated with Paralumbricillus gen. nov. in having only 2 or 3 chaetae per bundle Bayesian inference. Specimens marked in green are new to this study. and club-shaped spermathecae but have short sperm funnels (unlike Support values are posterior probabilities. Scale bar represents the most Paralumbricillus gen. nov.) and the testis sac is not irregularly estimated number of substitutions per site.

lobed or budding off into free floating cysts. There remains some dubious species in Lumbricillus with irregularly lobed testis sacs Figure S3 View Figure 3 . Majority-rule consensus tree for H3 estimated with that should most probably be transferred to another genus, perhaps Bayesian inference. Specimens marked in green are new to this study. even Paralumbricillus gen. nov. ( Table 1), but we believe that better Support values are posterior probabilities. Scale bar represents the descriptions in combination with genetic information are needed before estimated number of substitutions per site.

making such changes. Another future aspect concerning Lumbricillus is the re-evaluation of the numerous species still with incomplete Table S1. List of specimens used in this study, with specimen descriptions, including the unnamed ones treated in this study and in identification number, collection data, GPS coordinates (in decimal Klinth et al. (2017b). Of course, there are most likely a high number degrees, WGS84), GenBank accession numbers for seven different of unknown species yet to be discovered around the world. So far we markers (bold numbers are new sequences generated in this study) have focused mainly on the Scandinavian coasts and even here not all and voucher numbers. Letters for Lumbricillus pagenstecheri refer species have been identified. to barcoding clusters (see Klinth et al. 2017a). Country codes: In previous papers ( Klinth et al. 2017b; 2022), the arenarius group AU= Australia, DK= Denmark, ES= Spain, FR= France, GL= Greenland, was discussed in relation to Enchytraeoides Roule, 1888 (see Rota et NL= Netherlands, NO= Norway, SE= Sweden and UK =United al. 2008). In short, the type species Pachydrilus enchytraeoides Saint- Kingdom. * Unsuccessfully sequenced marker.

Loup, 1885 (possibly the same as Enchytraeoides marioni Roule, 1888 )

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