Knarr compressa ( Golemansky, 1979 ) Soler-Zamora & Useros & González-Miguéns & Gómez-Rodríguez & Lara, 2023

Soler-Zamora, Carmen, Useros, Fernando, González-Miguéns, Rubén, Gómez-Rodríguez, Pablo & Lara, Enrique, 2023, The problem of ‘ shadow species’ as illustrated with the taxonomic hotchpotch Cyphoderia ampulla (Rhizaria: Cyphoderiidae), Zoological Journal of the Linnean Society 199 (2), pp. 477-492 : 489-490

Knarr compressa ( Golemansky, 1979) comb. nov.

( Figs 1 , 7 D )

Zoobank registration: urn:lsid:zoobank.org:act:9325FB59-68C8-43B2-A556-401D4F4FEE0F .

Oleiformis Soler-Zamora, Useros, González-Miguéns & Lara gen. nov.

( Figs 1 , 3 , 5 )

Zoobank registration: urn:lsid:zoobank.org:act:D473BEDF-9357-4AD7-81DA-123888857181 .

Type species: Oleiformis major (Penard, 1890) comb. nov.

Description: Scales from the shell are non-imbricated, the species inhabits in freshwater ecosystems and they have a balloon-shaped shell that measures between 50–100 μm more than the largest species in the sister genus Cyphoderia .

Etymology: The genus name refers to the shape like an olive.

Included taxa: See Supporting Information, Appendix S1.

Oleiformis carmelae Soler-Zamora, Useros, González-Miguéns & Lara sp. nov.

( Figs 1 , 5 B 2 ; Model 2; Supporting Information, Fig. S1 )

Zoobank registration: urn:lsid:zoobank.org:act:CD3A3A3F-B2FC-4FBF-9B79-6F5EDC99CB6C .

Type locality: Morollón River , Fuente de la Gitana recreational area, La Peza, Spain (37º 51ʹ N 2º 47ʹ W) .

Habitat: River sediment.

Measurements: 169.92–190.83 μm length, 67.44–83.14 μm width, 22.51–27.85 μm pseudostome large axis, and

2.65–2.99 × 1.93–2.26 μm scales from the middle of the shell (N = 13).

Specific diagnosis: Elongated shell with an almost perfectly circular section. The shell is larger towards its middle part, giving an inflated appearance; rounded fundus. Short neck in comparison to other members of Cyphoderiidae . Rounded pseudostome with a thin organic layer surrounding it. Elliptical scales placed next to each other in a structured fashion; each scale is surrounded by six pores.

Intraspecific variability: Some individuals have a slightly more pointed fundus (Supporting Information, Fig. S1 ).

Differential diagnosis: It can be confused with O. major because of their similar shell shape. O. carmelae is smaller, with relatively larger scales. It differs also genetically in both COI and SSU.

Etymology: In honour of Professor Carmen Zamora-Muñoz, mother of the first author.

Holotype: MA-Amoeba 11264.

Figures

Figure 1. Bayesian phylogenetic tree based on SSU and COI sequences of the family Cyphoderiidae.At each node posterior probability values (Bayesian analysis) and bootstrap values (Maximum Likelihood analysis) are represented.The colours represent the molecular clades. GenBank accession codes are given on the tips. Species names correspond to the taxonomic assignation of the organisms based on morphology for each lineage.

Figure 2. A, Linear discriminant analysis of four continuous morphological traits.B, Number of shells classified into their own clade under the linear discriminant analysis.C, Linear discriminant analysis of four continuous morphological traits of clade C. D, Linear discriminant analysis of four continuous morphological traits of clade A.

Figure 3. Dichotomous key of the family Cyphoderiidae.New genera are in bold with their respective shell outline drawing (50 μm scale bar), scale arrangement under SEM (grey box with 10 μm scale bar), and shell outline under light microscopy (orange box) in order to differentiate them. (A) Cyphoderia; (B) Oleiformis gen. nov.; (C) Ichthyosquama gen. nov.: (D) Knarr gen. nov., (E) Psammoderia gen. nov.

Figure 5. SEM images of the different species of Oleiformis gen. nov. Insets show the typical habitat for each organism. At the bottom, an original drawing of the previously known species is shown. The coloured squares at the top left of each image have the same colour as the clades to which each taxon corresponds in Fig.1. The white bar represents a scale of 10 μm and the black bar 50 μm for each organism.

Figure 7. SEM images of the different species of Knarr gen. nov. and the original drawings of both genera, Knarr gen. nov. and Psammoderia gen. nov. Insets show the typical habitat for each organism. The coloured squares at the top left of each image have the same colour as the clades to which each taxon corresponds in Fig. 1. The white bar represents a scale of 10 μm and the black bar 50 μm for each organism.