Kleemannia Oudemans, 1930

Masan, Peter, 2017, A revision of the family Ameroseiidae (Acari, Mesostigmata), with some data on Slovak fauna, ZooKeys 704, pp. 1-228 : 1

Genus Kleemannia Oudemans, 1930

Kleemannia Oudemans, 1930: 135. Type species: Zercon pavidus C. L. Koch, 1839, by original designation.

Primoseius Womersley, 1956b: 116. Type species: Zercoseius macauleyi Hughes, 1948 (= Seiulus plumosus Oudemans, 1902), by original designation. Synonymy by Hughes (1961).

Kleemannia . - Evans 1963a: 229. not Zercoseius Berlese, 1916a: 33. Type species: Seius spathuliger Leonardi, 1899, by original designation. Incorrect synonymy by Oudemans (1939b).

Diagnosis (adults).

Dorsal shield variously sclerotised and ornamented, normally with 28-29 pairs of setae (z6 present or absent). Dorsal shield setae similar in adults of both sexes, well developed and relatively long, thickened, tubiform or flattened, mostly densely pilose or serrate. In female, st1 and st2 on sternal shield, st3 on small suboval or subcircular pseudo-metasternal platelets or sometimes on soft integument (in Kleemannia mirabilis sp. n. on sternal shield), and st4 on soft integument; endopodal platelets II-III present, subtriangular, and relatively small; anterior margin of epigynal shield convex, genital poroids on soft integument. Female with five or six pairs of opisthogastric setae (JV1-JV3, JV5, ZV2; JV4 sometimes absent), two pairs of which (JV2, JV3) on ventrianal shield (in Kleemannia insignis , JV2 occasionally on soft integument but contiguous to anterior margin of ventrianal shield), other on soft integument; male with five pairs of opisthogastric setae (JV4 always absent), 3-5 pairs of which on expanded ventrianal shield (JV1 and/or JV5 sometimes on soft integument). In both sexes, JV5 usually similar to those on dorsal shield, other ventrally inserted setae simple, smooth and needle-like. Peritrematal shields anteriorly connected to dorsal shield, and often with enlarged cavity-like poroid structure at level of coxae III. Metapodal platelets well developed, elongate. Corniculi well sclerotised and spaced, usually with bifid or trifid apex; setae h1 thickened, straight or slightly and regularly curved, progressively taperred and pointed. Fixed digit of chelicera with four subequal teeth on proximal masticatory area; male spermatodactyl relatively small, usually as long as movable cheliceral digit (if not shorter). Epistome with anterior margin produced into narrow and pointed central projection or, sometimes, short obtuse cusp; lateral margins mostly smooth. Palptarsal apotele usually two-tined. Coxae I-IV with delicate sculptural ornamentation, striate and reticulate. Genu III and tibiae III-IV with two anterolateral and two posterolateral setae. Tarsi I-IV each with empodium and claws; tarsi I usually well pigmented, brown in colour. Insemination apparatus with papilla-like sperm induction pores (solenostomes) associated with inner margin of coxae III.

Remarks.

Oudemans (1930) proposed the new genus Kleemannia with Zercon pavidus C. L. Koch, 1839 as type species, and added three further species in the genus ( Kleemannia plumosa , Kleemannia plumea and Kleemannia plumigera ). Since then, this taxon has been variously treated as a genus ( Vitzthum 1942; Hughes 1961, 1976; Evans 1963a; Ishikawa 1972; Domrow 1974, 1979; Evans and Till 1979), a subgenus of Ameroseius ( Athias-Henriot 1959, Hajizadeh et al. 2013a), or a junior synonym of Ameroseius ( Westerboer and Bernhard 1963; Bregetova 1977; Karg 1993, 2005; Halliday 1997; Karg and Schorlemmer 2009; Narita et al. 2013b), although it may be easily and reliably separated from Ameroseius by the presence of two posterolateral setae on genu III and tiabiae III-IV (one in Ameroseius ), four proximal denticles on fixed digit of chelicerae (at most three denticles in Ameroseius ), two pairs of opisthogastric setae on ventrianal shield (at most one pair in Ameroseius , but never JV2), two-tined claw on palptarsus (three-tined in Ameroseius ), thickened h1 (similar if compared to other hypostomal setae in Ameroseius ), epistome usually with sharply or obtusely pointed apex (never produced into a single pointed process in Ameroseius ). Therefore, the genus Kleemannia is considered to be a valid genus, and is removed from synonymy with Ameroseius .

The related genus Primoseius was proposed and very briefly characterised by Womersley (1956b). He included two species in the genus, the type species, mentioned as Lasioseius (Zercoseius) macauleyi Hughes, 1948 (sic, should be Zercoseius macauleyi ), and Lasioseius (Zercoseius) gracei Hughes, 1948 (sic, should be Zercoseius gracei ), which were subsequently synonymised with Kleemannia plumosa and Kleemannia plumigera , respectively ( Evans 1954). Later, the separate taxonomic status of this genus was rejected by Hughes (1961), Evans and Till (1979), Karg (1971a, 1993, 2005), Bregetova (1977), and many others, because the original concept of Primoseius was defined with the help of the same characters as they can be found in the most relative genus Ameroseius , or Kleemannia s. l. Womersley (1956b) based the diagnosis of his new genus especially on the possession of two pairs of setae (st1, st2) on sternal shield, a pair of pseudo-metasternal platelets each bearing a seta (st3), and metasternal setae (st4) on soft integument. Despite these facts, Primoseius is included as a subgenus of Kleemannia in the list of valid taxa of Ameroseiidae in this paper. It may be reliably distinguished from Kleemannia s. str. mainly by the absence of z6, and presence of specifically modified dorsal setae (conspicuously flattened, leaf- to feather-shaped). Primoseius currently comprises 12 described species ( Primoseius bisetae , Primoseius dipankari , Primoseius dubitatus , Primoseius eumorphus , Primoseius macauleyi , Primoseius mineiro , Primoseius parplumosa , Primoseius plumosoides , Primoseius plumosa , Primoseius potchefstroomensis , Primoseius pseudoplumosa and Primoseius wahabi ), of which only eight species are considered here to be valid ( Primoseius bisetae , Primoseius dipankari , Primoseius mineiro , Primoseius parplumosa , Primoseius plumosoides , Primoseius plumosa , Primoseius pseudoplumosa and Primoseius wahabi ). Narita et al. (2013b) classified eight species of " Ameroseius " into a species group named Ameroseius plumosus -group ( Ameroseius dipankari , Ameroseius eumorphus , Ameroseius mineiro , Ameroseius parplumosa , Ameroseius plumosa , Ameroseius potchefstroomensis , Ameroseius reticulatus and Ameroseius wahabi ). They defined the group by the following character states, of which some are vague and applicable also for other groups inside the genus: Ameroseius corniculus bifid or trifid; dorsal shield reticulate and without pit-like depressions; 26 or 28 pairs of dorsal shield setae, most of which are lanceolate to leaf-shaped; and five or six pairs of opisthogastric setae, two pairs of which are on ventrianal shield. The Ameroseius plumosus -group of Narita et al. (2013b) could be considered to be identical with Primoseius , and it omits three species described before 2013, namely Kleemannia bisetae , Kleemannia plumosoides and Kleemannia pseudoplumosa , and inexplicably includes Kleemannia reticulata , which has z6 clearly developed and present on the dorsal shield.

Kleemannia currently comprises 28 valid species described mostly from the Palaearctic, and only rarely from Neotropics (four species), Nearctic (one species), Afrotropics (one species) and Oriental Region (two species). Specimens reported as Kleemannia plumosa and Kleemannia plumigera have been reported from many countries around the world, from various natural and synanthropic habitats (stored grain and food, litter in sheds and stalls, mould in buildings). In Slovakia, this genus is represented by eight recorded species.

Key to species of Kleemannia occurring in Europe (females)

Partial keys to species of Kleemannia (classified in Ameroseius ) from Europe and former Soviet Union may be found in Karg (1971a, 1993) and Bregetova (1977), and they include eight and 13 species, respectively. The world species can be identified using keys from Narita et al. (2013b, 2015). A new key to the genus is needed to include a species here firstly reported from Europe ( Kleemannia kosi and Kleemannia parplumosa ), the new species described in this study ( Kleemannia dolichochaeta sp. n.), and a revised Berlese species previously considered to be dubious ( Kleemannia tenella ). The identification key for 12 species presented below is based on direct examination of type specimens or specimens considered to be conspecific with the type material, except for Kleemannia elegans .

Several diagnostic features can be useful in recognising of the Kleemannia species occurring in Europe, namely (1) presence/absence of z6 (absent in Kleemannia dolichochaeta , parlplumosa, Kleemannia plumosa and Kleemannia pseudoplumosa ); (2) placement of st3 (on soft integument in Kleemannia plumea and Kleemannia tenella ); (3) number of opisthogastric setae (with reduced number of five pairs in Kleemannia insignis , Kleemannia plumea and Kleemannia tenella ); (4) presence/absence of cavity-like poroid structure on peritrematal shield (absent in Kleemannia plumea , Kleemannia plumigera and Kleemannia tenella ); (5) placement of JV2 and JV3 on anterior portion of ventrianal shield (both setae on medial surface in Kleemannia elegans , Kleemannia kosi , Kleemannia pavida and Kleemannia plumigera ; JV2 on anterior margin or surface in Kleemannia dolichochaeta , Kleemannia insignis , Kleemannia parplumosa , Kleemannia plumosa and Kleemannia pseudoplumosa ; both setae on anterior margin or surface in Kleemannia delicata and Kleemannia tenella ; and (6) relative length of Z- and S-setae (Z-setae apparently shorter than S-setae in Kleemannia tenella ).

1	Dorsal shield with 28 pairs of setae (z6 absent); dorsal setae flattened, feather- to leaf-shaped ( subgenus Primoseius Womersley, 1956)	2
-	Dorsal shield with 29 pairs of setae (z6 present); dorsal setae variously formed, mostly tubiform, rod-shaped, lanceolate or oblanceolate ( subgenus Kleemannia s. str.)	5
2	Sternal shield with inverted U-shaped sculptural structure on anterior surface, the structure with anterior margin scalloped and heavily sclerotised	Kleemannia plumosa (Oudemans, 1902) (Plates 53 - 56 )
-	Sternal shield with simple sculpture on anterior surface, smooth or reticulate	3
3	Epigynal shield with inverted U-shaped sculptural structure on anterior surface, the structure smooth and well sclerotised	Kleemannia parplumosa Nasr & Abou-Awad, 1986 (Plate 45 )
-	Epigynal shield with simple sculpture on anterior surface, smooth or reticulate	4
4	Setae j5 apparently shorter than J2 and J4, with tips reaching between bases of j5 and j6	Kleemannia pseudoplumosa (Rack, 1972) (Plates 57 - 60 )
-	Setae j5 notably lengthened, similar in length to J2 and J4, with tips reaching beyond bases of j6	Kleemannia dolichochaeta sp. n. (Figure 3 , Plate 38 )
5	Epigynal shield with inverted U-shaped sculptural structure on anterior surface, the structure smooth and well sclerotised	Kleemannia plumea Oudemans, 1930 (Plates 48 , 49 )
-	Epigynal shield with simple sculpture on anterior surface, smooth or reticulate	6
6	Ventrianal shield with at least one pair of opisthogastric setae (JV2, sometimes also JV3) on or very close to its anterior edge	7
-	Ventrianal shield with both pairs of opisthogastric setae inserted well inside the anterior portion of the shield	9
7	Dorsal shield with j-setae apparently shorter than J-setae; setae j6 with tips reaching between bases of j6 and J2; dorsal shield densely granulated; ventrianal shield wider than long, rugose	Kleemannia insignis (Bernhard, 1963) (Plates 39 , 40 )
-	Setae j1-j6, J2 and J4 similar in length; setae j6 reaching beyond bases of J2; idiosomal shields not granulated or rugose; ventrianal shield about as wide as long	8
8	Setae z5, z6, and Z1-Z3 apparently shorter than adjacent setae in central and latero-marginal rows; setae st3 on soft integument; five pairs of opisthogastric setae present (JV4 absent); peritrematal shields without conspicuous poroid structures	Kleemannia tenella (Berlese, 1916) (Plates 61 - 63 )
-	Setae z5, z6, and Z1-Z3 relatively long, almost subequal to those in central and latero-marginal rows; setae st3 on small suboval pseudo-metasternal platelets; six pairs of opisthogastric setae present (JV4 developed); peritrematal shields each with a cavity-like poroid structure	Kleemannia delicata (Berlese, 1918) (Plates 35 - 37A , 37C , 76A , 76H , 77A )
9	Pseudo-metasternal platelets partly fused to sternal shield; basal part of sperm induction pores conspicuously thickened and heavily sclerotised	Kleemannia plumigera Oudemans, 1930 (Plates 50 - 52 , 76B , 76D , 77C , 77D )
-	Pseudo-metasternal platelets free from sternal shield; basal part of sperm induction pores weakly sclerotised	10
10	Ventrianal shield slightly constricted laterally (posteriad of setae JV3); transverse row of setae j6, z6, s5 and r5 with j6 longest; setae J4 relatively longer, with tips reaching to or beyond posterior margin of dorsal shield; dorsal shield unevenly reticulate	Kleemannia kosi El-Badry, Nasr & Hafez, 1979 (Plates 41 , 42 )
-	Ventrianal shield regularly curved laterally; transverse row of setae j6, z6, s5 and r5 with subequal setae (z6 slightly shorter); setae J4 relatively shorter, reaching between their bases and posterior margin of dorsal shield; dorsal shield entirely reticulate	11
11	Medial dorsal setae rather lanceolate	Kleemannia elegans (Bernhard, 1963)
-	Medial dorsal setae rather oblanceolate	Kleemannia pavida (C. L. Koch, 1839) (Plates 37B , 46 , 47 )

Figures

Figure 3. Kleemannia dolichochaeta sp. n., female. A Dorsal idiosoma B Ventral idiosoma. Not scaled.

Plate 35. Kleemannia delicata, female, dorsal idiosoma.

Plate 37. Kleemannia spp., female. A Kleemannia delicata, epistome B Kleemannia pavida, epistome C Kleemannia delicata, ventrianal shield.

Plate 38. Kleemannia dolichochaeta, female (holotype, from Spain), dorsal idiosoma.

Plate 39. Kleemannia insignis, female, dorsal idiosoma.

Plate 40. Kleemannia insignis, female. A Dorsal idiosoma with legs B Posterior margin of idiosoma C Setae j 1 D Ventrianal shield, a part E Dorsal shield sculpture F Sternal shield and metasternal area.

Plate 41. Kleemannia kosi, female (from Germany), dorsal idiosoma.

Plate 42. Kleemannia kosi, female (from Germany), dorsal idiosoma.

Plate 45. Kleemannia parplumosa, female (from Germany), dorsal idiosoma (pressed specimen).

Plate 46. Kleemannia pavida, female (from Germany), dorsal idiosoma.

Plate 47. Kleemannia pavida, female (from Germany), ventral idiosoma.

Plate 48. Kleemannia plumea, female, dorsal idiosoma.

Plate 49. Kleemannia plumea, female, ventral idiosoma.

Plate 50. Kleemannia plumigera, female, dorsal idiosoma.

Plate 52. Kleemannia plumigera, female (from Germany), ventral idiosoma.

Plate 53. Kleemannia plumosa, female, dorsal idiosoma.

Plate 56. Kleemannia plumosa, female, posterior part of dorsal shield. A A form with shorter dorsal setae B A form with longer dorsal setae.

Plate 57. Kleemannia pseudoplumosa, female, dorsal idiosoma (pressed specimen).

Plate 60. Kleemannia pseudoplumosa, female (from Greece), ventral idiosoma (pressed specimen).

Plate 61. Kleemannia tenella, female, dorsal idiosoma.

Plate 63. Kleemannia tenella, female, dorsal idiosoma.

Plate 76. Ameroseiidae spp., female. A-E Structures of sperm access system. A Kleemannia delicata, sperm induction pores (solenostomes) associated with inner margin of coxae III B and D Kleemannia plumigera, solenostomes associated with inner margin of coxae III C Epicriopsis suedus, sacculus foemineus and associated tubuli annulati E Sertitympanum nodosum, tubuli annulati F-H Chelicera. F Asperolaelaps rotundus G Ameroseius renatae H Kleemannia delicata.

Plate 77. Ameroseiidae spp., female. A-C Poroid structure associated with peritrematal shield. A Kleemannia delicata B Sertitympanum nodosum C Kleemannia plumigera D Metasternal region in Kleemannia plumigera.