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A new species and new records of thesubgenus Typhlodromus Scheuten fromSpain, with a key to the world species(Acari: Phytoseiidae)Francisco Ferragut a & Edward A. Ueckermann b ca Instituto Agroforestal Mediterráneo. Universidad Politécnica deValencia, Camino de Vera, s/n, 46022, Valencia, Spainb ARC-Plant Protection Research Institute, Private Bag X134,Queenswood, Pretoria, 0121, South Africac School of Environmental Sciences and Development, North-WestUniversity, Potchefstroom campus, Potchefstroom, 2520, SouthAfrica

Version of record first published: 25 Jun 2012

To cite this article: Francisco Ferragut & Edward A. Ueckermann (2012): A new species and newrecords of the subgenus Typhlodromus Scheuten from Spain, with a key to the world species (Acari:Phytoseiidae), Journal of Natural History, 46:27-28, 1731-1745

To link to this article: http://dx.doi.org/10.1080/00222933.2012.681318

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Journal of Natural HistoryVol. 46, Nos. 27–28, July 2012, 1731–1745

A new species and new records of the subgenus Typhlodromus Scheutenfrom Spain, with a key to the world species (Acari: Phytoseiidae)

Francisco Ferraguta* and Edward A. Ueckermannb,c

aInstituto Agroforestal Mediterráneo. Universidad Politécnica de Valencia. Camino de Vera,s/n. 46022 Valencia, Spain; bARC-Plant Protection Research Institute, Private Bag X134,Queenswood, Pretoria 0121, South Africa; cSchool of Environmental Sciences and Development,North-West University, Potchefstroom campus, Potchefstroom 2520, South Africa

(Received 25 October 2011; final version received 22 March 2012; printed 26 June 2012)

A new mite species of the family Phytoseiidae (Acari: Mesostigmata), Typhlodromus(Typhlodromus) thuriferus sp. nov. collected from the Cupressaceae Juniperus thu-rifera in Spain is described. Three species of the same subgenus, new records forSpain, are also included. A key to all the species of this subgenus is given.

Keywords: Acari; Phytoseiidae; Typhlodromus (Typhlodromus); taxonomy; Spain

Introduction

The subgenus Typhlodromus Scheuten (Acari: Phytoseiidae) was first recognized as aninfrageneric taxon included in the genus Typhlodromus by Chant (1957), to accommo-date species with more than four pairs of anterior lateral setae on the dorsal shield;it belongs to the tribe Typhlodromini of the subfamily Typhlodrominae (Chant andMcMurtry, 2007). The species in this subgenus have setae j3, z2, z3, z4, s4 and s6 onthe margins of podoscutum and morphologically resemble species in the subgenusAnthoseius De Leon except in the absence of seta S5. At present it includes 52 validspecies (Kolodochka 2003; Moraes et al. 2004; Ehara et al. 2006; Chant and McMurtry2007; Ueckermann et al. 2008; Papadoulis et al. 2009; Palevsky et al. 2009; Tixier et al.2010; El-Banhawy and Knapp 2011). Apparently their centre of distribution is thePalaearctic region (Chant and Yoshida-Shaul 1987) where at least 90% of the knownspecies occur (Tixier et al. 2008). Most members of this subgenus exhibit a considerablemorphological resemblance and species recognition is particularly difficult. Althoughspecies in this group have been reviewed several times (Chant and Yoshida-Shaul 1987;Karg 1989; Denmark 1992), understanding of the variation among these species hasyet to be established.

This paper refers to a new phytoseiid mite belonging to this subgenus collectedfrom Juniperus thurifera L. in Spain. This species has three pairs of preanal setae,an unusual character for the group. Notes on three species, new records for Spain,are also given. Although previous revisions provide keys for identification, no key toidentify all the known members of this subgenus is available; for this reason a key toworld species is proposed at the end of the paper. We resolved to include the unstable

*Corresponding author. Email: fjferrag@eaf.upv.es

ISSN 0022-2933 print/ISSN 1464-5262 online© 2012 Taylor & Francishttp://dx.doi.org/10.1080/00222933.2012.681318http://www.tandfonline.com

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1732 F. Ferragut and E.A. Ueckermann

Typhlodromus (Typhlodromus) rarus Wainstein with setae j3 and S5 (one or both mem-bers) absent or present (Kolodochka 1985), excluded from their key by Chant andYoshida-Shaul (1987) but included in their list of species of this subgenus by Chantand McMurtry (2007). We also experienced problems in distinguishing T. (T.) phy-laktioticus Papadoulis and Emmanouel, 1990 from T. (T.) tubifer Wainstein, 1961;T. (T.) zaheri Denmark, 1992 from T. (T .) athiasae Porath and Swirski, 1965; andT. (T.) difficilis Kolodochka, 2003 from T. (T.) olympicus Papadoulis and Emmanouel,1993 and therefore tentatively consider them synonyms until the type specimens canbe compared. Although T. (T.) neomagdalenae Zannou, Moraes and Oliveira wasdescribed as having five pairs of dorsal solenostomes (Ueckermann et al. 2008) subse-quent re-examination of type material confirms that it has four pairs of solenostomes:gd2, gd6, gd8 and gd9 (G.J. de Moraes, personal communication).

Materials and methods

Specimens of phytoseiid mites from Spain were collected by the first author fromundisturbed habitats. Mites were collected under a stereomicroscope or extractedfrom plant material using Berlese funnels. They were then cleared in Nesbitt’s fluidand mounted in Heinze-PVA medium. Observations and drawings were made usinga Nikon Optiphot 2 interference contrast microscope. Voucher specimens of thespecies collected were deposited in the Laboratory of Acarology, Instituto AgroforestalMediterráneo, Universidad Politécnica de Valencia, Spain. Deposition of type spec-imens is indicated under the new described species. Several paratype specimens ofthe new species will also be deposited in the Arachnida Collection of ARC-PlantProtection Research Institute, Pretoria, South Africa.

All measurements are given in micrometres (µm), with those of the holotype fol-lowed by the range in the measurements of the paratypes in brackets in the case ofthe new species. Measurements of setae and other morphological characters in thenew records are provided whenever there is insufficient morphological information inthe literature or when differences with known data are found. Setal nomenclature fol-lows Lindquist and Evans (1965) as adapted by Rowell et al. (1978) and Chant andYoshida-Shaul (1991) for the dorsal and ventral surfaces of phytoseiid mites, respec-tively. Nomenclature used for dorsal solenostomes is that proposed by Athias-Henriot(Athias-Henriot 1975; Swirski et al. 1998) and the world distribution of each species isbased on Moraes et al. (2004).

Results

Typhlodromus (Typhlodromus) Scheuten

Typhlodromus Scheuten 1857: 111Typhlodromus (Typhlodromus) Chant 1957: 528; Chant and McMurtry 1994: 255;

Moraes et al. 2004: 360; Chant and McMurtry 2007: 157Wainsteinius Arutunjan 1969: 180 (synonymy according to Chant and McMurtry

1994)Typhlodromus (Trionus) Denmark 1992: 32 (synonymy according to Chant and

McMurtry 1994)

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Journal of Natural History 1733

Typhlodromus (Oudemanus) Denmark 1992: 34 (synonymy according to Chant andMcMurtry 1994)

pyri species group, Chant and Yoshida-Shaul 1987: 1771Type species: Typhlodromus pyri Scheuten 1857: 111.

Typhlodromus (Typhlodromus) baccettii Lombardini

Typhlodromus baccettii Lombardini 1960: 19; Ragusa and Swirski 1978: 215; Moraeset al. 1986: 243; Chant and Yoshida-Shaul 1987: 1780; Denmark 1992: 30; Edlandand Evans 1998: 282.

Typhlodromus (Typhlodromus) baccettii Lombardini, Moraes et al. 2004: 362; Barbaret al. 2005: 147

Typhlodromus (Typhlodromus) helenae Schicha and Dosse, 1974: 79 (synonymy accord-ing to Chant and Yoshida-Shaul 1987: 1780)

Anthoseius oligadenus Athias-Henriot, 1978: 697 (synonymy according to Chant andYoshida-Shaul 1987: 1780)

Specimens examined

Five females, Els Angels, Girona Province, 41◦59′23′′ N, 2◦54′49′′ E, 445 m above sealevel (a.s.l.), on Erica arborea L. (Ericaceae), 3 August 2004, F. Ferragut.

Previous records

Australia, France, Italy, Norway.

Remarks

Measurements of the five females collected were similar to the species redescriptionprovided by Ragusa and Swirski (1978) for specimens from Italy. Measurements of thefive females are: dorsal shield, 314 long (306–320), 164 wide (158–166); genital shield,69 wide (66–72); ventrianal shield, 94 long (84–100), 96 wide (92–98). Setae j1: 21(20–22), j3: 25 (24–27); j4: 12 (11–12), j5: 14 (13–14), j6: 16 (16–17), J2: 17 (16–18), J5:7 (7–8), z2: 16 (14–16), z3: 18 (15–20), z4: 20 (18–22), z5: 14 (14–15), Z4: 30 (28–32),Z5: 51 (48–54), s4: 24 (22–25), s6: 26 (24–27), S2: 27 (25–28), S4: 29 (26–32), r3: 23(22–25), R1: 20 (19–22), JV5: 41 (38–42). SgeIV: 21 (20–22); StiIV: 26 (25–26); St IV:45 (42–48). This species has three pairs of dorsal solenostomes (gd2, gd6 and gd9).Chaetotactic formula of genus II: 22/0, 2/01. Although previous authors have con-sidered that it bears three subapical teeth on fixed digit of chelicerae, the specimensstudied here clearly show four teeth, with a strong tooth proximal to the pilus dentilis.

Typhlodromus (Typhlodromus) corticis Herbert

Typhlodromus corticis Herbert, 1958: 429; Rasmy and MacAphee 1970: 173; Chantet al. 1974: 1274; Moraes et al. 1986: 243; Chant and Yoshida-Shaul 1987: 1776;Corino 1989: 399; Chiappini 1991: 87; Denmark 1992: 17; Wu et al. 1992: 50;Meshkov 1999: 430; Karg and Huhta 2009: 518; Denmark and Evans 2011: 389.

Typhlodromus (Typhlodromus) corticis Herbert, Moraes et al. 2004: 363; Papadouliset al. 2009: 153.

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1734 F. Ferragut and E.A. Ueckermann

Typhlodromus rodovae Wainstein and Arutunjan, 1968: 1241 (synonymy according toChant and Yoshida-Shaul 1987: 1776)

Specimens examined

Two females, Uña, Cuenca Province, on Pinus nigra subsp. salzmannii (Dunal) Franco(Pinaceae), 22 October 1994, F. Ferragut.

Previous records

Armenia, Azerbaijan, Canada, China, Finland, Greece, Italy, Moldova, Norway,Russia, Ukraine.

Remarks

The morphological characters of the specimens collected resemble those given byChant and Yoshida-Shaul (1987) except for the shorter s6. Average measurementsof the two females: dorsal shield, 362 long and 168 wide; genital shield, 74 wide;ventrianal shield, 114 long, 96 wide. Setae j1: 22, j3: 28; j4: 16, j5: 16, j6: 18, J2: 21,J5: 8, z2: 18, z3: 25, z4: 22, z5: 16, Z4: 32, Z5: 51, s4: 24, s6: 24, S2: 29, S4: 32, r3: 25,R1: 26, JV5: 44. SgeIV 25; StiIV: 26; St IV: 48. This species bears five pairs of dorsalsolenostomes (gd2, gd4, gd6, gd8 and gd9). Chaetotactic formula of genus II: 22/0,2/01.

Typhlodromus (Typhlodromus) ernesti Ragusa and Swirski

Typhlodromus ernesti Ragusa and Swirski, 1978: 211; Moraes et al. 1986: 243; Chantand Yoshida-Shaul 1987: 1798; Denmark 1992: 20; Steeghs et al. 1993: 24; Edlandand Evans 1998: 284; Swirski and Amitai 1997: 13; Meshkov 1999: 430.

Typhlodromus (Typhlodromus) ernesti Ragusa and Swirski, Moraes et al. 2004: 364;Barbar et al. 2005: 147.

Typhlodromus ernesti postici Karg 1989: 275.Typhlodromus ernesti ernesti Karg 1989: 280.

Specimens examined

Three females, one male, Caldes de Boi, Lerida Province, 42◦33′48′′ N, 0◦50′44′′ E,1530 m a.s.l., on Abies alba Mill. (Pinaceae), 7 July 2000, F. Ferragut.

Previous records

Canada, France, Israel, Italy, Norway, Russia, Sweden.

Remarks

The measurements of three adult females are similar to those of the holotype, exceptfor the shorter z3 and z4. Measurements of the three females are: dorsal shield,344 long (330–356), 160 wide (146–170); genital shield, 71 wide (70–72); ventrianalshield, 109 long (98–116), 100 wide (92–110). Setae j1: 22 (20–25), j3: 25 (24–26); j4:15 (15–16), j5: 14 (14–15), j6: 18 (16–20), J2: 19 (18–22), J5: 5 (5–6), z2: 16 (15–16),z3: 17 (17–18), z4: 18 (16–20), z5: 14 (14–15), Z4: 34 (31–38), Z5: 53 (50–58), s4: 23

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Journal of Natural History 1735

(21–25), s6: 25 (22–28), S2: 30 (27–32), S4: 29 (28–32), r3: 20 (19–22), R1: 22 (20–25),JV5: 46 (45–48). SgeIV 20 (18–22); StiIV: 27 (24–30); St IV: 45 (42–48). Four pairsof dorsal solenostomes (gd2, gd6, gd8 and gd9). Chaetotactic formula of genus II:22/0, 2/01. Considerable variability has been reported in the literature for the num-ber of teeth on the fixed digit of the chelicera, ranging from one tooth (Chant andYoshida-Shaul 1987; Denmark 1992) to four teeth (Ragusa and Swirski 1978; Chantand Yoshida-Shaul 1987). Specimens collected in Spain have two teeth in addition tothe pilus dentilis.

Typhlodromus (Typhlodromus) thuriferus Ferragut, sp.nov.(Figures 1, 2)

Diagnosis

Medium-sized phytoseiid mite weakly sclerotized, with elongated palps reaching tarsusof leg I. Dorsal setae short, S4 similar in length to Z4, r3 similar to R1 and Z5 1.5 timeslonger than Z4. With three pairs of dorsal solenostomes, gd2, gd6 and gd9. Peritremerelatively short reaching level between setae z2 and z3. Ventrianal shield pentagonal,with three pairs of preanal setae and without preanal pores. Chelicerae with three orfour teeth on fixed digit and one tooth on movable digit. Calyx of spermatheca cup-shaped with the atrium inserted at the base of calyx. Genu II with seven setae. Threemacrosetae on leg IV.

Female (Figure 1)Measurements of holotype followed by the respective ranges of nine paratype femalesin parentheses.

Dorsum (Figure 1A). Dorsal shield suboval, weakly ornamented on podonotum,except for lateral margins, opisthonotum strongly reticulated 325 (325–352) long and164 (154–170) wide (at level of s4). With three pairs of dorsal solenostomes: gd2,gd6 and gd9. Seventeen pairs of short, smooth and acute setae on dorsal shield (dorsalsetal pattern 12A:7A according to Chant and Yoshida-Shaul 1991) except Z5, which isslightly serrated. Measurements of dorsal and sublateral setae as follows: j1 17 (17–20);j3 18 (16–20); j4 12 (11–13); j5 12 (12–13); j6 14 (13–15); J2 15 (13–15); J5 5 (5–6); z2 13(12–16); z3 16 (14–17); z4 14 (14–16); z5 12 (11–14); Z4 18 (18–20); Z5 28 (27–31); s417 (16–18); s6 17 (16–18); S2 18 (17–19); S4 21 (18–22); r3 19 (18–21); R1 18 (16–20).Ratio S4/Z4: 1.1; ratio Z5/Z4: 1.5; ratio r3/R1: ≈1.0. Seta r3 blunt, seta R1 pointed.

Peritreme. Reaching the level of seta z2 or between z2 and z3.

Venter (Figure 1B). Sternal shield smooth, with two pairs of setae and two pairsof lyrifissures. Posterior margin concave. Setae st3 and st4 on metasternal platelets.Distances between setae st1–st1: 46 (44–48), st2–st2: 55 (53–56), st3–st3: 68 (65–71),st1–st3: 35 (32–36). Genital shield narrower than ventrianal shield, 68 (68–77) wide.Four slender platelets between the genital and ventrianal shields. Ventrianal shieldsmooth, with three pairs of setae (JV1, JV2, ZV2), without preanal solenostomes andwidest at level of ZV2. Anterior margin convex, lateral margins concave behind ZV2;98 (98–110) long, 78 (78–92) wide. Bacillary platelets near the anterior corners. Fourpairs of setae (including JV5) and four pairs of poroids flanking ventrianal shield;

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1736 F. Ferragut and E.A. Ueckermann

Figure 1A–E. Typhlodromus (Typhlodromus) thuriferus sp. nov. Female. (A) Dorsal view. (B)Ventral view. (C) Spermathecae. (D) Chelicera. (E) Leg IV.

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Journal of Natural History 1737

JV5 smooth and pointed, 29 long (28–30). Bacillary or fusiform primary metapodalshield, 26 (25–30) long.

Spermatheca (Figure 1C). Calyx of spermatheca cup-shaped, 15 (15–17) long, withsclerotized walls. Atrium located at base of calyx.

Chelicerae (Figure 1D). Fixed digit with three teeth distal to pilus dentilis; movabledigit with one tooth, length 26 µm.

Legs (Figure 1E). Measurements of legs and palps as follows: leg I 312 (310–330);leg II 262 (260–280); leg III 270 (248–276); leg IV 350 (330–354); palp: 186 (182–196).Genu II with seven setae. Chaetotactic formula of genus II: 22/0, 2/01. Threemacrosetae on leg IV with bulbous tips; Sge IV: 22 (21–23); Sti IV: 30 (29–33); StIV:46 (46–54).

Male (Figure 2)Measurements with mean followed by range of seven male paratypes.

Dorsum (Figure 2A). Oval dorsal shield with lateral margins irregular and withlobed projections, 272 long (268–280), 159 (154–166) wide at the level of s4. Dorsalsurface striated and reticulated, especially on opisthoscutum. With three pairs of

Figure 2A–C. Typhlodromus (Typhlodromus) thuriferus sp. nov. Male. (A) Dorsal view. (B)Chelicera. (C) Ventrianal shield.

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1738 F. Ferragut and E.A. Ueckermann

solenostomes, gd2, gd6 and gd9 and 19 pairs of short, pointed and smooth setae.In one specimen seta R1 is inserted off the dorsal shield. Measurements of setae j1:16 (14–18); j3: 16 (15–18); j4: 11 (10–12); j5: 12 (11–12); j6: 14 (13–17); J2: 13 (12–16);J5: 5; z2: 12 (11–14); z3: 15 (14–16); z4: 14 (14–15); z5: 12 (12–13); Z4: 21 (20–23); Z5:28 (26–31); s4: 16 (16–17); s6: 16 (15–17); S2: 17 (17–18); S4: 19 (17–20); r3: 16 (14–18);R1: 16 (15–16).

Peritreme. Short, reaching to level of seta z3 or between z3–z4.

Venter (Figure 2C). Ventrianal shield almost smooth, with scanty striae anterior topreanal setae; 105 long (102–112) and 137 wide (134–142). With three pairs of preanalsetae and without preanal solenostomes. Seta JV5 smooth, 18 long (17–18) (ventralsetal pattern JV: ZV according to Chant and Yoshida-Shaul 1991).

Chelicerae (Figure 2B). Fixed digit 22 µm long, with three or four teeth; mov-able digit unidentate. Spermatodactyl attenuated distally; margin of shaft bears amembranous border.

Legs. Measurements of legs and palp: leg I: 261 (258–265); leg II: 233 (225–240); legIII: 234 (230–240); leg IV: 325 (300–348); palp: 175 (160–186). Macrosetae present ongenu, tibia and tarsus IV; Sge: 17 (16–18); Sti: 27 (25–28); Sta: 42 (40–44).

Type material

Holotype female, seven paratype females and two paratype males from Juniperus thu-rifera L. (Cupressaceae), Prádena, Segovia Province, Spain; geographic coordinates41◦07′19′′ N, 3◦42′22′′ W, 1145 m a.s.l., collected on 27 October 2000. Eleven paratypefemales and three paratype males from Juniperus thurifera, Puebla de Valverde, TeruelProvince (40◦10′14′′ N, 1◦01′07′′ W), 1500 m a.s.l., collected on 1 May 2008. Nineparatypes females and two paratypes males from Juniperus thurifera, Pedraza, SegoviaProvince (41◦08′14′′ N, 3◦49′19′′ W), 1115 m a.s.l., collected on 28 July 2008. Holotypeand some paratypes were deposited at Museo Nacional de Ciencias Naturales,Madrid, Spain (MNCN). Reference number of holotype: MNCN 20.02/17118; otherparatypes: MNCN 20.02/17121, MNCN 20.02/17124, MNCN 20.02/17125. Otherparatypes are deposited in the Natural History Museum, London, UK, under the ref-erence collection ENT 2011-208 and in the National Collection of Arachnida (NCA),ARC-Plant Protection Research Institute, Pretoria, South Africa.

Etymology

The specific name thuriferus refers to the host plant from which all the specimens werecollected.

Remarks

Typhlodromus (Typhlodromus) thuriferus sp. nov. belongs to the pyri-species group(Chant and Yoshida-Shaul 1987) or longipalpus-group (Denmark 1992) character-ized by having three pairs of preanal setae on ventrianal shield. It closely resemblesT. (T.) leptodactylus Wainstein, 1961 in having palps that are distinctly long in rela-tion to idiosoma and legs. However, it can be distinguished from T. (T.) leptodactylusin the shape of the spermatheca, without a long neck between the atrium and calyx,

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Journal of Natural History 1739

and by having only three pairs of dorsal solenostomes (gd8 absent). Other eightspecies in the subgenus have also three pairs of dorsal solenostomes: T . (T.) bac-cettii Lombardini; T. (T.) confusus Narayanan, Kaur and Ghai; T. (T.) kykladiticusPapadoulis and Emmanouel; T. (T.) oresibious Papadoulis, Emmanouel and Kapaxidi:T. (T.) pentelicus Papadoulis and Emmanouel; T. (T.) phylaktioticus Papadoulis andEmmanouel; T. (T.) pyri Scheuten and T. (T.) tubifer Wainstein. However, all of themhave four pairs of preanal setae on the ventrianal shield and so can be easily separatedfrom the new species.

The new species appears to be the predominant phytoseiid mite on Juniperus thu-rifera in Spain. The thuriferous or Spanish juniper, J. thurifera, is an Ibero-Magrebianpalaeo-endemism which has persisted since the Tertiary. At present this tree onlyoccurs in isolated areas of Spain, Morocco, Algeria and France under continental andcold Mediterranean climatic conditions (Quézel and Pesson 1980).

Key to species of the subgenus Typhlodromus Scheuten. Females

1. Ventrianal shield with three pairs of preanal setae . . . . . . . . . . . . . . . . . . . . . . . . 2Ventrianal shield with four pairs of preanal setae . . . . . . . . . . . . . . . . . . . . . . . . . 5

2. Dorsal setae short and acute; s4, s6, S2 and J2 between 15 and34 µm long;palps greatly elongated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3Dorsal setae longer, serrate and knobbed; s4, s6, S2 and J2 greater than 40 µmlong; palps not greatly elongated . . . . . . . . . T. (T.) armiger Ehara and Amano

3. Dorsal shield with three pairs of solenostomes (gd8 absent); spermathecawithout a neck between calyx and atrium . . . . . . . . . . T. (T.) thuriferus sp. nov.Dorsal shield with four pairs of solenostomes (gd8 present); spermatheca witha neck between calyx and atrium . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4. Dorsal setae shorter, S4, Z4 and Z5 between 19 and 25 µm long; ventrianalshield pentagonal . . . . . . . . . . . . . . . . . . . . . . . . . . . T. (T.) leptodactylus Wainstein. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= T. (T.) longipalpus Swirskiand Ragusa 1976 sensu Abbasova 1980 and Chant and Yoshida-Shaul 1987)Dorsal setae longer, S4 39, Z4 45 and Z5 72 µm long; ventrianal shield oval. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. (T) shoshae Palevsky et al.

5. Ventrianal shield with preanal pores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6Ventrianal shield without preanal pores . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26

6. Dorsal shield with three to five pairs of solenostomes; leg four with at leastone macroseta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7Dorsal shield with one pair of solenostomes; leg IV without macrosetae . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. (T.) roshanlali Narayanan and Ghai

7. Dorsal shield with three pairs of solenostomes . . . . . . . . . . . . . . . . . . . . . . . . . . . 8Dorsal shield with four or five pairs of solenostomes . . . . . . . . . . . . . . . . . . . . 10

8. Peritreme extends to level between z2 and z4; calyx of spermatheca bell-shaped; movable digit with one or two teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9Peritreme extends anterior to setae j1; calyx a short broad tube flared distally;movable cheliceral digit without teeth . . . . . T. (T.) confusus Narayanan et al.

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9. Peritreme extends to level of setae z4; fixed cheliceral digit with two or threeteeth, movable digit with two teeth; posterior margin of sternal shield straight. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. (T.) pentelicus Papadoulis and EmmanouelPeritreme extends to level of setae z2; fixed cheliceral digit with four teeth,movable digit with one tooth; posterior margin of sternal shield lobed . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. (T.) oresibious Papadoulis et al.

10. Dorsal shield with five pairs of solenostomes . . . . . . . . . . . . . . . . . . . . . . . . . . . 11Dorsal shield with four pairs of solenostomes . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

11. Solenostome gd9 absent; macroseta on leg IV acute . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. (T.) psidium Basha et al.Solenostome gd9 present; macrosetae on leg IV acute or knobbed or blunt . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

12. Leg IV with three macrosetae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13Leg IV with one macroseta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

13. Peritremes reaching seta j1; setae Z4 and S4 similar in length; j3 40, J2 39, Z570 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. (T.) nerminae El-Banhawy and KnappPeritremes reaching seta j3; seta Z4 shorter than S4; j3 24, J2 21, Z5 50 . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. (T.) wawerus El-Banhawy and Knapp

14. Peritremes extend to level between z2 and z3; fixed cheliceral digit with threeteeth; macroseta on leg IV acute . . . . . . . . . . . . . . . . . . . . T. (T.) corticis HerbertPeritremes extend to level between z3 and z4; fixed cheliceral digit with fourteeth; macroseta on leg IV knobbed or blunt . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. (T.) accessorius Kolodochka

15. Sternal shield with three pairs of setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16Sternal shield with two pairs of setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

16. Most or all dorsal setae smooth (except Z5) and shorter than distances to setaenext behind . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17All dorsal setae serrate and longer than distances to setae next behind, exceptfor setae j1, j4 and z5 which are smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. (T.) magdalenae Pritchard and Baker

17. Peritreme extends to level between j1 and j3; most dorsal setae long, all later-ally placed setae serrate as well as Z4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. (T.) neomagdalenae Zannou et al.Peritreme extends to level between setae z3 and z4; all dorsal setae muchshorter and smooth except for Z5 . . . . T. (T.) sapphicus Ragusa and Tsolakis

18. Setae J2 12–35, half or less distance to Z4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19Setae J2 49, two-thirds distance to Z4 . . . . . . . . . . . . . T. (T.) swirskii Denmark

19. Peritreme extends to level between j1 and j3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20Peritreme extends to levels between z2 and z3 or z3 and z4 . . . . . . . . . . . . . . 21

20. Calyx short, saccular, with atrium incorporated in calyx; setae j3 can be absentor present, one member of S5 can be present; fixed cheliceral digit with threeteeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. (T.) rarus (Wainstein)

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Calyx longer, tubular, atrium apparently not incorporated in calyx; setaej3 present and S5 completely absent; fixed digit with four teeth . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. (T.) kadii Kandeel and El-Halawany

21. Calyx of spermatheca saccular, with or without slight constriction betweencalyx and atrium; movable cheliceral digit unidentate or bidentate . . . . . . . 22Calyx with long, slender neck, funnel-shaped distally, thick-walled, atriumnodular; movable digit bidentate . . . . . . . . . . . . . . . . . . . . . . . T. (T.) andrei Karg. . . . . . . . . . . . . (= T. (T.) eucervix Karg and Edland 1998 sensu Denmark 1992)

22. Peritremes reaching to level between setae z2 and z3 . . . . . . . . . . . . . . . . . . . . 23Peritremes reaching to level between z3 and z4 . . . . . . . . . . . . . . . . . . . . . . . . . . 24

23. Setae j6 28, J2 36, z3–4, 32 and S2–4 46–48 µm long; ventrianal shieldapparently smooth . . . . . . . . . . . . . . . . . . . . T. (T.) pseudopyri Ehara and AmanoSetae j6 15–16, J2 18, z3–4, 15–20 and S2–4 24–28 µm long; ventrianal shieldreticulated . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. (T.) inhabilis Kuznetzov

24. Calyx of spermatheca saccular without constriction between it and atrium;setae St3 on round, oval or wider than long platelets . . . . . . . . . . . . . . . . . . . . 25Calyx of spermatheca saccular with slight constriction between it and atrium;setae St3 on round or oval platelets . . . . . . . . . . . . T. (T.) pritchardi Arutunjan

25. Posterior margin of sternal shield lobed; J2 17, S4 35, Z5 46; St3 on wider thanlong platelet . . . . . . . . . . . . . . . . . . . . . . . . T. (T.) moricola Ehara and ToyoshimaPosterior margin of sternal shield almost straight; J2 31, S4 50, Z5 64; St3 onoval or longer than broad platelet . . . . . . . . . . . . . . T. (T.) quercicolus Denmark

26. Dorsal shield with five pairs of solenostomes . . . . . . . . . . . . . . . . . . . . . . . . . . . 27Dorsal shield with three or four pairs of solenostomes . . . . . . . . . . . . . . . . . . . 28

27. Fixed cheliceral digit with six teeth, movable digit with two teeth . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. (T.) griekwensis SchultzFixed cheliceral digit with three teeth, movable digit with one tooth . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. (T.) atticus Swirski and Ragusa

28. Dorsal shield with three pairs of solenostomes . . . . . . . . . . . . . . . . . . . . . . . . . . 29Dorsal shield with four pairs of solenostomes . . . . . . . . . . . . . . . . . . . . . . . . . . . 32

29. Calyx of spermatheca with a neck . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30Calyx bell or cup-shaped, without neck . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . T. (T.) kykladiticus Papadoulis and Emmanouel

30. Genu II with eight setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31Genu II with seven setae . . . . . . . . . . . . . . . . . . . . . . T. (T.) baccettii Lombardini. . . . . . . . . . . . . . . . . . . . . . . . . . . . (= T. (T.) helenae Schicha and Dosse 1974 andT. (T.) oligadenus Athias-Henriot 1978 sensu Chant and Yoshida-Shaul 1987)

31. Calyx of spermatheca bell-shaped with short neck . . . . . T. (T.) pyri ScheutenCalyx of spermatheca funnel-shaped with long neck . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. (T.) tubifer Wainstein. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= T. (T.) ajselAbbasova 1980 sensu Chant and Yoshida-Shaul 1987 and probably. . . . . . . . . . . . . . . . . . . . T. (T.) phylaktioticus Papadoulis and Emmanouel 1990)

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32. Setae Z4 half length of Z5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33Setae Z4 0.6–0.75 length of Z5 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34

33. Posterior margin of sternal shield with a lobe . . . . . . T. (T.) laurae ArutunjanPosterior margin without a lobe . . . . . . . . . . . . . . . . . . T. (T.) knisleyi Denmark

34. Calyx of spermatheca variously shaped . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35Calyx a slender tube with undifferentiated atrium . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. (T.) moroccoensis Denmark

35. Calyx of spermatheca with proximal part tube-like or thick/sclerotized anddistal part strongly flared, funnel or cup-shaped . . . . . . . . . . . . . . . . . . . . . . . . 36Calyx tubular, relatively broad, slightly flared distally or not or short, bell- orcup-shaped with a short neck or not . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 38

36. Distal half of calyx of spermatheca cup- or funnel - shaped and neckthick/sclerotized or tube-like; fixed cheliceral digit with four teeth; peritremeextends to level of j3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37Proximal two-thirds of calyx tube-like and distal third flared; fixed digit withthree teeth; peritreme extends to level of z2 . . . . . . . . . T. (T.) personatus Karg

37. Distal half of the spermatheca cup-shaped; leg IV with one macroseta . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. (T.) cotoneastri WainsteinDistal half of the spermatheca funnel-shaped; leg IV with two macrosetae . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. (T.) bichaetae Karg

38. Calyx of spermatheca tubular . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39Calyx bell- or cup-shaped with a short neck or not . . . . . . . . . . . . . . . . . . . . . . 40

39. Calyx of spermatheca a short broad tube shorter than 20 long; Z5 shorter than55; peritreme extends to level of j1 . . . . . . . . . T. (T.) phialatus Athias-Henriot. . . . . . . . . . . . . . . . . . . (= T. (T.) ulex Evans 1988 sensu Edland and Evans 1998)Calyx tube-like, longer than 20; Z5 longer than 55; peritreme extends to levelof z2–j3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. (T.) athiasae Porath and Swirski. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (= T. (T.) siwa Elbadry1967, T. (T.) pelargonicus El Badry 1968, T. (T.) perbibus Wainstein andArutunjan 1968, T. (T.) hellenicus Swirski and Ragusa 1977 sensu Chant andYoshida-Shaul 1987, Denmark 1992 and Papadoulis et al. 2009 and probably. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. (T.) zaheri Denmark 1992)

40. Genu II with six or seven setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41Genu II with eight setae . . . . . . . . . . . . . . . . . . T. (T.) octogenipilus Kreiter et al.

41. Genu II with six setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. (T.) setubali Dosse. . . . . . . . . . . . . . . . . . . . . . . . . (= T. (T.) laurentii Ragusa and Swirski 1978, sensu

Chant and Yoshida-Shaul 1987)Genu II with seven setae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 42

42. Spermatheca with neck or constriction between calyx and atrium . . . . . . . . 43Neck or constriction absent . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45

43. Peritreme extends to level between j1 and j3 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. (T.) exhilaratus Ragusa. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (=T. (T.) exhilaratus exhilaratus; Chant and

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Yoshida-Shaul 1987 and T. (T.) exhilaratus americanus; Chant and Yoshida-Shaul 1987, sensu Papadoulis et al. 2009)

Peritreme extends to level between j3 and z2 . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44

44. Setae JV5 longer than Z5 and S4 longer than Z4; ventrianal shield reticulated. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. (T.) beglarovi Kuznetzov

Setae JV5 similar in length toZ5, S4 equal in legth to Z4; ventrianal shieldsmooth . . . . . . . . . . . . . . . . . . . . . T. (T.) olympicus Papadoulis and Emmanouel. . . . . . . . . . . . . . . . . . . . . . . . . . . . (probably = T. (T.) difficilis Kolodochka 2003)

45. Leg IV bearing one or two macrosetae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 46Leg IV with three macrosetae . . . . . . . . . . . . T. (T.) ernesti Ragusa and Swirski

46. Surface of peritreme uniformly stippled, reaching to level between setae . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . j3–z2 47Surface of peritreme not entirely stippled, with central core, extending to levelof setae z3–z4 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. (T.) klimenkoi Kolodochka

47. Setae Z5 65–70, Z4 38–39 µm long . . . . . . . . . . . . . . . . . T. (T.) tiliae OudemansSetae Z5 41–47, Z4 24–30 µm long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48

48. Posterior margin of sternal shield without a lobe; leg IV with one macroseta. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. (T.) morellensis Ferragut. . . . . . . . . . . . . . . . . . . . . . . (T. (T.) norvegicus Edland and Evans 1998. syn nov.)

Posterior margin of sternal shield with a lobe; leg IV with two macrosetae . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . T. (T.) mutatus Kolodochka

Acknowledgements

The authors wish to thank Drs. Jim McMurtry of Oregon State University, USA and FaridFaraji of MITOX, Amsterdam, The Netherlands, for their critical review of this paper. Partof this work was based upon research supported by the National Research Foundation (NRF),South Africa. Any opinion, findings and conclusions or recommendations expressed in the mate-rial are those of the authors and therefore the NRF does not accept any liability in regardthereto.

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