Zambedania sekhukhunensis, Camerik & Magowski & Hawkes & Ueckermann & Ochoa & Bauchan, 2016

Camerik, Anne M., Magowski, Wojciech Ł., Hawkes, Peter G., Ueckermann, Edward A., Ochoa, Ronald & Bauchan, Gary R., 2016, A New Species of Zambedania (Acari: Heterostigmatina: Pygmephoridae) from the Two Rivers Platinum Mine in South Africa and Notes on the Life-cycle of the Genus, Zoological Studies 55 (11), pp. 1-21 : 5-18

Zambedania sekhukhunensis sp. n. Camerik and Magowski ( Figs. 3-20 )

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Diagnosis: The female of Z. sekhukhunensis is most similar to that of Z. africana in having leg IV terminating in two claws, the stigmata being undivided and aggenital plate smooth (Ag). However, the species is uniquely different from Z. africana by the following combination of characteristics: stigmatal peritremes about half (vs. about one third) the length of the prodorsal shield (PdS); except for setae f, no other idiosomal dorsal setae on protuberances (vs. setae c, d, e, f and h1 on protuberances); setae e about 0.9x as long as setae f (vs 0.4 as long); h1 about 1.5x as long as h2 (vs. subequal in length); tibiotarsal solenidion ω is stemmed and about 1.3x the length of Φ1, 2 (vs. sessile and 1.7x); TaII: pv’, pv” barbed (vs. feathered).

D e s c r i p t i o n: (measurements- Table 1). Female. ( Figs. 3-9 , Table 2).

Gnathosoma ( Figs. 4A, B ). Orientation: hypognathous; shape ovoid. Cheliceral setae page 5 of 21

(ch1 smooth, ch2 barbed) setiform, subequal in size; ch 1 located antero- mediad of ch 2. Palpal supracoxal setae (pp) present. Dorso-median apodeme prominent. The antero-laterad pedipalpal segments are fused, bearing smooth setae dFe and dGe. Subcapitulum (= infracapitulum) with a pair of slender, simple, smooth setae (su) and a mushroom-shaped accessory setigenous structure (ass) located anteromedially and laterally flanked by a clavate, non-striated solenidion.

Pharyngeal system ( Figs. 4C ) subdivided into three striate, muscular pumps; in ventral view extending from the gnathosomal foramen to beyond apodeme 2. The first (anterior) pump is suboval, rather short and is immediately attached to the second pump. The second is narrow, rectangular and the longest of the three pumps. A short stretch of pharyngeal tube connects the second to the third pump, which is the widest and sclerotized at its anterior and posterior edges.

Idiosomal dorsum ( Figs. 3A , 9A ). Cuticle strongly sclerotized with uniform sparse punctate sculpturing. Prodorsal shield (PdS) with long, slitlike, obliquely placed stigmata reaching its anterior edge; capitate sensillae (sc1) with tiny barbs ( Figs. 3A , 9B ) are placed in circular bothridia; two pairs of well-developed tapered, setiform, barbed setae located posteriad of the stigmata; of these, vertical (v2) anterolaterad and shorter than the long and more robust scapular setae (sc2). Opisthosoma elliptical, five segmented with four dorsal segments: C, D, EF (coalesced) and H; the fifth segment, pseudanal shield (Ps) located caudoventrally. All segments bear two pairs of setae, except D which has one pair only. All dorsal setae very long, tapered, setiform and barbed, except setae v2 which is similar in form to the other dorsal setae but less than one third as long. Only seta f on a short protuberance. No cupules observed.

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Idiosomal venter ( Fig. 3B ). All ventral setae short (at least half the size of dorsal opisthosomal setae) and slightly barbed. Ap. 1, 2 complete, well-sclerotized, originating at the anterior margin of coxisternal plates I and II respectively. Ap. 1 forming an Y-shaped juncture with prosternal apodeme (appr). Ap. 2 slightly oblique, originating from the anterior base of legs II, extending across plates I, II and uniting postero-medially with the appr. The appr is incomplete, not fused with the well developed and strongly sclerotized sejugal apodeme (apsej). Ap. 3, 4 complete, anteriorly arched, medially joined to the poststernal apodeme (appo). Distribution of coxisternal setae typical for the genus. Setae 2b and 4b are the longest coxisternal setae. Ps with three pairs of setae,

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ps 1, 2, 3, of which ps 2 is the longest pseudanal seta.

Legs ( Figs. 5 - 8 ). The first two pairs of legs are subequal in length; legs III, IV are distinctly longer. All tectal leg setae are setiform, tapering, distally acute and more or less barbed, pilose or one-sided barbed with long barbs (feathered, see Fig. 9 C, D ).

Leg I. ( Figs. 5A, B ; 9C, D ). Tr: v’ <Fe v”; Fe: long l’> v”, d strongly barbed longest seta on segment; Ge: l’ strongly barbed, ~ Fe d; v’> l”> v”. TiTa with a long and strongly sclerotized, single, longitudinally ribbed apical claw closing into a butterfly-shaped counterpiece made up of the setae u’ and u” ( Fig. 9D ). Claw with small auxiliary blade located inwards near its base, and similarly small blunt protrusion even more proximally at its

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base. The “lock” on the dorsal side of tarsal part of the segment at the base of the flexed claw is well exposed in figure 9C. Barbed seta k <d. Setae v’ ~ v”, both feathered; l’ ~ l”, both barbed; feathered pv’ <pv” typically brush-like barbed at distal end ( Fig. 5B ); pl’ <pl” either barbed. Seta s is situated anterolaterad of the claw, but difficult to observe with a light microscope because of the accumulation of setae there. However s is clearly visible in figure 9D and easily observed on the larva and male. Three striated solenidia: stemmed solenidion ω> Φ 1, 2. Striation of Φ 2 wider apart than both of Φ 1 and ω. Eupathidia on pinnaculi (p’ missing): p”, ft’ <ft”. Distalmost seta on the verticil tc’ <tc”.

Leg II ( Fig. 6 ). Tr: barbed v’ <barbed Fe v”; Fe: l’> v”; d longest seta of leg; Ge: l’ ~ l” ~ v’ with strongly sclerotized annulus at its base; Ti: d shortest seta on segment, close to slim striated Φ, feathered l’ <v’ <barbed v”; Ta: barbed pl”> barbed tc’; tc’> smooth tc”; pv’ <barbed pv” ~ slim, smooth u. Solenidion ω stout, club-shaped.

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Terminal two simple claws with a fan-shaped empodium.

Leg III ( Fig. 7). Tr: one-sidedly barbed v’ ~ Fe v’, Fe d longest seta on the leg; Ge: l’> v’; Ti: Φ stout, club-shaped; slightly barbed l’ ~ barbed d; smooth v’<barbed v”; Ta: barbed pl” longest seta on verticil, barbed tc’ ~ smooth tc”; smooth pv’ <barbed pv”, distal-most barbed u ~ pv’. Terminal pair of simple claws with a fan-shaped empodium.

Leg IV ( Fig. 8). Tr: v’ ~ Fe v’, each barbed; Fe: robust, barbed d with strongly sclerotized annulus, longest seta on leg; Ge: barbed v’ <Ti v’; Ti: smooth l’ ~ v’ ~ v”, both barbed; d long, barbed with strongly sclerotized annulus and tiny solenidion Φ in sacculus at its base. Ta: pl” pilose; whipped tc’> smooth tc”, pv” ~ pv’ both barbed; and pv’> barbed u’.

Etymology: The specific name sekhukhunensis refers to the collecting locality of the spider hosts, Sekhukhuneland, near Lydenburg, Mpumalanga, South Africa.

(A)

(B)

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Male ( Figs. 10-15 , Table 3).

Gnathosoma ( Fig. 10C, D ). Gnathosomal capsule of the male much smaller than that of female. However, most of the external structures of the female capsule present on the male gnathosoma except that the mouthparts are absent. The stylophore extends anteriorly into a truncated-triangular distal edge bearing two pairs of tiny, sub-equal in size, smooth cheliceral setae: ch1 and ch2. Unlike the female, the male lacks palpi, palpal supracoxal setae (pp) and chelicerae. Subcapitulum with a pair of relatively long slender, simple setae (su); anteriad setigenous accessory structures (ass) laterally flanked by relatively long clavate, non-striated solenidia (sol).

Pharyngeal system: absent.

Idiosomal dorsum ( Figs 10A , 11A ). Males on average much smaller than females (range of lengths 172-186 vs 270-309) and with well sclerotized, sparsely punctate exoskeleton. Prodorsal shield (PdS) triangular, dome-shaped, without stigmata, tracheal system or bothridia; verticals (v2) antero-mediad of setiform short page 8 of 21

scapulars sc1 and long sc2. Opisthosoma with tergites CD fused, EF fused, and dorsodistally partly covers greatly reduced tergite H which coalesced with caudoventral pseudanal shield (Ps) to form HPs, the genital capsule. All dorsal setae barbed, except for smooth setae e, h1 and h2. Fused shield CD bearing two pairs of very long tapered, setiform barbed setae (c1, c2) anteriorly and a pair (d) of similar setae postero-laterally. Shield EF with setae e less than half as long as barbed f, covering HPs. Shield H ( Fig. 11A ) with two pairs of setae: anteriormost setae (h1) smooth, short, thorny and pointed, h2 setae strongly modified into button-like structures in suckershaped rings. Membranous copulatory flanges border the H shield posteriorly.

Idiosomal venter ( Figs 10B , 11B ). Compared with the long clearly barbed dorsal setae the ventral setae short, smooth or more or less barbed. Coxisternal plates: anterior and posterior edges of ventral shields smooth, not subdivided, and widely curved. The setae are distributed as in the females. Ap1 very short, seemingly a mere widening of the

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anterior part of the prosternal apodeme (appr). The appr extends uninterruptedly caudad to form a Y-shaped juncture with complete ap2, to continue posteriorly towards the apsej. Ap3, 4 complete, anteriorly arched; ap3 medially concavely and ap4 convexly joined to completely formed poststernal apodeme (appo). Appo meeting posteriorly of setae 4a with ap5 from the posterior condyle of Tr IV. Setae 4b proximal-most on coalesced plate HPs. On the genital capsule modified pseudanal setae ps2 on a slight elevation, medio-laterad of and very close to ps3. Setae ps1 ( Fig. 11A ) probably implanted medio-anteriorly of ps2, 3 and covered by copulatory flanges, on either side of the folded aedeagus. No cupules seen.

Legs. Leg I ( Fig. 12). All tactile leg setae setiform, tapering, distally acute and barbed.

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Trochanter length v’ ~ Fe v”; Fe d setiform and longest seta of leg I; l’> ½ d, v” ~ ⅔ l’. Genu v’ ~ l”> v”, l’ longest seta of verticil. All tibial setae barbed on one-side, except smooth l’. Setae v’ ~ l” <k; v” longest, d shortest seta on verticil. Solenidion slim Φ 1> Φ2. Tarsal solenidion ω 2 proximal-most on the verticil, close to and much shorter and thinner than robust ω 1. One-sidedly barbed pv” ~ pv’, pl” ~1½ pl’. Seta s ~ ω 1. Setae u’ <u”. Eupathidia (ξ) ft’ ~ ft”; tc’ <tc”; p’ <p”. Tectals and prorals on pinnaculum, dorso-laterad of single claw.

Leg II ( Fig. 13 ). Trochanter: ”v’ <Fe v”,” the latter barbed on one side. Fe l’ barbed on one side, lacking on right leg, but present on the left, shortest seta on segment, d the longest. Genual smooth l” ~ slightly barbed l’ <one-sidedly barbed v’. Tibial one-sidedly barbed l’ longest seta on verticil, v’> v”,

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d shortest seta, closest to solenidion. Tarsus with robust solenidion ω proximal-most on segment; pl” longest seta, on protuberance; barbed tc’ <smooth tc”, pv’ <one-sidedly barbed pv”, u’ <to u”, both one-sidedly barbed. Cordate empodium with medial fold between a pair of simple claws,

Leg III ( Fig. 14 ). Third leg very much like the second leg; trochanteral setae barbed v’> barbed Fe v’. Femur with longest, robust, barbed seta d extending to tarsus. Genu with barbed setae v’ ~ l’. Tibial d closest to solenidion Φ and <one-sidedly barbed l’. Seta v ’ ~ v”, both slightly barbed; similar to leg II, leg III bears a robust tarsal solenidion ω;

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one-sidedly barbed pl” longest seta on segment, slightly barbed tc’> smooth tc”, slightly barbed pv’ ~ one-sidedly barbed pv”; barbed u’ <smooth u”; as in leg II terminating in a pair of simple claws with a cordate empodium in-between.

Leg IV ( Fig. 15 ). Leg IV modified; trochanteral seta barbed v’> Fe v’; Fe d slightly barbed ~ barbed v’. Genu: barbed v’> Fe v ’; l’ absent. Tibia: short Φ close to smooth d; smooth l’ <d; one-sidedly barbed v’ longest on segment, smooth v” the shortest. Tarsus very short with modified claw that seems to fit in a hooked claw-like structure u’. Setae pl”, pv” and pv’ short and smooth; very long

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tc’ ~ 1½ tc”, both setae barbed. Seta u” inserted laterad of claw.

Larva ( Figs 16-20 , Table 4).

Gnathosoma ( Figs. 17A, B ). The gnathosoma about the same size and shape as that of the adult female. Dorsally the stylophore terminates in a trapezoid antero-distal edge. The pedipalps situated antero-laterad and subdivided into four segments, palptrochanter, fused palpfemoro-genu with smooth setae dFe and dGe, and palptibiotarsus. The palptrochanter fused with the stylophore. The palptrochanter and palpfemoro-genu connected by membranes as are the palpfemoro-genu and the palptibiotarsus. The palptibiotarsus, laterad of the stylophore clearly page 13 of 21

subdivided into three parts. The distal claw medially with thorn-like structures. Stylophore with very short, blunt peglike ch 1 setae and slightly postero-laterad thereof smooth setiform ch 2, which is about three and a half times the length of ch 1. Palpal supracoxal setae pp absent. Subcapitulum with a pair of slender, simple, smooth subcapitular setae (su); antero-mediad a mushroom-shaped setigenous accessory structure (ass) and laterad at its base a clavate, non-striated solenidion (sol).

Pharyngeal system ( Fig. 17C ) similar to that of the adult female situated in the propodosomal area. The pharyngeal pumps shaped similarly to those of the adult female, but more compact and very thin.

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Idiosomal dorsum ( Fig. 16A ). The larval exoskeleton very lightly sclerotized and without clearly defined tergites and sculpture. The PdS with three pairs of tapered setiform setae: one page 14 of 21

pair of short, smooth verticals (v) anterolaterad of barbed scapulars sc1, followed posterolateral by a very long pair sc2. Bothridia, stigmata or podocephalic canals absent. Opisthosomal tergites fused in pairs: CD (partly), EF, and dorsoventrocaudal HPs. All dorsal opisthosomal setae very long and barbed. Segment C is subdivided into three shields, the median one coalesced with plate D bearing setae c1 and two separate laterals with c2. Setae c1 are situated mediad of c2 and

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anteriad of d. Setae d on CD and f on plate EF implanted on protrusions. Setae e laterad of seta f on segment EF. On segment HPs setal pair h1 positioned slightly antero-mediad of h2 on the dorsal side, and the pseudanal ps1, 2, 3 ventrocaudad.

Idiosomal venter ( Fig. 16B ). Coxisternal plates I and II lightly sclerotized bearing two pairs of setae on each plate (1a, 1b and 2a, 2b respectively). Anterior edges of the plates form apodemes (ap) 1 and 2, medially separated from each other and do not form the prosternal (appr)

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apodeme. Sejugal apodeme (apsej) absent. CX III with two pairs of setae (3a, 3b). Ap3 and poststernal apodeme (appo) absent. In place of appo, some creases between CX III plates present. All ventral setae short, tapered and smooth.

Legs. Leg I ( Fig. 18 ). Femur: barbed d longest seta on segment, slightly barbed l’> smooth v”, both on pinnaculum. Genu: all setae barbed, except smooth v” <v’; l’ (on pinnaculum) longest seta and l” the shortest; all tibial setae on pinnaculi and smooth except feathered v’: v”> l’> l”> d ~ k. Striation of solenidion Φ wider than of ω (both

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on pinnaculi). Tarsal setae all smooth except little barbed pl’; pl’ ~ pl”; pv”> pv’; ft’~ ft”> s> u’> u”; ft’ <tc’ <tc”. Setae pv’, pl”, ft’, tc’ and tc” on pinnaculi. On tarsus two distal claws present (only one on the right leg- artefact); no empodium between the claws.

Leg II ( Fig. 19). Femur: slightly barbed l’> smooth v” (both on pinnaculi), d longest on segment and barbed. Genu: v’ ~ l’, both one-sidedly barbed; slightly barbed l” shortest seta. Tibia: one-sidedly barbed l’> slightly barbed d; pilose v’ <one-sidedly barbed v”; solenidion j <Ta ω. Tarsus: smooth pl”> smooth tc’> smooth tc”; slightly barbed pv’> smooth pv”; slightly barbed u’> u”. Solenidion ω large. Two claws with a spatulate empodium in between. Tibial setae d, v’, v”, Φ and tarsal pl”, tc’ and ω on pinnaculi.

Leg III ( Fig. 20). Femur: barbed v’ <long one-sidedly feathered d (on pinnaculum). Genu: smooth v’ <one-sidedly barbed l’. Tibia: all setae one-sidedly barbed: l’ <v’ <v” <d which is closest to solenidion j. Tarsus: smooth pl” <smooth, robust page 16 of 21

tc’> smooth tc”; slightly barbed pv’> smooth pv”> smooth u’ ~ u”. Two claws with a fan-shaped empodium in between.

Material examined: Holotype adult female, 28 paratype adult females, 5 paratype adult males and 1 paratype larva. Females primarily aggregated in and around the fovea ( Fig. 2 ) of the cephalothorax and around the coxae, near the spinnerets and anus of spiders; in addition, females found with associated males and larva in and around spider’s nests.

Host: Harpactirella overdijki Gallon, 2010 ( Araneae : Theraphosidae ). Specimens were found on 4 females and 2 immature spiders. Site: most spiders were collected under a large tree ( Fig. 1 ); 200m transect; midpoint of transect: 24.9086 S, 30.09971 E; altitude 945 m (range 938-950 along transect). Locality: Two Rivers Platinum Mine, 38 km SE of Burgersfort, 40 km WNW of Lydenburg, Sekhukhuneland, Limpopo Province, South Africa. Dates: 25 March - 25 April 2008, 3-7 November 2008, 19 April 2010, 12-13 January 2011 and

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7-11 February 2011. Collectors: Peter Hawkes, Jonathan Fisher and assistants: Faith Kgafane, Katarina Jovanovic, Maphilisi Zulu, Matimba Baloyi, Michael Pierce, Nkosinathi Babu and Philippa Wing.

Deposition of types: Holotype female, 4 paratype females, 1 paratype male and 1 paratype larva, deposited in the USNM mite collection, Beltsville , Maryland, USA; 6 paratype females and 1 paratype male in the collection of the University Adam Mickiewicz University in Poznań, Poland;

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4 paratype females and 1 paratype male in the collection of the Zoologisches Institut und Museum of Universität Hamburg, Germany; 4 paratype females and 1 paratype male in the Muséum d’ Histoire naturelle, Département des Arthropodes et d’ Entomologie, Genève, Switzerland; 10 paratype females and several non-type female specimens and 1 paratype male in the Arachnida Collection of the ARC Plant Protection Research Institute, Pretoria, South Africa.

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Figures

Fig. 3. Zambedania sekhukhunensis sp. n. Female holotype. Idiosoma: A- dorsum, B- venter; sculpture omitted.

Fig. 4. Zambedania sekhukhunensis sp. n. Female holotype. Gnathosoma A- dorsum, B- venter, C- pharyngeal pumps.

Fig. 5. Zambedania sekhukhunensis sp. n. Female holotype. Leg I A- (left): antaxial view, B- tibiotarsus (left): paraxial view.

Fig. 6. Zambedania sekhukhunensis sp. n. Female holotype. Leg II (right): dorsal view.

Fig. 9. Zambedania sekhukhunensis sp. n. Female. A- dorsal view; B- caput of bothridial seta, dorsal view; in front “pilose” setae; C- Leg I (right). Note TiTa in dorsal view. At distal end “turret” carrying claw with at its base dorsal pinnaculum of the “locking system”, flanked by part of counterpiece. Eupathidia dorsad on pinnaculi. The cluster d and k associated with the three solenidia. Ventro-laterad near to counter piece seta s followed by feathered pv and v. D- detail distal end of left TiTa I in lateral view, claw and butterfly-shaped counter piece made up of setae u’ - u”, below seta s, followed by “feathered” primiventrals pv’ and pv” (photos by G. Bauchan and R. Ochoa).

Fig. 10. Zambedania sekhukhunensis sp. n. Male paratype. Idiosoma: A- dorsum, B- venter; gnathosoma: C- dorsum, D- venter; ascalebar for idiosoma, b- scalebar for gnathosoma; sculpture omitted.

Fig. 11. Zambedania sekhukhunensis sp. n. Male paratype. Genital capsule: A- dorsal view, B. ventral view.

Fig. 13. Zambedania sekhukhunensis sp. n. Male paratype. Leg II (right): dorsal view.

Fig. 14. Zambedania sekhukhunensis sp. n. Male paratype. Leg III (right): dorsal view.

Fig. 15. Zambedania sekhukhunensis sp. n. Male paratype. Leg IV (left): dorsal view.

Fig. 16. Zambedania sekhukhunensis sp. n. Larva paratype. Idiosoma: A- dorsum, B- venter.

Fig. 17. Zambedania sekhukhunensis sp. n. Larva paratype. Gnathosoma: A- dorsal view, B. ventral view, C- pharyngeal pump system.

Fig. 18. Zambedania sekhukhunensis sp. n. Larva paratype. Leg I (right): dorsal view.

Fig. 2. Zambedania sekhukhunensis sp. n phoretic female mites aggregated in and around the fovea of the cephalothorax and around the coxae of Harpactirella overdijki Gallon, 2010 (Aranaeae: Theraphosidae) (photo by I. Engelbrecht).

Fig. 1. Soon to be exploited North Open Pit (NOP) area at the Two Rivers Platinum Mine at Leydenburg, South Africa (photo by P. Hawkes).