Acari: Mesostigmata: Laelapidae - BioOne

Systematic & Applied Acarology 20(6): 707–720 (2015) http://dx.doi.org/10.11158/saa.20.6.12 Article

ISSN 1362-1971 (print) ISSN 2056-6069 (online)

http://zoobank.org/urn:lsid:zoobank.org:pub:9163E164-01E2-429E-8335-5FA3895A4B95

A new species of Scissuralaelaps (Acari: Mesostigmata: Laelapidae) from millipedes in the Philippines OWEN D. SEEMAN1* & GERD ALBERTI2 1

Queensland Museum, PO Box 3300, South Brisbane, Qld, 4101, Australia Zoologisches Institut und Museum, Ernst-Moritz-Arndt-Universität Greifswald, J.-S.-Bach-Str. 11/12, 17489 Greifswald, Germany *Corresponding author: [email protected] 2

Abstract A new species of Scissuralaelaps (Acari: Laelapidae: Iphiopsidinae), S. huberi sp. nov., is described from the millipede Acladocricus sp. (Spirobolida: Rhinocricidae) in the Philippines. The new species is close to S. bipartitus Ishikawa, which also has males with heavily sclerotised anterior and lateral margins of the peritrematal and metapodal shields. Key words: Iphiopsidinae, Myriapoda, Spirobolida, Cebu, sexual dimorphism

Introduction Scissuralaelaps Womersley (Mesostigmata: Laelapidae: Iphiopsidinae) is a genus of ten species found on julid, spirobolid and spirostreptid millipedes in the Philippines and Papua New Guinea (eight species) and burrowing cockroaches in Queensland, Australia (Womersley 1945; Rosario 1981; Ishikawa 1988; Fain 1992; Halliday 1993). The nature of the relationship between the mite and their hosts is unknown, although Halliday (1993) speculated those associated with cockroaches could be predators in their burrows, based on the form of their chelicerae and widespread presence of astigmatan mites found with the cockroaches. Here we describe a new species of Scissuralaelaps from the Philippines, from where three other species have already been described: Scissuralaelaps philippinensis Rosario from Mindanao Island on an undetermined millipede, and Scissuralaelaps bipartitus Ishikawa and Scissuralaelaps breviseta Ishikawa, also from Mindanao Island, but on the millipedes Acladocricus philippinus Wang and Trigoniulus sp., respectively (Rosario 1981; Ishikawa 1988).

Materials and methods Mites were preserved in 80% ethanol, cleared in Nesbitt’s fluid, and slide-mounted in Hoyer’s medium. Specimens were examined, measured and drawn with a Nikon Eclipse 80i equipped with DIC optics, stage-calibrated ocular micrometer and a drawing tube. Specimens for scanning electron microscopy were dehydrated with Hexamethyldisilazane, sputter-coated with gold, and examined in a Hitachi TM-1000 Tabletop microscope. Chaetotaxy follows Evans (1963) and Evans and Till (1965). Beaulieu (2009) was used as a general template for a laelapid description. Distances between setae were measured between their bases. Measurements are presented mostly as ranges and in micrometres. © Systematic & Applied Acarology Society

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Taxonomy Scissuralaelaps Womersley, 1945 Type species: Scissuralaelaps novaguinea Womersley, 1945: 225. By original designation.

Diagnosis. Female sternal shield divided longitudinally, either completely or almost so. Tibia I lacking seta av2; genu IV with pl2. Gnathosoma with labrum well developed, with fimbriate lateral margins, extending well beyond tips of corniculi; internal posterior hypostomatic setae h3 subequal to or shorter than external posterior hypostomatic setae h2. Associated with julid, spirobolid and spirostreptid millipedes or blaberid cockroaches. Scissuralaelaps huberi sp. nov. (Figs. 1–33) Material examined. Holotype, ♀, near Busay Cave, Moalbal, south-west Cebu, Republic of the Philippines, 9°54′57.5′′ N 123°26′13.2′′ E, ex Acladocricus sp. (Spirobolida: Rhinocricidae; Fig. 31), 4 Dec. 2011, S. Huber. Paratypes, 6 ♀♀, 4 ♂♂, same data as holotype. 2 ♀♀, 2 ♂♂, Campus of San Carlos University, Cebu, Republic of the Philippines, ex large millipede, Aug. 1986, G. Alberti. Holotype and paratype female and male deposited in San Carlos University, Cebu, Republic of the Philippines. Paratype male and female deposited in Senckenberg-Museum, Görlitz, Germany. Remaining paratypes deposited in Queensland Museum, Brisbane, Australia. Diagnosis. Dorsal shield with 60 pairs of short setae. Female sternal shield completely divided longitudinally; surface entirely reticulate. Endopodal shields absent. Female epigynal shield expanded posterior of setae st5. Peritremes extend past CxII. Female with anterior margin of gnathotectum smooth medially, denticulate laterally. Femur II ad1 a macroseta. Male with anterior margin of idiosoma extending over most of gnathosoma; separate sternogenital and anal shields; anal shield large; sternogenital shield not expanded posterior of setae st5; metapodal shields large with heavily sclerotised lateral margin that extends along peritrematal shields to anterior margin. Description. Female (n = 9) Dorsal idiosoma (Fig. 1). Dorsal shield length 730–830, width 460–500, suboval, reticulate marginally and submarginally, smooth areas medially, with 60 pairs of short (length 5–14) setae, no unpaired setae, with 21 pairs of pore-like structures (one lyrifissure; all others surrounded by lacunae), sigilla arranged in nine groups on either side of shield. Marginal setae short, in soft lateral cuticle. Ventral idiosoma (Figs 2–4). Tritosternum with pilose laciniae, length 145–152, essentially unfused (fused for 4–6 from bases), laciniae with a basal, irregular arched dorsal fringe of 50–60 denticles (Fig. 3), base trapezoidal (length 28–33). Presternal area weakly sclerotised. Sternal shield divided longitudinally, maximum length 185–200, width of each half at level of setae st2 60–65, maximum width of each half 100–105 at level of coxae II–III, ratio length/width of half sternal shield at st2 2.9–3.1; sternal shield entirely reticulate, with longitudinally elongated cells medially on each half; each shield section bearing short smooth slender setae st1–3 (26–33, 22–55, 22, respectively) and lyrifissures iv1–2. Setae st4 (20–29) and lyrifissure iv3 in soft cuticle; free endopodal shields absent. Epigynal shield (full length 220–245; 62–67 from level of st5 to posterior tip) with hyaline anterior margin, broadly rounded, reticulate throughout, with eight (one specimen with six, Fig. 4) large cells posterior to st5, narrowest width between coxae IV (45–50), and round posteriorly (maximum width 86–93), bearing setae st5 on margins and flanked by pair of lyrifissures iv5 on soft cuticle near st5. Exopodal plate surrounding posterior margin of coxae IV. External, primary (33–37 × 18–21) metapodal platelets oval, inner (17–23 × 5–13) metapodal platelets rectangular. Anal shield 708

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(length 105–120 × greatest width 125–140) subtriangular, reticulate, with post-anal seta (16–18) shorter than para-anal setae (25–28) and with posterolateral poroids; cribrum narrow; sigilla in two anterolateral pairs. Soft cuticle with ca. 30 pairs of setae, hypertrichous laterally, lengths 12–15. Peritrematal shield smooth, free posteriorly, narrowly fused to dorsal shield near z1, bearing 7–8 pore-like structures (one ip lyrifissure, two gland pores on shield and 4–5 pore-like structures within peritreme); peritreme extending anterior of coxa I, to level of z1.

FIGURE 1. Scissuralaelaps huberi sp. nov., female, dorsum. 2015

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FIGURE 2–4. Scissuralaelaps huberi sp. nov., female. 2, Venter; 3, Dorsal surface of the tritosternum; 4, Epigynal shield, showing variation in reticulation (scale bar as for Fig. 2).

Gnathosoma (Figs 5–10). Gnathotectum (Fig. 5) with anterior medial margin smooth, rounded; lateral margins denticulate. Subcapitular groove bearing six rows of denticles, preceded anteriorly by a smooth ridge devoid of teeth; rows 1–6 (anterior–posterior) with 10–13, 8–10, 11–12, 10–12 (one specimen with 6), 6–7, 4–5 denticles (Fig. 6). Anterior subcapitulum bearing: fringed internal 710


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FIGURES 5–10. Scissuralaelaps huberi sp. nov., female. 5, Gnathotectum (scale bar as for Fig. 6); 6, Subcapitulum and right palp; im = internal mala; lb = labrum; ss = salivary stylet; 7, Palp tarsus, detail; tarsal setae are numbered and shaded grey, tibial setae are indicated with a * if fine-tipped or ** if blunt-tipped and on small tubercles; 8, Left corniculus, ventral view, detail showing small sub-basal spine (scale bar as for Fig. 9); 9, Right palp, detail of dorsodistal spine of femur; 10, Chelicera, antiaxial view.

malae comprising thin internal processes that extend past tips of corniculi, and smaller, broad, anteriorly notched lateral processes (Fig. 6, im); large labrum, extending to mid-genu of palp (Fig. 6, lb); horn-like corniculi (55–60), extending to middle of femur of palp, with sub-basal ventral spine (Fig. 8). Hypostomal setae short and fine, h1 longer (36–40) than outer posterior setae h2 (20–22), 2015


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FIGURES 11–16. Scissuralaelaps huberi sp. nov., female. 11, Leg I, trochanter to tibia, dorsal view; 12, Leg II, ventral-anterolateral view; 13, Leg III, femur-tibia, dorsal view; 14, Leg IV, trochanter to tarsus, dorsal view; 15, Ambulacrum I; 16, Ambulacrum III, sclerotised parts in both ambulacral figures are shaded grey (dorsal) or dotted (ventral).

inner posterior setae h3 (14–15), and palpcoxal setae pc (19–20); setae h2 70–74 apart, and pc 19– 20 apart (Fig. 6). First cheliceral segment length 70–80; second cheliceral segment (155–160) ending in fixed digit, bearing an offset apical tooth and three teeth subapically, followed by a large tooth at

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the level of a short, setiform pilus dentilis, a row of six small teeth of irregular size, and a proximal tooth; dorsal seta small (10), posteriad dorsal lyrifissure; movable digit (61–65) bidentate, with long arthrodial corona (32–40) (Fig. 10). Palps (205–230) (Figs 6–7), setation of palp segments from trochanter to tarsus: 2-5-6-14-15; palp trochanteral setae simple; palp femoral setae d1–3 thickened, al truncate, pl simple, femur with dorsal spine (Fig. 9); palp genual setae al1 and al2 truncate, seta d3 spine-like and other setae simple; two apical blunt-tipped tibial setae on small tubercles; tarsus (Fig. 7) placed entirely ventrally, bearing apical cluster of short setae; apotele 2-tined, with tines tapered and flanked by a small cusp, ventral tine shorter (16 from fork) than dorsal tine (26 from fork, 31 from base). Legs (Figs 11–16). Excluding ambulacra, lengths of leg I 680–725, leg II 560–600, leg III 545– 560, leg IV 630–660. Tarsus I without claws, ambulacrum 54–55, stalk 20–21 long from base of ventral insertion; ambulacra II–IV with small claws, lengths (including claws and pulvillus, excluding short stalk) II–III 44–48, IV 46–48; opercula not well developed (Figs 15–16). Dorsal lengths of femora I 100–110, II 95–100, III 95–105, IV 115–130; genua I 105–110, II 90–100, III 75–80, IV 95–100; tibiae I 110–115, II 80–85, III 70–75, IV 90–95; tarsi I 185–190, II–III 140–150, IV 165–170. Setation of leg segments I–IV: Coxae 2-2-2-1. Trochanters 6-5-5-5, TrI al1, pd1 thickened, TrIV with ad1 thickened. Femora 12-11-6-6 or, as (al–ad/av, pd/pv, pl), I (2–3/1, 2/2–2), II (2–3/1, 2/2–1), III (1–2/1, 1/0–1), IV (1–2/1, 1/0–1), FeI ad1–3, al1, av1, pd2 thickened, FeII seta ad1 greatly thickened, FeIII with all setae except av1 thickened, FeIV with setae ad1–2, pd1 thickened. Genua 12-11-10-11, or I (2–3/1, 3/1–2), II (2–3/1, 2/1–2), III (2–2/1, 2/1–2), IV (2–2/1, 3/1–2), GeI pv1 thickened, GeII ad2–3, pv1 thickened, GeIV with setae pl2, av1 thickened. Tibiae 12-10-9-10, or I (2–3/1, 3/1–2), II (2–2/1, 2/1–2), III (2–1/1, 2/1–2), IV (2–1/1, 3/1–2), TiI without modified setae, TiII–III with seta pv1 thickened, TiIV with seta av1 thickened. Tarsus I same as for male (Fig. 22), with four short blunt-tipped setae (length 20) among dorsodistal cluster of weakly to strongly tapered setae of various lengths (15–45); small subapical sclerotised platelet with four setae: one tapered, one blunt-tipped, two tiny. Tarsi II–IV 18-18-18. Male (n = 6) Dorsal idiosoma. Dorsal shield (length 810–930 × width 550–620), similar in ornamentation and setation to female. Anterior margin of idiosoma extending over most of gnathosoma (Fig. 33). Caudal region of opisthosoma bends ventrally almost perpendicularly (Figs. 30, 33). Peritrematal shield fused to dorsal shield at level of CxI–II. Ventral idiosoma (Fig. 17). Sternogenital shield (355–400 long × 225–265 wide), bearing slender setae st1–5, st4 longest (39–60), other setae (25–35); reticulated throughout except near anterolateral margin; inverted V-shaped region between coxa IV demarcates anterior darker, coarsely punctate cuticle and posterior lighter, finely punctate cuticle. Genital opening in anterior concavity of sternogenital shield. Anal shield extensive (210–235 long × 200–245 greatest width) but bearing only para-anal setae (26–29) and postanal seta (20–22), reticulate anteriorly, smooth to lightly reticulate elsewhere; sigilla in two anterolateral pairs. Peritrematal and metapodal shields with heavily sclerotised lateral margins extending anteriorly around edge of dorsum, becoming thicker anteriorly; five pairs of pores in margins, one pore in cuticle laterad stigmata, and peritreme bearing 3–4 small pores. Peritrematal and exopodal shields fused; metapodal shield large, with eroded medial margins and one pair of sigilla. Peritreme extends to anterior margin of coxa II. Two pairs of small platelets between metapodal and sternogenital shield, one subrectangular, one elongate. Soft cuticle with three pairs of small (12–20) setae between anterior part of anal shield and metapodal shields; lateral cuticle hypertrichous, with 30–37 pairs of setae. Six pairs of pores in soft cuticle plus one pair of pores on posterolateral margin of anal shield.

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FIGURE 17. Scissuralaelaps huberi sp. nov., male, venter, tritosternum not shown (as in female).

Gnathosoma (Figs. 18–20). Gnathotectum (Fig. 18) with denticulate anterior margin. Subcapitulum and palps as in female, setae h1 34–37, h2 11–16, h3 12–15, pc 14–17, palp length 215–220, corniculi 52; chelicerae (Figs. 19–20) with fixed digit (34–36, from lyrifissure) bearing an offset blunt tooth near apex, three small teeth and a small pilus dentilus; cheliceral seta small (5–6);

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movable digit (38–40) with one tooth; spermatodactyl appears divided into two processes, curves upwards, length (56–64).

FIGURES 18–20. Scissuralaelaps huberi sp. nov., male. 18, Gnathotectum; 19, Chelicera, paraxial view; 20, Chelicera, antiaxial view.

Legs (Figs. 21–25). Chaetotaxy as in female. Lengths of leg I 805–825, leg II 660–700, leg III 660–715, leg IV 800–860. Modified setae: Trochanters – TrI with setae al1, pd1 thickened; TrII with setae al1, pl1 thickened; TrIII with setae ad1, al1 thickened; TrIV with setae ad1, al1, pv1 thickened. Femora – FeI with setae ad1–3, al1–2, av1, pd1–2 thickened; FeII with seta pv1 a large spine, setae av1, pv2 with bulbous base, setae ad1, pd2 thickened; Fe III with setae av1 with bulbous base, other setae thickened; FeIV with seta av1 with bulbous base, setae ad1–2, al1, pd1 thickened. Genua – GeI with setae al1–2 thickened; GeII with setae av1, pv1 with bulbous base, setae ad1–3 thickened; GeIII with setae av1, pv1 with bulbous base, setae al1–2, pl2 thickened; GeIV with seta pv1 with bulbous base, seta pl2 thickened. Tibiae – TiI without modified setae; TiII–III with setae av1, pv1 with bulbous base; TiIV without modified setae. Tarsi – TaI same as female; TaII with setae al2–3, av2, mv with bulbous base; TaIII with setae al2–3, av2, mv, pl2 with bulbous base; TaIV with setae al2, av2, mv with bulbous base. Etymology. It is with great pleasure that we name this species for Mr Siegfried Huber, who collected most of the mites. Remarks. The new species is very close to S. bipartitus by sharing a similar number of setae on the dorsal shield (60–64), similar body size and general shape of ventral shields, the same host genus and females with a smooth medial margin on the gnathotectum. The most obvious difference is found in the ventral shielding of males: S. bipartitus has a holoventral shield, but S. huberi sp. nov. has separate genitosternal and anal shields. Examination of the holotype and only male paratype specimen revealed that males also have different posterodorsal setae on femur II, which in S. bipartitus are long and thickened (length 62–64; Fig. 26) but smaller and finer in S. huberi sp. nov. (pd1 20–25, pd2 28–45; Fig. 23). The new species also bears ten setae on genu III in both sexes (seta pl2 present; Figs 13, 24), but in S. bipartitus genu III of the female bears nine setae (pl2 absent; Fig. 2015


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28) while the male retains ten setae (Fig. 27). Male S. bipartitus also appear to have massive legs in comparison to the female (Figs 26–28) and S. huberi sp. nov. This difference is likely real, but is exaggerated by the flattening of its legs on the slide. The anterior width of the femur and genu in male S. bipartitus for leg II is 120 and 110, respectively, and for leg III 115 and 110, respectively. In comparison, the width in male S. huberi sp. nov. for the femur and genu of leg II is 95–110 and 90– 110, respectively, and for leg III 95–105 and 85–100, respectively.

FIGURES 21–25. Scissuralaelaps huberi sp. nov., male. 21, Leg I, ventral view, coxa–tibia; 22, Leg I, tarsus, dorsal view; 23, Leg II, anterolateral view; 24, Leg III, anterolateral view; 25, Leg III, anterolateral view.

The males of S. bipartitus and S. huberi sp. nov. present significant sexual dimorphism in having a strong thickening of the margins of the peritrematal and metapodal shields, and legs with numerous modified setae. The former feature is not mentioned for any other species, but could have been 716


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overlooked in descriptions, while the male’s modified setae are described only in S. philippinensis. Male S. huberi sp. nov. also show further striking dimorphisms in that the idiosoma projects over most of the gnathosoma, while the latter is mostly exposed in females, and the male’s opisthosoma bends downwards almost perpendicularly, while no such bend occurs in the female. These features, especially the latter, are largely lost in slide-mounted males yet are easily appreciated under a stereomicroscope and scanning electron microscope (Figs 29–30, Figs 32–33).

FIGURES 26–28. Scissuralaelaps bipartitus Ishikawa, 1988. Male, paratype, 26. Leg II, femur–tibia, posterodorsal view; 27, Leg III, femur–tibia, dorsal view. Female, holotype, 28, Leg III, femur–tibia, dorsal view.

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FIGURES 29–31. Scissuralaelaps huberi sp. nov. 29, S. huberi on millipede Acladocricus sp.; 30, S. huberi, showing (a) female and (b) males, with bent opisthosoma; 31, Host millipede Acladocricus sp. (Spirobolida: Rhinocricidae). Photographs by S. Huber.

FIGURES 32–33. Scissuralaelaps huberi sp. nov., scanning electron micrographs. 32. Female, ventral view; 33, Male, ventral view, showing prodorsal hood over gnathosoma (a) and highly curved caudal bend (b).

Discussion The family Iphiopsididae sensu Casanueva (1993), and as also presented in Lindquist et al. (2009), is defined by the loss of pl2 on genu IV and av2 on tibia I, having weakly-developed internal malae, a smooth gnathotectal margin, and by its association with Myriapoda, Araneae and Crustacea. The group has also been defined by having a reduced peritreme (Evans 1955; Lindquist et al. 2009). Scissuralaelaps only has one of these characters (loss of av2 on tibia I), only some species have smooth gnathotectal margins, where studied they have pl2 on genu IV, their peritremes are well developed and two of its members associate with cockroaches. Consequently, if Scissuralaelaps is 718


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considered part of this family, the family-rank is based on the loss of av2 on tibia I alone, which seems a flimsy basis for a family. Therefore, we have followed the treatment of the group as a subfamily of the Laelapidae (e.g., Keegan 1950; Evans 1955) and note that the subfamily is in need of review because Scissuralaelaps is not alone in lacking the characters that define this group: Ljunghia Oudemans, which associates with mygalomorph spiders, has well-developed internal malae and peritremes, and a denticulate gnathotectal margin (Moraza et al. 2009). The family name appears variously as Iphiopsididae or Iphiopsidae, as do the subfamily names Iphiopsidinae or Iphiopsinae. The first use of the family name was Iphiopsidae Kramer, 1886, but this name is based on the incorrect stem as derived from the type genus Iphiopsis Berlese, 1882. The correct Greek stem is Iphiopsid- rather than Iphiops-, even though the International Code of Zoological Nomenclature permits the ending –id to be elided from the stem prior to adding family group suffixes (ICZN article 29.3.1.1). The names Iphiopsidae and Iphiopsinae were used exclusively for over a century since Kramer (1886) (e.g., Keegan 1950; Evans 1955; Rosario 1981) and remain in use (e.g, Moraza et al. 2009; Masan et al. 2012; Seeman & Nahrung 2012) and thus an argument could be made to retain this family name due to prevailing usage (ICZN article 29.5). However, since Casanueva (1993) used the name Iphiopsididae, this correct form has been used in publications such as Lindquist et al. (2009), Farfan and Klompen (2012) and Moraza and Kazemi (2012). As there is no longer consistent prevailing use of the original imperfect spelling, and the correct Greek stem is Iphiopsid-, then family names based on the stem Iphiopsid- should be used in future, i.e., Iphiopsididae and Iphiopsidinae.

Acknowledgements We are very grateful for the input of Dr Matthew Shaw, who provided comments on a draft of the manuscript and pointed out Kramer’s original use of the name Iphiopsidae. We also thank Dr Hirotsugu Ono for lending type specimens of S. bipartitus, and Dr Bruce Halliday (Australian National Insect Collection, Canberra) and an anonymous referee who provided useful corrections and improvements to this manuscript.

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