Davisella trevisani, Rodrigues, Daniela F. S., Navia, Denise, Oliveira, Anibal R., Ferragut, Francisco & Flechtmann, Carlos H. W., 2017

Rodrigues, Daniela F. S., Navia, Denise, Oliveira, Anibal R., Ferragut, Francisco & Flechtmann, Carlos H. W., 2017, Two new eriophyoid mite species (Trombidiformes: Eriophyoidea) from the cocoa tree, and a note on Shevtchenkella biseta (Nalepa), Zootaxa 4237 (1), pp. 112-130 : 114-123

Davisella trevisani n. sp. Rodrigues, Navia & Oliveira

( Figs. 1–2 )

Diagnosis. A Davisella species presenting three wax-like bearing ridges on dorsal opisthosoma, median ridge extending along whole opisthosoma. Enlarged subcylindrical tubercle bearing pedipalp genual seta d. Prodorsal shield without tubercles; ocellar-like structures on lateral area; broad-based and rounded frontal lobe; median and admedian lines not joined by transversal lines. Each empodium branch 5-rayed. Coverflap anteriorly with dashes and posteriorly with 16-24 radiating longitudinal lines. Dorsal annuli lacking microtubercles.

FEMALE (n=10). Body fusiform, brown-yellowish, 134 (126–158), 76 (66–76) wide. Gnathosoma abruptly bent down in lateral view; pedipalp coxal seta (ep) 2 (2); dorsal pedipalp genual seta (d) 2 (2), thickened, curved, slightly hook-like, inserted in apex of enlarged cylindrical tubercle, ca. three times longer as wide, which extends dorsally over pedipalp tibia allowing seta d reach tarsal pedipalp base; subapical pedipalp tarsal seta (v) 1 (1); cheliceral stylets 48 (40–48); base of gnathosoma (anterior to prodorsal shield) finely microtuberculated. Prodorsum downcurved. Prodorsal shield 32 (30–34) including frontal lobe, 58 (53–58) wide, subelliptical; short, broad-based frontal lobe 4 (4–5), 18 (17–19) wide, apically curved. Shield design of faint sinuous longitudinal lines (median, admedian, submedian and sublateral); median and admedian lines extending along 3/4 anterior shield, other lines shorter; ocellar-like structures in lateral area; tubercles and scapular setae (sc) absent. Posterior margin medially pointed, resembling a subtriangular projection. Legs with all segments present; paraxial tibial seta (l’) and basiventral femoral seta (bv) lacking on both legs; genual seta (l”) absent on leg II. Leg I 26 (25– 29); femur 13 (9–13); genu 3 (3–5), antaxial genual seta (l") 38 (35–39); tibia 3 (3–5); tarsus 9 (8–9), antaxial fastigial tarsal seta (ft”) 27 (25–30), paraxial fastigial tarsal seta (ft’) 22 (21–26), unguinal seta (u') angulated 5 (4– 5), tarsal solenidion (ω) blunt, apically expanded, knobbed 5 (5); empodium 6 (6), divided, symmetrical, each branch 5-rayed and each ray bifurcated at base (except apical ray). Leg II 20 (20–23); femur 8 (8–10); genu 3 (3– 4); tibia 4 (3–4); tarsus 6 (5–8), ft" 20 (16–20), ft' 5 (4–5), ω 5 (5); u' 5: empodium 6 (6), 5-rayed as in leg I. Coxisternal region finely granulated, mainly on coxa I. Anterolateral seta on coxisternum I (1b) 11 (7–11), 8 (6– 10) apart; proximal seta on coxisternum I (1a) 13 (10–13), 8 (6–9), apart; proximal seta on coxisternum II (2a) 20 (16–20), 25 (21–26) apart; coxigenital region with 4 (4) annuli, finely microtuberculated. External genitalia 20 (16–23), 32 (28–33) wide, coverflap semicircular, anteriorly with short irregular dashes or with granules and short dashes, posteriorly with 23 (16–24) radiating longitudinal lines; proximal seta on coxisternum III (3a) 6 (5–8). Internal genitalia, not seen. Opisthosoma with three well defined dorsal ridges, wax-bearing; median ridge extending along whole opisthosoma (except for last 4–5 annuli) and lateral ridges extending along 3/4 anterior opisthosoma; 56 (54–64) dorsal annuli, smooth, 53 (47–56) ventral annuli with microtubercles on posterior margin, which are more elongated in between ventral setae or on annuli posterior to ventral seta f. Lateral seta c2 absent; ventral seta d 47 (42–48), on annulus 11 (9–12), 30 (25–34) apart; ventral seta e 29 (21–29), on annulus 26 (18–28), 19 (18–21) apart; ventral seta f 20 (16–20), on annulus 44 (37–46), 23 (14–23) apart; caudal seta h2 34 (28–34); accessory seta h1 minute.

MALE (n=5). Most males smaller than females and similar in shape to them. All measurements of males are given in Table 1. External genitalia 9–13, 21–24 wide; 3a 6–8, granules not seen, eugenital setae seen on anterior median area.

TABLE]. MeaSuremeŊŧS of ŧhe ŊeW erỉophyoỉđ mỉŧeS from Theobroma cacao (L leŊgŧh. Wỉđŧh)

Davisella trevisani n. sp. Tetra theobromae n. sp.

Characŧer Female (Ŋ=10) Male (Ŋ=5) Immaŧure (Ŋ=3) Female (Ŋ=14) Male (Ŋ=1) MeaŊ ± ’Đ MỉŊ Max MeaŊ ± ’Đ MỉŊ Max MeaŊ ± ’Đ MỉŊ Max MeaŊ ± ’Đ MỉŊ Max MeaSure

ỉoSoma L. 138 ±9.2 126 158 123±5.2 120 132 102±4.8 98 107 145±8.3 130 159 115

ỉoSoma. 72 ±3.7 66 76 62±1.4 60 64 52±5.1 49 58 51±2.8 46 56 44

ProđorSal Shỉelđ L. 32 ±1.4 30 34 31±3.3 26 35 30±1.8 29 32 24±0.7 23 25 22 Femur I L. 11±1.2 9 13 10 ±1.2 9 11 10 ±0.7 9 10 8 ±0.8 7 9 7

VeŊŧral ƀaSỉfemoral I (bv) L. 8±0.8 7 8 8

GeŊu I L. 4±0.9 3 5 4±0.7 4 5 4±0.0 4 4 4±0.6 3 5 3 AŊŧaxỉal faSŧỉgỉal Seŧa I (ft' ') L. 25 ±1.9 21 26 25±2.4 21 26 19±0.0 19 19 16±0.9 15 18 15 ……continued on the next page TABLE]. (coŊŧỉŊueđ)

Davisella trevisani n. sp. Tetra theobromae n. sp.

Characŧer Female (Ŋ=10) Male (Ŋ=5) Immaŧure (Ŋ=3) Female (Ŋ=14) Male (Ŋ=1) TABLE]. (coŊŧỉŊueđ)

Davisella trevisani n. sp. Tetra theobromae n. sp.

Characŧer Female (Ŋ=10) Male (Ŋ=5) Immaŧure (Ŋ=3) Female (Ŋ=14) Male (Ŋ=1) MeaŊ ± ’Đ MỉŊ Max MeaŊ ± ’Đ MỉŊ Max MeaŊ ± ’Đ MỉŊ Max MeaŊ ± ’Đ MỉŊ Max MeaSure

ỉSŧerŊal aŊŊul ỉ 4 ±0.0 4 4 2±0.0 2 2 6±0.6 5 7 5 GeŊ ỉŧalỉa L. 19±2.5 16 23 10 ±2.1 9 13 11 ±1.1 9 13 10 GeŊỉŧalỉa. 30±2.1 28 33 22±1.1 21 24 19±0.5 18 20 13 LoŊgỉŧuđỉŊal rỉƀS 19±3.0 16 24 14±1.2 12 16

GeŊỉŧal Seŧa (3a) L. 7±0.9 5 8 7±0.7 6 8 3±0.7 3 4 10 ±0.9 8 11 7 Laŧeral Seŧa (c2) L. 19±1.3 18 22 19 Laŧeral Seŧa (c2) aŊŊuluS 4±0.8 2 5 3 VeŊŧral Seŧa I (d) L. 47±2.8 42 48 45±5.3 42 49 13±0.0 13 13 34±1.3 32 36 31 ỉSŧaŊce ƀeŧWeeŊ (d) 30±4.2 25 34 24±1.6 23 26 19±2.2 18 21 32±1.9 27 33 28 VeŊŧral Seŧa I (d) aŊŊuluS 11±1.2 9 12 12 ±1.6 10 14 8 ±0.6 8 9 15 ±0.8 13 16 13 VeŊŧral Seŧa II (e) L. 25±5.6 21 29 30±2.7 28 32 8 ±0.7 8 9 7±0.9 5 8 5 ỉSŧaŊce ƀeŧWeeŊ (e) 19±1.3 18 21 15±1.1 14 16 12 ±0.7 11 13 17 ±1.7 14 19 16 VeŊŧral Seŧa II (e) aŊŊuluS 25±3.2 18 28 28±2.2 25 30 18±1.2 17 19 28±1.7 26 32 23 VeŊŧral Seŧa III (f) L. 17±1.5 16 20 14±0.0 14 14 9 ±1.7 8 11 13 ±0.8 12 14 11 ỉSŧaŊce ƀeŧWeeŊ (f) 20±2.4 14 23 16±3.5 10 19 12 ±1.5 11 14 15 ±1.1 13 16 14 VeŊŧral Seŧa III (f) aŊŊuluS 42±2.7 37 46 43±3.9 39 47 34±1.2 33 35 44±2.5 41 50 38 Toŧal đorSal aŊŊul ỉ 60 ±3.4 54 64 57±2.4 53 58 51±1.7 49 52 22±0.9 21 24 21 Toŧal veŊŧral aŊŊul ỉ 51 ±3.0 47 56 53±2.5 49 55 41±0.6 41 42 48±2.5 45 54 43 Cauđal Seŧa (h2) L. 29±2.8 28 34 31±2.1 29 34 11 ±0.7 10 11 37 ±1.9 35 40 42 AcceSSory Seŧa (h1) L. 2±0.0 2 2 2

IMMATURE STAGE (n=3). Body slightly fusiform, 98–107, 49–58 wide. Gnathosoma setae not seen, probably due to reduced size, cheliceral stylets 30–35; d minute inserted on an elongated cylindrical tubercle (as in adults). Prodorsal shield as in adults, except for ocellar-like structures which were not seen, 29–32, 40 wide. Coxisternal region. 1b 4, 6 apart, 1a 5, 6 apart, 2 a 9, 9–14 apart. Legs as in adults. Leg I 21–23; femur 9–10; genu 4, l" 23–28; tibia 3–4; tarsus 5–6, ft" 19, ft' 18–19, u' 4, ω 4, as in adults; empodium 5, 5-rayed. Leg II 14–15; femur 5; genu 3; tibia 1–3; tarsus 4–5, ft" 11–13, ft' 4, u' 4, ω 4; empodium 4–5, 5-rayed. Genitalia absent, 3a 3–4, on ventral annulus 9. Opisthosoma evenly curved (without dorsal ridges); 49–52 dorsal annuli, smooth, except for three longitudinal rows with small slightly elongated microtubercles (probably corresponding to openings [pores] of wax-like material secreting glands observed in adults); 41–42 ventral annuli posterior to seta 3a, with small, slightly elongated microtubercles. Annuli equal dorso-ventrally; c2 absent; d 13, on annulus 8–9, 18–21 apart; e 8 – 9, on annulus 17–19, 11–13 apart; f 8–11, on annulus 33–35, 11–14 apart. h 2 10–11; h1 not seen.

Type material. Twenty-nine specimens collected from Theobroma cacao L. ( Malvaceae ) leaves, at " Estação Experimental de Ouro Preto do Oeste (ESTEX-OP), Comissão Executiva do Plano da Lavoura Cacaueira (CEPLAC)", municipality of Ouro Preto do Oeste, Rondônia, Brazil (10°44'30"S, 62°13'30"W), on 6 September 2012, coll. Francisco Ferragut and Denise Navia. Female holotype (red labelled slide, indicated by a red ring and in the specimen map at the right side label) and nine paratypes (7 females, 1 male and 1 immature) on four slides deposited in the Acarological Collection at Universidade Estadual de Santa Cruz ( UESC), Ilhéus, BA, Brazil . Ten paratypes (6 females and 4 males) in two slides deposited in the Acarological Collection at Embrapa Recursos Genéticos e Biotecnologia, Brasilia , DF, Brazil . Nine paratypes (5 females, 1 male and 3 immatures) in two slides deposited in the Acarological Collection at “ Departamento de Entomologia e Acarologia, Universidade de São Paulo, Escola Superior de Agricultura ‘ Luiz de Queiroz’ ( ESALQ)", Piracicaba , SP, Brazil .

Relation to host. A vagrant species found on lower surfaces of young leaves, no symptoms observed.

Etymology. The species is named in honor to Dr. Olzeno Trevisan, entomologist, CEPLAC, Rondônia, Brazil, for his remarkable enthusiasm and contribution to the knowledge of arthropods associated with cocoa trees and other plants in the Amazon forest. The specific name " trevisani " is derived from the surname of the honored researcher in the genitive possessive case.

Remarks. Davisella trevisani n. sp. is the 8th species to be assigned to this genus ( Table 2). It can be distinguished from all other species by the dorsal opisthosomal wax-bearing ridges. In other species these dorsal ridges are slight (not very well defined) or extend just along the anterior opisthosoma as in D. breitlowi ( Davis, 1964) , D. paucisetosa ( Flechtmann, 1999) , D. palmea ( Flechtmann, 1998) , D. notosa ( Flechtmann, 1995) , and D. spondias Reis & Navia, 2010 . Though D. haramotonis ( Keifer, 1974) and D. globosa ( Keifer, 1969) also present a noticeable median ridge extending along the whole opisthosoma, the lateral ridges are not present and/or ridges are not described as wax-like bearing.

The prodorsal shield of the new species is very similar to that of D. spondias in the ornamentation pattern of free longitudinal lines and circumscribed with a semi-elliptical line. However in the new species the frontal lobe is rounded while in D. spondias it is trapezoidal. In addition D. trevisani n. sp. presents ocellar-like structures in its lateral area (not observed in D. spondias ). The new species is similar to D. palmea , D. notosa and D. spondias in the absence of prodorsal shield tubercles (in D. breitlowi , D. globosa , D. haramotonis and D. paucisetosa tubercles are present ahead of rear shield margin). It is also similar to D. breitlowi in the 5-rayed empodium (all other species present 6-7 rays in each empodial branch, see Table 2). The new species can be distinguished from D. breitlowi , D. globosa , D. palmea and D. paucisetosa by the absence of microtubercles on the dorsal annuli (dorsal microtubercles present in these species, see Table 2). The coverflap of the new species is similar to that of D. haramotonis , D. spondias and D. paucicetosa , by presenting microtubercles or short irregular dashes anteriorly and less than 30 radiating longitudinal lines posteriorly.

Ocellar-like structures are present in the lateral area of the prodorsal shield of D. trevisani n. sp., Flechtmann (1998) represented two circles in the prodorsal shield median area near rear shield margin of D. palmea . These circles could also constitute ocellar-like structures. If this is the case, then D. trevisani n. sp. and D. palmae would be similar in the presence of ocellar-like structures in the prodorsal shield, although these structures are placed in different positions in the two species (lateral in D. trevisani n. sp. and median in D. palmae ).

Enlarged subcylindrical tubercles bearing the pedipalp genual seta, like those observed in D. trevisani n. sp., have never been reported in other eriophyoid mites. Chetverikov & Craemer (2015) conducted a detailed study on the gnathosoma morphology of about 80 mite species in the three recognized families of Eriophyoidea, Species D. breitlowi D. globosa D. haramotonis D. notosa D. palmea D. paucisetosa D. spondias D. trevisani n.

( Davis, 1964) ( Keifer, 1969) ( Keifer, 1974) (Flechtmann, (Flechtmann, (Flechtmann, Reis & Navia, sp. 1995) 1998) 1999) 2010

plant, Magnolia Anacardium Psidium guajava Miconia Syagrus Trema micrantha Spondias Theobroma

type locality grandiflora L., occidentale L., L., Myrtaceae candolleana romanzoffiana (L.) Blum. mombin L., cacao L., Magnoliaceae Anacardiaceae Oahy , Hawaii, USA Triana, (Cham.) Glassm. Ulmaceae Anacardiaceae Malvaceae Georgia, USA São Paulo, Brazil Melastomataceae Arecaceae São Paulo, Brazil Pernambuco, Rondônia, Brazil Rio de Janeiro, Rio de Janeiro, Brazil

Brazil Brazil

number of rays 5 7 6 7 6 – 7 7 6 5 empodium

branch

prodorsal shield minute, ahead minute, ahead minute, ahead rear absent/absent absent/circles near minute or absent absent/absent absent/presents tubercles/ rear shield rear shield shield margin/ shield margin (missing in 5% of in the lateral area ocellar-like margin/ absent margin/ absent absent (probably ocellar- specimens)/

structures like) absent

prodorsal shield longitudinal lines longitudinal lines faint longitudinal longitudinal lines smooth longitudinal lines longitudinal lines faint sinuous

ornamentation (median and (weak median lines (median line (admedian lines (median line long (median and median and admedian lines) line; admedian on rear posterior meeting curved flanked by two admedian, not admedian lines complete; half; admedian cross lines at its admedian narrow, joined) longitudinal lines submedian lines) lines complete; 1/4 posterior; elongated bosses)

submedian lines) submedian lines)

……continued on the next page Species D. breitlowi D. globosa D. haramotonis D. notosa D. palmea D. paucisetosa D. spondias D. trevisani n.sp.

( Davis, 1964) ( Keifer, 1969) ( Keifer, 1974) (Flechtmann, Flechtmann, (Flechtmann, Reis & Navia,

1995) 1998) 1999) 2010

dorsal slight median median and lateral median ridge slight median and slight median and slight median slight median three well defined

opisthosoma ridge ridges lateral ridges lateral ridges ridges ridge wax-like bearing

ridges ridges

coverflap distally 32 short basally granulated, basally with short basally granulated, 44 – 52 basally with basally with basally with short

ornamentation longitudinal distally 22 – 25 short dashes mainly set distally 20 – 22 longitudinal lines irregular short granules or short irregular dashes lines irregular in longitudinal radiating lines occupying whole dashes, distally dashes; distally and/or granules, longitudinal lines lines, distally 16-18 coverflap (median 22 – 26 radiating 35 – 38 radiating apically with 16 – short radiating lines 12 – 16 lines longer lines longitudinal lines 24 radiating interrupted longitudinal lines halfway)

coxigenital smooth lines of fine dashes curved lines of granulated around granulated smooth granulated finely

ornamentation short dashes tubercles microtuberculate

. dorsal annuli 62 78 75 78 – 95 45 – 53 76 – 83 69 – 77 54 – 64 opisthosoma small, beadlike, dorsally absent; dorsally faint or dorsally absent; dorsally small dorsally spaced; dorsally absent dorsally absent; microtubercles on rear annulus fine on ventral side, absent; laterally as ventrally very beadlike ventrally fine (except for last ventrally with margin, spacer weak or absent fine points; small beadlike microtubercles; twelve); ventrally microtubercles, dorsally laterally ventrally present ventrally fine pointed whose are more elongated in between ventral

Morphological traits were extracted from text and/or drawings of species original descriptions.

including three genera of Diptilomiopidae : two in the Diptilomiopinae , Diptacus Keifer, 1951 and Diptilomiopus Nalepa, 1916 ; one in the Rhyncaphytoptinae , Rhyncaphytoptus Keifer, 1939 . These authors did not observe similar structures on the pedipalp segments of any of the studied species. Minute gnathosomal structures which seem to be involved in keeping the mouthparts in position, especially the stylets sheath, named the "gnathosomal interlocking apparatus", were observed as being restricted to the basal parts of chelicerae, medioproximal palpcoxae and anterodorsal margin of the prodorsal shield. The enlarged pedipalp genual tubercle observed in D. trevisani n. sp. possibly acts as stopper to prevent "losing" stylets from the "envelop" between palps, a similar function that of the "cheliceral retainer" observed in Eriophyidae and Diptilomiopidae species by Chetverikov & Craemer (2015). These authors described four types of gnathosomal interlocking apparatus. Further morphological studies should be conducted to confirm function of this interesting structure as an interlocking apparatus, which would be classified as a new type.

Figures

FIGURE 1. Davisella trevisani n. sp. D. Dorsal habitus, female; V. Ventral habitus, female; GF. External genitalia, female; GM. External genitalia, male; PI. Pedipalp, immature (enlarged). PD. Detail of dorsal pedipalp, female (enlarged). PL. Detail of lateral pedipalp, female (enlarged).

FIGURE 2. Davisella trevisani n. sp. L. Dorsolateral habitus, female; IL. Ventrolateral habitus, immature; ED. Dorsal view of empodium, female (enlarged); EL. Lateral view of empodium, female (enlarged); L 1. Leg I, antaxial view, female; L 2. Leg II, antaxial view, female.