European Journal of Taxonomy 202: 1-25

http://dx.doi.org/10.5852/ejt.2016.202

This work is licensed under a Creative Commons Attribution 3.0 License.

ISSN 2118-9773

www. europeanj ournaloftaxonomy. eu

2016 • Paredes-Leon R. et al.

Research article

um:lsid:zoobank.org:pub:55CBC031-F369-48A2-BE0E-2249AB7A43Dl

Camerobiid mites (Acariformes: Raphignathina: Camerobiidae)

inhabiting epiphytic bromeliads and soil litter of tropical dry forest

with analysis of setal homology in the genus Neophyllobius

Ricardo PAREDES-LEON '•*, Angelica Maria CORONA-LOPEZ 2 ,

Alejandro FLORES-PALACIOS 3 & Victor Hugo TOLEDO-HERNANDEZ 4

\, 2,3 ,4 c en t ro d e Investigacion en Biodiversidad y Conservacion (CIByC), Universidad Autonoma del

Estado de Morelos, Avenida Universidad 1001, Col. Chamilpa, C.P. 62209, Cuernavaca, Morelos, Mexico.

* Corresponding author: [email protected]

2 Email: [email protected]

’Email: [email protected]

4 Email: [email protected]

1 urn :lsid: zoobank.org: author: 3 A3 A9078-178C-41AD-8520-B2E72BDF C21C

2 urn:lsid:zoobank.org:author:D9D501D6-5883-4C9A-877D-5567149BC542

3 urn:lsid:zoobank.org:author:DF49E2C9-D57A-4AF9-92AB-AD3C828A97Dl

4 um:lsid: zoobank.org: author :EEB4 IE AF-BA41-4EEF -BA 11-3F CFAF37EE93

Abstract. A survey of the camerobiid mites living on epiphytic bromeliads and the forest floor of a

Mexican tropical dry forest was carried out. We found three new species of the genus Neophyllobius,

which are described in this paper; the first two, namely N. cibyci sp. nov. and N. tepoztlanensis

sp. nov., were both found inhabiting bromeliads ( Tillandsia spp.) and living on two tree species

(Quercus obtusata and Sapium macrocarpum ); the third, N. tescalicola sp. nov., was found in soil

and litter under Q. obtusata. These three new species can be differentiated from other species in the

genus by a combination of morphological characters in adult females, mainly those setae on femora

and genua I. The idiosoma and leg setal ontogeny of a camerobiid mite is presented for the first

time in this paper, illustrating chaetotaxic notations and their relative positions in N. cibyci sp. nov.

larva, protonymph and adults (female and male), and establishing setal homologies among instars.

Setal homology with other species in the cohort Raphignathina is briefly discussed. Additionally,

a compilation and an identification key to all known species of camerobiid mites in Mexico is

provided.

Keywords. Acari, chaetotaxy, Mexico, predator. Prostigmata, tree canopy, Tillandsia spp.

Paredes-Leon R., Corona-Lopez A.M., Flores-Palacios A. & Toledo-Hernandez V.H. 2016. Camerobiid mites

(Acarifonnes: Raphignathina: Camerobiidae) inhabiting epiphytic bromeliads and soil litter of tropical dry forest

with analysis of setal homology in the genus Neophyllobius. European Journal of Taxonomy 202: 1-25. http://

dx.doi.org/10.5852/ejt.2016.2Q2

European Journal of Taxonomy 202 : 1-25 ( 2016 )

Introduction

The family Camerobiidae Southcott, 1957 (Acariformes: Prostigmata: Raphignathina: Raphignathoidea)

comprises about 161 species in seven genera of predatory mites (Fan & Walter 2011; Zhang et al. 2011;

Khanjani etal. 2012a, 2012b, 2013, 2014; Akyol 2013; Khanjani & Hoseini2013; Zeity & Gowda 2013;

Ulu 9 ay & K 09 2014). Camerobiids are usually found in different microhabitats like humus, tree bark,

vegetation or inside epiphytes in temperate and tropical ecosystems. Some camerobiid species feed on

plant pests such as scale insects (Coccoidea), plant-parasitic mites (Eriophyidae and Tenuipalpidae)

and on other small mites that scavenge or feed on fungi on plant surfaces (Tarsonemidae and Tydeidae)

(McGregor 1950; De Leon 1967; Bolland 1983; Gerson et al. 2003; Walter et al. 2009).

Neophyllobius Berlese, 1886 species have world-wide distribution, and the genus is the richest in

Camerobiidae with more than 100 species. The genus was revised by Bolland (1991) and later works

mainly include species descriptions (e.g., K 09 & Ayyildiz 1996; Bolland 2001; Khanjani & Ueckermann

2006; Khanjani et al. 2010; Ulu 9 ay & K 09 2014). Like many other groups of mites, there is insufficient

knowledge regarding their ecology and ontogeny which, in turn, restricts our knowledge of habitat

specificity and the identification of homologies among species. Also, many descriptions of camerobiid

mites are based on a single specimen because they are usually found singly or in low numbers (Bolland

1986), and rarely is setal nomenclature detailed.

Members of Raphignathoidea generally include four active postembryonic instars: larva, protonymph,

deutonymph and adult (Walter et al. 2009). However, in Neophyllobius it is questioned whether the life

cycle includes one nymphal instar instead of two. Protonymphs are frequently found, but deutonymphs

have been mentioned briefly only in a few species, for example N. saxatilis Halber, 1938, N. ambulans

Meyer, 1962, N. gonzali Zaher & Gomaa, 1979, N. aesculi Bolland, 1983, N. ceratoniae Bolland, 1991,

N. dichantii Bolland, 1991, N. muscantribii Bolland, 1991 and N. piniphilus Bolland, 2001 (Bolland

1983, 1991, 2001). Only the deutonymphs of the last two species has been partially illustrated (Bolland

1983, 1991). Because of the scarcity of specimens the distinction between deutonymph and adult female

is not always clear. Besides, two types of protonymphs (male and female) has been recognized at least

in N. piniphilus (Bolland 2001).

The family Camerobiidae in Mexico comprises 13 species of Neophyllobius (Bolland 1991) (Table 1).

This number is clearly low, suggesting that the group is under-represented, and shows that camerobiid

mites have been reported throughout tropical and northern areas and in a variety of different habitats

(inside epiphytes, tree bark or in the forest floor).

Analysis of leg chaetotaxy for different instars in rich groups, like the genus Neophyllobius , could provide

systematic and phylogenetic clues for a better understanding of the groups at different hierarchical

levels (Swift 2001), and may be helpful in distinguishing between lineages (Norton 1977). It is important

to homologize the setal organs in the different species and among different instars because to ignore the

homology and only take into account the numerical formulas in descriptions ofnewtaxa is not phylogenetically

informative. In the Camerobiidae, Grandjean’s idiosomal chaetotaxy system was firstly applied by Kethley

(1990), whereas the leg chaetotaxy system was implemented in adult females belonging to Acamerobia

inflatus Fan & Walter, 2011 (Fan & Walter 2011). However, leg chaetotaxy is frequently omitted in most

recent species descriptions, with a few noteworthy exceptions (e.g., Khanjani et al. 2013, 2014).

In this paper, we provide descriptions of three new species of Neophyllobius : two that have been found

exclusively on epiphytic bromeliads and one on soil litter in Mexico. An identification key to females

of all the species of Neophyllobius in Mexico is provided. The ontogeny of idiosoma and leg setation,

setal notations (chaetotaxy) and their relative positions on the podomeres are presented and illustrated

for one of these new species.

PAREDES-LEON R. et al ., New species of camerobiid mites and setal homology in Neophyllobius

Table 1 . Species of Neophyllobius Berlese, 1886 recorded in Mexico.

SPECIES

LOCALITY

HABITAT

REFERENCE

REMARKS

N. mexicamis McGregor,

1950

Mexico, intercepted at

Brownsville, TX

On avocado budwood

McGregor (1950) Unknown locality

N. quadrisetosus De Leon, SONORA: Guaymas,

1958 intercepted at Nogales

Soil from cactus

De Leon (1958)

N. trisetosus De Leon,

OAXACA: Oaxaca

Ouercus sp.

De Leon (1958)

1958

OAXACA: Siete

Cabrillas

Ouercus sp.

OAXACA:

Tamazulapan

MICHOACAN: near

Quiroga

NAYARIT: Santa

Maria del Oro

NAYARIT: Ixtlan del

Rio

Unknown tree

N. farrieri De Leon, 1958

OAXACA: Puenta

[.s'/c] de Nejapa

Spanish moss

De Leon (1958)

The common name “Spanish

moss” is commonly used to

refer to Tillandsia usneoides

(L.) L.

N. inequalis De Leon,

1958

VERACRUZ:

Veracruz

Erythrina sp., Terminalia

catappa L.

De Leon (1958)

N. equal is De Leon, 1958

NAYARIT: Navarrete

SAN LUIS POTOSI:

near Valles

Guazuma sp.

De Leon (1958)

According to De Leon

(1958) and Bolland (1991),

it is open to doubt whether

the specimens recorded in

San Luis Potosi belong to

this species

N. consobrinus De Leon,

1958

CHIAPAS: Tuxtla

Gutierrez

Jaquinia pungens A. Gray

De Leon (1958)

N. lobatus De Leon, 1958

NAYARIT: San Bias

From “guasamolleta”

De Leon (1958)

On Guasa molleta sensu

Bolland (1991)

N. marginatus De Leon,

1958

NAYARIT: Ixtlan del

Rio

Asteraceae

De Leon (1958)

NAYARIT: nine miles

west of Tepic

Ouercus sp.

N. longulus De Leon,

1958

OAXACA: Matias

Romero

Calophyllum sp.

De Leon (1958)

N. curtipilus De Leon,

1958

CHIAPAS: Tuxtla

Gutierrez

Cordia dentata Poir.,

Guazuma tomentosa

Lam.,

Rhus schideana Schltdl.,

Lonchocarpus rugosus

De Leon (1958)

De Leon first (1958) named

this species as N. curtipilus

(p. 175, 178, 180), but later

the name was misspelled as

N. curtipilis (p. 181)

Benth.,

Ouercus sp., and

avocado

Also, Bolland (1991)

misspelled it as N. curtipilis

N. spatulus De Leon,

1958

NAYARIT: Tepic

Two undetermined

species of Asteraceae

De Leon (1958)

The type specimens are lost

(Bolland 1991)

OAXACA: km post

666, Route 190 north

of Tehuantepec

A Malpighiaceae shrub

European Journal of Taxonomy 202 : 1-25 ( 2016 )

SPECIES

LOCALITY

HABITAT

REFERENCE

REMARKS

N. deleoni Bolland, 1991

JALISCO: route 35,

18 miles southeast of

Guadalajara

Unknown

Bolland (1991)

Originally described as

NeophyUobius horridus

by De Leon (1958) but as

the name was preoccupied,

Bolland (1991) re-named it

DOUBTFUL AND UNDETERMINED SPECIMENS

NeophyUobius spp.

CAMPECHE:

Calakmul

Soil litter

Vazquez et al.

(2011)

CAMPECHE: Zoh-

Laguna

Soil litter

QUINTANA ROO:

Reserva de la Biosfera

de Sian Ka’an

Soil litter

QUIN TANA ROO:

Bahia de Chetumal

Leaf litter and mangrove

Vazquez-

Gonzalez (2008)

MORELOS: Mpio.

Tepoztlan: “derrame

lavico del volcan

Chichinautzin”

Tillandsia spp.

This study

See comments about

Camerobia sp.

MORELOS: Mpio.

Tepoztlan: “derrame

lavico del volcan

Chichinautzin”

Tillandsia spp.

Palacios-Vargas

(1982)

All the species of

Camerobia Southcott,

1957 are not distributed

throughout the Americas.

We reviewed six adult

specimens from this series

(CNAC009249-009254)

and they are NeophyUobius

Camerobiidae gen. et sp.

MORELOS

VERACRUZ

Tillandsia prodigiosa

(Lemaire) Baker

Hoffmann &

Lopez-Campos

(2000)

Undetermined genus and

species

Material and methods

This research was carried out in the tropical dry forest about 1 km south of San Andres de la Cal,

Tepoztlan, Morelos, Mexico (18.95617° N, 99.11394° W, 1495 m a.s.l.). Mean annual temperature is

about 20.5°C and mean annual precipitation is 1091.8 mm. The tropical dry forest in the study area has

an open canopy with short tree species (maximum 16 m in hight); two forest units can be identified by

plant composition and the parent rock of the soil (Cortes-Anzures 2015; Vergara-Torres et al. 2010).

Sampling was done in the forest unit emerged on a lava flow; here, epiphytes are abundant (25

species) and the dominant tree species are Sapium macrocarpum Miill.Arg. (Euphorbiaceae), Ipomoea

pauciflora M.Martens & Galeotti (Convolvulaceae) and Quercus obtusata Bonpl. (Fagaceae), which

together comprise 46% of tree individuals. The most abundant epiphytes are widespread bromeliads

Tillandsia hubertiana Matuda and T. schiedeana Steud. (Bromeliaceae), which comprise 76% of all

adult individuals (Cortes-Anzures 2015).

Camerobiid mites were collected from two species of epiphytic plants (Bromeliaceae): Tillandsia

hubertiana and T. schiedeana. Both bromeliads are xerophytic, but T. schiedeana has been deemed an

atmospheric epiphyte (gray, narrow leaves that do not form an impounding tank) whereas T. hubertiana

is a tank epiphyte (green, wide leaves that do form an impounding tank) (Ruiz-Cordova et al. 2014).

T. hubertiana is endemic to Mexico (Espejo-Serna et al. 2004). Both bromeliads were collected from

PAREDES-LEON R. et al., New species of camerobiid mites and setal homology in Neophyllobius

trunks or branches of two tree species: Quercus obtusata and Sapium macrocarpum. Quercus obtusata is

an oak species endemic to Mexico (Arizaga et al. 2009). Additionally, soil litter samples were collected

from around the base of the trunk of both Q. obtusata and S. macrocarpum.

Mites were extracted from soil litter samples and bromeliads with Berlese-Tullgren funnels and preserved

in 80% ethanol; previously, the bromeliad leaves were cut into small pieces. Then mites were observed

under a stereoscopic microscope (Olympus SZ) and were cleared in potassium hydroxide (KOH) solution.

Microscope slides were made using Hover's medium as a preserver for observation and illustration using

a drawing tube adapted to a compound microscope (Nikon Optiphot-2). Additional observations were

made with an optic microscope (Nikon Labophot-2). Line drawings were edited using the GNU Image

Manipulation Program (GIMP) (The GIMP team 2014). Holotype and paratype specimens are deposited

in the “Coleccion Nacional de Acaros” (CNAC) kept at the “Instituto de Biologia, Universidad Nacional

Autonoma de Mexico” in Mexico City, and three paratype specimens in the “Coleccion de Insectos de

la Universidad de Morelos” (CIUM) at the “Centro de Investigacion en Biodiversidad y Conservacion,

Universidad Autonoma del Estado de Morelos”, in Cuernavaca, Morelos, Mexico.

The nomenclature for describing palp and leg chaetotaxy follows Grandjean (1944, 1946) and Fan &

Walter (2011), and the terminology of idiosomal chaetotaxy follows Kethley (1990) and Fan & Walter

(2011). In descriptions of male, protonymph and larva, only specific features that were found to be

different from those of females are mentioned. All measurements are given in micrometres (pm).

Results

Subclass Acari Sundevall, 1833

Superorder Acariformes Zakhvatkin, 1952

Order Trombidiformes Reuter, 1909

Suborder Prostigmata Kramer, 1877

Supercohort Eleutherengonides Oudemans, 1909

Cohorte Raphignathina Kethley, 1982

Superfamily Raphignathoidea Kramer, 1877

Family Camerobiidae Southcott, 1957

Genus Neophyllobius Berlese, 1886

Neophyllobius cibyci sp. nov.

um:lsid:zoobank.org:act:66ED492F-lC66-4CD7-8FFl-5340094F49A4

Figs 1M; Table 2

Diagnosis

This species is unique due to a combination of following characters: female with dorsal setae reaching

setae immediately behind, femur II with three setae id, v ’ and v ”), femur III with two setae (d and v ’),

setae d on genua I-IV reaching tibiae, short setae (about % of podomere length) on femur I, setae / ’

and d on femur I nearly located horizontally at same level, dorsal setae cl and dl distinctly longer than

distance between setae cl-dl and dl-el respectively, idiosomal setae dl longest, and setae d and V on

palpal femur heavily spinose.

Etymology

The specific name incorporates the acronym of the “Centro de Investigacion en Biodiversidad y

Conservacion” (CIByC) at the “Universidad Autonoma del Estado de Morelos” in recognition of its

work on biological conservation.

European Journal of Taxonomy 202 : 1-25 ( 2016 )

Type material

Holotype

MEXICO: §, CNAC009229, ex Tillandsia hubertiana at 2.5 meters (m) on Quercus obtusata. Collected

from the type locality on 12 Mar. 2014, O. Cortes and R. Paredes coll. (RPL1219).

Paratypes

MEXICO: 1 §, CNAC009230, ex T. schiedeana at 3 m on Q. obtusata (RPL1220); 1 $, CNAC009231,

ex T. schiedeana at 3.05 m on Q. obtusata (RPL1223); 1 §, CNAC009232, 1 L, CNAC009243, ex

T. hubertiana at 2.1 m on Q. obtusata (RPL1225); 1 CNAC009233, ex T. hubertiana at 2.7 m on

Sapium macrocarpum (RPL1237); 1 CNAC009234, ex T. schiedeana at 2.7 m on S. macrocarpum

(RPL1238); 1 §, CIUM, ex T. schiedeana at 2.7 m on S. macrocarpum (RPL1241); 1 §, CNAC009235,

ex T. schiedeana at 2.9 m on Q. obtusata , 22 Aug. 2014, S. Gomez and R. Paredes coll. (RPL1295); 1 $,

CNAC009236, ex T. schiedeana at 2.6 m on S. macrocarpum , 22 Aug. 2014, S. Gomez and R. Paredes

coll. (RPL1304); 1 §, CNAC009237, on soil litter of S. macrocarpum , 23 Oct. 2014, R. Reyes and R.

Paredes coll. (RPL 1341); 1 S, CNAC009238, ex T. hubertiana at 3.6 m on Q. obtusata (RPL1222);

1 $, CNAC009239, ex T. hubertiana at 2.7 m on S. macrocarpum , 30 Apr. 2014 (RPL1276); 1 $,

CNAC009240, ex T. schiedeana at 2.4 m on S. macrocarpum , 22 Aug. 2014, S. Gomez and R. Paredes

coll. (RPL1301); 1PN, CNAC009241, ex T. hubertiana at 3.4 m on Q. obtusata , 30 Apr. 2014 (RPL1255);

1 L, CNAC009242, same data as holotype (RPL 1219); 1 L, CNAC009244, ex T. hubertiana at 2.6 m

on S. macrocarpum , 30 Apr. 2014 (RPL 1279). All collected from the type locality, with same data as

holotype except where noted.

Type locality

MEXICO: Morelos, Tepoztlan, 1 km S of San Andres de la Cal, 18.94305° N, 99.11786° W, 1490 m

a.s.l.

Description

Female (n =10) (Figs 1, 3A-D, 4A-D)

Holotype female (followed in parentheses by range of holotype and nine paratype females).

Gnathosoma. 87 (81-98) long and 93 (77-93) wide. Subcapitulum with subcapitular setae m 28 (28-31)

and two pairs of adoral setae Orl 11 (9-14) and Or2 9 (8-13); these three pairs are nude and slender,

m longest (Fig. IB); distance m-m 25 (21-25). Chelicerae 38 (30M4) long. Palp, five-segmented with

following setal distribution: trochanter without setae; femur with two serrate setae, d 20 (17-20) and l’

39 (31-39); genu with one long, slender, nude dorsal id) seta 39 (33-39); tibia with three tactile setae (/ ’,

l” and d) and one claw (sword-like seta); tarsus with two eupathidia (acmC and sulQ, two simple setae

(. ba and va) and one small solenidion (co) (Fig. 1A). Setae elcp present.

Idiosoma. Longer than wide, 280 (270-350) long (gnathosoma excluded), 235 (195-320) wide.

Cuticle striated, except on coxae and attenuated between setal pairs cl, dl, el and fl (Fig. 1C).

Dorsum. With 15 pairs of serrated setae set on small tubercles; all setae are long, dl longest and h2

shortest; all setal pairs are longer than distance to setal pairs immediately behind. Two pairs of eyes

are positioned between setae sci and see. Length of setae: vi 58 (47-58), ve 50 (42-56), sci 46 (40-

53), see 46 (42-52), pdx 67 (55-71), cl 80 (66-83), c2 70 (53-70), dl 82 (64-82), d2 52 (43-54),

el 77 (60-77), e2 52 (45-57),// 67 (54-72), f2 41 (36-43), hi 35 (34-39), h2 34 (29-34). Distances

between setae: vi—vi 66 (45-66), ve-ve 96 (70-100), vi-ve 26 (26-32), sci-sci 115 (91-120), sce-sce

145 (125-150), sci-sce 45 (35—45), cl-dl 45 (34-58), dl-dl 16 (10-16), dl-d2 83 (82-94), dl-el

42 (39-46), ve-sci 22 (15-22), pcbc-pcbc 23 (13-24), pdx-cl 21 (22-37), cl-cl 16 (11-17), cl-c2

98 (86-100), c2-c2 185 (155-190), el-el 14 (11-16), el -fl 46 (37-59), fl-h1 50 (39-57), el -e2

PAREDES-LEON R. et al ., New species of camerobiid mites and setal homology in Neophyllobius

Fig. 1 . Neophyllobius cibyci sp. nov., §, holotype. A. Palp. B. Subcapitulum. C. Dorsal idiosoma.

D. Ventral idiosoma. E. Trochanter-tibia of leg I. F. Tarsus I.

European Journal of Taxonomy 202 : 1-25 ( 2016 )

Fig. 2. Neophyllobius cibyci sp. nov. A-B. paratype (CNAC009238). A. Dorsal idiosoma. B. Ventral

idiosoma. C-D. Protonymph, paratype (CNAC009241). C. Dorsal idiosoma. D. Ventral idiosoma.

E-F. Larva, paratype (CNAC009242). E. Dorsal idiosoma. F. Ventral idiosoma.

PAREDES-LEON R. et al.. New species of camerobiid mites and setal homology in Neophyllobius

66 (56-70), d2-e2 42 (39-45),/?-/? 14 (9-14), fl-f2 66 (46-66), f2-f2 115 (105-125), e2-f2 42

(33-71), hl-hl 10 (6-11), hl-h2 30 (21-30), h2-h2 61 (50-68), f2-h2 33 (29-40). Venter. Coxal

setae thinner than dorsal setae. Setal pairs la, 3a, 4a, ag, g,psl, ps2 and ps3 nude and short. Setae

lb, lc, 2c, 3b, 3c, 4b and 4c serrated. Setae lc longest and psl—3 shortest. Setal pairs 3a and 4a

located on individual platelets. Genito-anal valves with one pair of genital setae (g). Coxa I grouped

with coxa II, and coxa III with IV but not completely fused (Fig. ID). Length of setae: la 26 (25-29),

lb 29 (22-30), lc 69 (53-69), 2c 54 (33-56), 3a 34 (33-38), 3b 37 (25-40), 3c 49 (34-50), 4a 18

(18-26), 4b 21 (17-22), 4c 30 (22-31), ag 15 (15-19), g 10 (10-15), psl 12 (10-14), ps2 12 (10-14),

ps3 12 (9-12).

Legs. Slender and long, leg I and IV longest. Lengths (excluding ambulacra): leg I 565 (480-565), leg II

495 (440-500), leg III 475 (465-530), leg IV 550 (510-590). Podomere lengths: femur I 180 (155-180),

femur II 145 (125-145), femur III 145 (130-145), femur IV 165 (135-165), tibia I 185 (165-185),

tibia II 160 (135-160), tibia III 185 (165-190), tibia IV 195 (180-200), tarsus I 62 (60-70), tarsus II

69 (60-70), tarsus III 71 (59-71), tarsus IV 71 (64-76). Leg setae as indicated in Figs 1E-F, 3A-D and

4A-D. Tarsus I with five setal pairs: vs serrated, tc, a, p and u slender and nude, u bifurcated at tip. Setae

tc longest (Fig. IF). Setae lengths: v’ on femur I 26 (21-27), v” on femur I 27 (20-30), /’ on femur I

28 (21-30), d on femur I 24 (19-24), dorsal seta on genu I 250 (210-265), dorsal seta on genu II 275

(210-280), dorsal seta on genu III 335 (230-340), dorsal seta on genu IV 360 (215-360), solenidion

co on tarsus I 6 ( 6 ), co on tarsus II 5 (4-5). All tarsi with ambulacrum bearing a pair of claws and an

empodium with two rows of tenent hairs. Counts of setae on legs I—IV, indicating additional solenidia

in parenthesis: coxae: 3-1-2-2, trochanters 1—1—1—1, femora 4—3-2-2, genua l(/c)-l(/c)-l-l, tibiae

9(p)-&((p)-8((p)-l((p), tarsi 10(m)-10(m)-8-8.

Male (Figs 2A-B, 3E-H, 4E-H), range of three paratypes

Gnathosoma. 72-81 long and 65-72 wide. Subcapitular setae m 25-26, setae Orl 9, Or2 8-9, distance

m-m 17-19. Chelicerae 20-24 long. Length of palpal setae: femur d 18-20 and / ’ 33-34, and genu d 35.

Idiosoma. 195-210 long (gnathosoma excluded), 170-180 wide. Dorsum (Fig. 2A). Setae shorter than

those of females, cl longest and hi shortest. Length of setae: vi 35-38, ve 42, sci 38—42, see 37-38,

pebe 42-44, cl 51-53, c2 47, dl 40, d2 40-41, el 45-46, e2 34-35,// 24-25, f2 30, hi 9-10, h2 22.

Distances between setae: vi-vi 40—45, ve-ve 65-69, vi-ve 20-25, sci-sci 80-87, sce-sce 105-112,

sci-sce 10-11, cl-dl 39-44, dl-dl 8-10, dl-d2 55-59, dl-el 30-33, ve-sci 15-17, pdx-pdx 9-10,

pdx-cl 25-31, cl-cl 9-10, cl-c2 65-75, c2-c2 124-133, el-el 8 , el-fl 32-35, fl-hl 22-25, el-e2

48-50, d2-e2 26-31,//-// 8 ,fl-f2 35-42 ,f2-f2 63-66, e2-f2 40^13, hl-hl 8 , hl-h2 14-15, h2-h2

34-38, f2—h2 15-16. Venter (Fig. 2B). Agenital setae (ag) absent. Genital seta (g) very close to ps3.

Length of setae: la 16-18, lb 20-23, lc 44-41, 2c 35-37, 3a 17-18, 3b 22-24, 3c 25, 4a 12-13, 4b

12-13, 4c 16-2 1,psl 9,ps2 9,ps3 9, g 10.

Legs. Leg IV longest. Lengths (excluding ambulacra): leg I 465^175, leg II 405^110, leg III 440^150,

leg IV 475—485. Podomere lengths: femur I 150-155, femur II 115-125, femur III 120, femur IV 130—

135, tibia I 155, tibia II 125-130, tibia III 155-160, tibia IV 155-160, tarsus I 58-60, tarsus II 58-60,

tarsus III 60, tarsus IV 65. Tibia I with two solenidia (cp and cp2), and all tarsi with one long basal

solenidion (co) (Fig. 4E-H). Leg setae as indicated in Fig. 3E-H and 4E-H. Setae lengths: v ’ on femur I

21-22, v” on femur 125-27, / ’on femur 122, d on femur 124-25, dorsal seta on genu I 110-125, dorsal

seta on genu II 130, dorsal seta on genu III 185-195, dorsal seta on genu IV 215-230, solenidion co on

tarsus 127-28, co on tarsus II 30, co on tarsus III 30, co on tarsus IV 27-28. Counts of setae on legs I-IV,

indicating additional solenidia in parenthesis: coxae: 3-1-2-2, trochanters 1—1—1—1, femora 4—3-2-2,

genua l(/c)-l(/c)-l-l, tibiae 9 (cp, cp2)-%(cp)-%(cp)-l(cp), tarsi 10 (m)- 10 (m)- 8 (m)- 8 (m).

European Journal of Taxonomy 202 : 1-25 ( 2016 )

Protonymph (Figs 2C-D, 3I-L, 4I-L)

Gnathosoma. 70 long and 65 wide. Subcapitular setae m 20, setae Orl 5, Or2 6, distance m—m 14.

Chelicerae 24 long. Length of palpal setae: femur d 13 and /’ 28, and genu d 27.

Idiosoma. 295 long (gnathosoma excluded), 230 wide. Dorsum (Fig. 2C). Setae shorter than those of

females, cl longest. Length of setae: vi 41, ve 40, sci 38, see 43,pdx 46, cl 60, c2 46, dl 56, d2 39, el

56, e2 40, fl 50, f2 35, hi 30, h2 22. Distances between setae: vi—vi 48, ve-ve 83, vi—ve 18, sci-sci 97,

sce-sce 130, sci-sce 39, cl-dl 42, dl-dl 11, dl-d2 69, dl-el 38, ve-sci 14, pdx-pdx 17, pdx-cl 26,

cl—cl 10, cl-c2 78, c2-c2 150, c7-c7 11, c7-/7 45, fl-hl 38, c7-c2 54, <72-c2 36,/7-/7 ll,/7-^2 44,

/V/2 86, e2-^2 42, hl-hl 5, hl-h2 18, h2-h2 35, f2—h2 23. Venter (Fig. 2D). Setal pairs 4b , 4c, ag and

g absent. Length of setae: 7a 21, lb 23, 7c 35, 2c 34, 3a 26, 3/? 27, 3c 11, 4a 14, jot 7 9,p^2 9,/?.s3 9.

Legs. Leg I longest. Lengths (excluding ambulacra): leg I 375, leg II 340, leg III 315, leg IV 325.

Podomere lengths: femur I 130, femur II 95, femur III 100, femur IV 100, tibia I 115, tibia II 95, tibia

III 115, tibia IV 105, tarsus I 55, tarsus II 55, tarsus III 55, tarsus IV 55. Leg setae as shown in Figs 3I-L

and 4I-L. Setae lengths: v” on femur I 21, V on femur I 20, d on femur 118, dorsal seta on genu I 185,

dorsal seta on genu II205, dorsal seta on genu III 240, dorsal seta on genu IV 210, solenidion co on tarsus

I 5, co on tarsus II 4. Counts of setae on legs I—IV, indicating additional solenidia in parenthesis: coxae:

Fig. 3. Schematic leg setations of Neophyllobius cibyci sp. nov. A-D. holotype. A. Trochanter-tibia of

leg I. B. Trochanter-tibia of leg II. C. Trochanter-tibia of leg III. D. Trochanter-tibia of leg IV. E-H.

paratype (CNAC009238). E. Trochanter-tibia of leg I. F. Trochanter-tibia of leg II. G. Trochanter-tibia

of leg III. H.Trochanter-tibia of leg IV. I-L. Protonymph, paratype (CNAC009241). I. Trochanter-

tibia of leg I. J. Trochanter-tibia of leg II. K. Trochanter-tibia of leg III. L. Trochanter-tibia of leg

IV. M-O. Larva, paratype (CNAC009242). M. Trochanter-tibia of leg I. N. Trochanter-tibia of leg IT

O. Trochanter-tibia of leg III.

PAREDES-LEON R. et al., New species of camerobiid mites and setal homology in Neophyllobius

3-1-2-0, trochanters 1—1—1—0, femora 3—2—1—1, genua 1 (/c)—1 (/c)—1—1, tibiae 6((p)-5((p)-5{(p)-3((p),

tarsi 9(co)-9(oj)-7-5.

Larva (Figs 2E-F, 3M-0, 4M-0), range of three paratypes

Gnathosoma. 45-50 long and 60 wide. Subcapitular setae m and palpal femur seta d absent, setae Orl

88, Or2 8. Chelicerae 20-22 long. Length of palpal setae: femur /’ 20-21 and genu d 20.

Idiosoma. 165-180 long (gnathosoma excluded), 130-135 wide. Dorsum (Fig. 2E). Setae shorter than

those of females, c2 longest. Setal pair pdjc absent. Length of setae: vi 28-29, ve 30-34, sci 34, see 34, cl

35-38, c2 42M3, dl 32-34, d2 28, el 35-36, 25-27, fl 30-34 ,f2 23-24, hi 17-18, h2 15. Distances

between setae: vi—vi 30-35, ve-ve 58-60, vi—ve 18-20, sci-sci 65-68, sce-sce 85, sci-sce 28-29, cl-dl

43M6, dl-dl 10, dl-d2 37-38, dl-el 25-27, ve-sci 10-11, vi—cl 35—41, cl-cl 10, cl-c2 55-60, c2-

c2 105-110, el-el 10, el-fl 25-30, fl-hl 22-24, el-e2 35-39, d2-e2 21-49 Jl-fl 9-\9Jl-f2 25-26,

f2-f2 50-55, e2-f2 30-32, hi-hi 8, hl-h2 9, h2-h2 20-22, f2-h2 22-24. Venter (Fig. 2F). Setal pairs

lc, 2c, 3b, 3c, 4b, 4c, ag and g absent. Length of setae: la 20-24, lb 22-23, 3a 25, psl 1,ps2 1 ,ps3 8.

Legs. Leg I longest. Lengths (excluding ambulacra): leg 1260-265, leg II240, leg III 250-255. Podomere

lengths: femur I 80, femur II 60, femur III 70-75, tibia I 60-65, tibia II 55, tibia III 65-70, tarsus I 45,

tarsus II 45, tarsus III 45. Leg setae as indicated in Figs 3M-0 and 4M-0. Setae lengths: v” on femur I

32-34, d on femur 122, dorsal seta on genu 143M5, dorsal seta on genu II 33, dorsal seta on genu III 45,

solenidion co on tarsus I 3, co on tarsus II3. Counts of setae on legs I—III, indicating additional solenidia in

parenthesis: coxae: 2-0-0, trochanters 0-0-0, femora 2-2-1, genua 1 (k)-\ {k)-\ , tibiae 3{(p)-3{(p)-3{(p),

tarsi 7(co)-7(co)-5.

Intraspecific variation

In the case of Neophyllobius cibyci sp. nov., we observed that leg setation teratologies are relatively

rare, but include the following: 1) adult specimens with certain setae in different locations; for example,

female right femur I with v ” distal to d instead of proximal (CNAC009236), female right femur I with

/’positioned before d (CNAC009234), female left femur I with / ’positioned before d (CNAC009233),

female tibiae I with seta dm between v’ and v” (CNAC009231 and CNAC009235) instead of distal,

female right tibia I with dm between v ’ and v ” (CNAC009232) instead of distal, and male right tibia I

with / p distal to /”p instead of proximal (CNAC009238); 2) female specimens without setae on left or

right legs, for example left femur I without /’ (CIUM), left femur I without v” (CNAC009236), and left

tibia I without v” (CNAC009236 and CIUM).

Remarks

Neophyllobius cibyci sp. nov. resembles N. farrieri De Leon, 1958 collected on “Spanish moss”

(probably Tillandsia usneoides ) in Oaxaca, Mexico. In both species, dorsal setae on genua I-IV are very

long, extending beyond the tip of the tarsus, setae on femur I are short (about Ve of podomere length),

and setae /’ and d on femur I are nearly located at the same level. These species can be differentiated

by the following characters: in N. cibyci sp. nov., dorsal setae cl and dl are distinctly longer than the

distance between setae cl-dl and dl-el respectively, setae dl are the longest of the dorsal setae, and

setae d and / ’ on palpal femur are heavily spinose. In N. farrieri, the lengths of dorsal setae cl and dl

are the same as the distance between setae cl-dl and dl-el respectively; setae el are the longest of the

dorsal setae, and setae d and / ’ on palpal femur are weakly spinose.

Neophyllobius cibyci sp. nov. includes at least three postembryonic instars (larva, protonymph and

adult). We found only one nymphal instar (protonymph) which resembles the female type proposed by

Bolland (2001).

European Journal of Taxonomy 202 : 1-25 ( 2016 )

Fig. 4. Schematic tarsal setations of Neophyllobius cibyci sp. nov. A-D. 5 , holotype. A. Tarsus I.

B. Tarsus II. C. Tarsus III. D. Tarsus IV. E-H. S, paratype (CNAC009238). E. Tarsus I. F. Tarsus II.

G. Tarsus III. H. Tarsus IV. I-L. Protonymph, paratype (CNAC009241). I. Tarsus I. J. Tarsus II. K. Tarsus

III. L. Tarsus IV. M-O. Larva, paratype (CNAC009242). M. Tarsus I. N. Tarsus II. O. Tarsus III.

PAREDES-LEON R. et al., New species of camerobiid mites and setal homology in Neophyllobius

Ecology

Neophyllobius cibyci sp. nov. was the most abundant camerobiid species with 17 specimens, most of

which (76.5%) were collected in dry season (March and April). This species is deemed to be an inhabitant

of the tree canopy because most specimens (94%) were collected on epiphytic bromeliads at 2.8 m (± 0.4)

and only one female specimen (CNAC009237) was found on soil litter. Neophyllobius cibyci sp. nov.

was found in low numbers, usually one specimen per sample, except for two samples which had two

specimens each. The closely resembling species, N. farrieri , also inhabits epiphytic bromeliads of the

genus Tillandsia.

Distribution

This species is only known from San Andres de la Cal, Morelos, Mexico. The type locality of N. cibyci

sp. nov. is about 440 km as the crow flies from the known distribution of N. farrieri (Puenta [sic. ] de

Nejapa, Oaxaca, Mexico).

Setal homology among Neophyllobius cibyci sp. nov. instars

The collection of different instars provides material for study and suggests the ontogenetic development

pattern of gnathosoma, idiosoma and leg chaetotaxy. Although, Grandjean’s idiosomal and leg chaetotaxy

systems were previously applied to camerobiid mites (Kethley 1990; Fan & Walter 2011), this is the

first study of complete chaetotaxy (on gnathosoma, idiosoma and legs) involving three different instars

(larva, protonymph and adult) of a camerobiid mite species. Swift’s (2001) study of leg chaetotaxy

of Caligonellidae (Raphignathina: Raphignathoidea), included one larva and one adult female of an

undetermined species of Neophyllobius. We found only one nymphal instar (protonymph); for this

reason and because the presence of deutonymph is questionable in Neophyllobius , we prefer not to

hypothesise the presence of setae in this particular instar (deutonymph).

There are different concepts about the leg chaetotaxy. Fundamental setae are those present when the

appendages in question are first formed, that is on legs I—III in the larva or on leg IV in the protonymph.

Accessory setae are any which are formed during subsequent moults (Grandjean 1941; Norton 1977;

Swift 2001). Setal priority for each podomere refers to a list of setal organs that appear at the beginning

of ontogeny (fundamental setae) and have greater priority, or force, than organs than develop later

(accessory setae), which suggests that these setal organs are less susceptible to evolutionary regression

(Norton 1977; Swift 2001). Setae of equal priority are listed together within parentheses as proposed by

Norton (1977).

Gnathosoma. Larvae have Orl and Or2. Setal pattern on palps is consistent across larva, protonymph

and adults, except that dorsal seta (d) on femur do not appear until the protonymphal instar; count of

setae from palp coxa to tarsus: 1 ( elcp ), 0, 2, 1, 3 + 1 claw, 4 + co. Setal pair m on subcapitulum do not

appear until the protonymphal instar.

Idiosoma. The dorsal chaetotaxy in larvae includes 14 setal pairs: vi, ve, sci, see, cl, c2, dl, d2, el, e2,

fl,f2, hi and h2 (Fig. 2E). Setal pair pdx appears in protonymph (Fig. 2C). The ventral chaetotaxy in

larvae includes six setal pairs: la, lb, 3a,psl,ps2 and ps3 (Fig. 2F). Setal pairs lc, 2c, 3b, 3c and 4a are

added in protonymph (Fig. 2D). Setal pairs 4b and 4c are added in adults (Figs ID, 2B). Setal pair ag is

present only in adult females (Fig. ID).

Legs (Table 2). Setae v are absent on trochanters I—III in larva (Fig. 3M-0), and do not appear until the

protonymphal instar (Fig. 3I-K). Seta v on trochanter IV does not appear until adulthood (Fig. 3D, H).

Femora I—II of the larva with setae v ” and d (Fig. 3M-N), femur III of the larva (Fig. 30) and femur IV

of the protonymph (Fig. 3L) only with seta d. Seta V is added on femur I of the protonymph (Fig. 31).

Setae v ’ on femora I—III (Fig. 3A-C, E-G), and /” on femur IV (Fig. 3D, H) are added in adults. The setal

European Journal of Taxonomy 202 : 1-25 ( 2016 )

Table 2. Setal leg ontogeny of Neophyllobius cibyci sp. nov., parentheses represent a setal pair and

square brackets indicate additional setae in 3.

LEG

INSTAR

TROCHANTER

FEMUR

GENU

TIBIA

TARSUS

Larva

d,v”

d, k

dm, (I), cp

vs ’, (tc), (p), (u), CO

Protonymph

d, v ’

d, k

dm, (J),(p, dp, V’p, v 'm

vs”, (tc), (p), (u), (a), co

Adult

d,(v),V

d, k

d, (/), (ym), dp, (Ip), dm, cp, [cp2]

(vs), (tc), (p), (u), (a), co

Larva

d,v”

d, k

dm, (I), cp

vs (tc), (p), (u), co

Protonymph

d,v”

d, k

dm, (/), cp, dp, 1 ”p

vs (tc), (p), (u), (a), co

Adult

d, (v)

d, k

d, (I), v’m, dp, (Ip), dm, cp

(vs), (tc), (p), (u), (a), co

Larva

dm, (I), cp

vs ’, (tc), (u)

III

Protonymph

dm, (1), (p, dp, V’p

vs”, (tc), (p), ( u)

Adult

d, v’

dm, (/), cp, (Ip), dp, d, v 'm

(vs), (tc), (p), (u), [co]

Protonymph

dp, v 'm, 1”, cp

vs \ (tc), (u)

Adult

d, 1”

dp, v ’m, (l), cp, d, V’p, dm

(vs), (tc), (p), (u), [co]

priorities on femora I and II for Neophyllobius cibyci sp. nov. are: (d, v”), l v \ Priorities on femora III

and IV are: d, l”, v ’. Genua I—II in all instars with seta d and k (Fig. 3A-B, E-F, I—J, M-N), and genu III

only with d (Fig. 3C, G, K, O). Genu IV of the protonymph and adults only with seta d (Fig. 3D, H, L).

Tibiae I—III of larva with setae dm, l’, l” and cp (Fig. 3M-0). Tibia IV of the protonymph with setae dp,

v ’m, /” and (p (Fig. 3L); dp and l”p are added to tibiae I—III (Fig. 3I-K) of the protonymph. Seta v ’m is

added to tibia I (Fig. 31) of the protonymph. Setae / ’p, and d are added to tibiae I—II of adults (Fig. 3A-B,

E-F). Seta v ”m appears only on tibiae I of adults (Fig. 3A, E). Setae l p and v ’m are added to tibia III of

adults (Fig. 3C, G). Setae l \ dp, l”p and d are added to tibia IV of adults (Fig. 3D, H). Tibia I of males

with (p2 (Fig. 3E). The setal priorities for tibiae I and II in Neophyllobius cibyci sp. nov., are: (dm, l’, /”),

(dp, V’p, v ’m), (d, l ’p, v”m). Priorities on tibiae III and IV are similar except v”m is not formed.

Tarsi I—II of the larva with vs’, ( tc ), (p), (u) and co (Fig. 4M-N), tarsus III with vs’, ( tc ), (u) and co

(Fig. 40). In tarsi I—II of protonymph, setal pair a is added (Fig. 4I-J). Seta vs ’ is lost on tarsi I and III

of protonymph but vs ” appears (Fig. 41, K). Tarsus IV of the protonymph with vs ’, (tc) and (u) (Fig. 4L).

Setae vs’ reappears in tarsi I and III of adults (Fig. 4A, C, E, G). Seta vs” appears on tarsi II and IV

(Fig. 4B, D, F, H). Setal pair p is added in tarsus IV of the adults (Fig. 4D, H). Tarsi III-IV of males

with co (Fig. 4G-H). The setal priorities for tarsi I and II are: (tc ’, tc ”, u’,u ”, vs ’,p’,p ”), (a ’, a ”, vs ”).

Priorities on tarsi III and IV are: (tc ’, tc ”, u’,u ”, vs ’), (p ’, p ”, vs ”).

Neophyllobius tepoztlanensis sp. nov.

um:lsid:zoobank.org:act:A8A8CE4B-824C-40D7-A435-45A8E4401A4D

Fig. 5A-F

Diagnosis

This species is characterized as follows: dorsal setae reaching setae immediately behind, seta d on femur

I positioned in front of / ’, femur II with three setae (d, v ’ and v ”), femur III with two setae (d and v ’),

setae d on genua I-IV slightly less than half as long as tibiae I-IV respectively, dorsal seta dl less than

half as long as width of body; dorsal setae cl as long as setae dl, setae v” on femora I—II positioned

distinctly in front of v ’.

PAREDES-LEON R. et al., New species of camerobiid mites and setal homology in Neophyllobius

Etymology

The specific name makes reference to the municipality Tepoztlan, where the type locality is situated.

The name of the municipality derives from two words in the nahuatl language, “Tepoztecatl” (a nahua

divinity) and “tlan” (beside), so its meaning is “in the company of Tepoztecatl".

Type material

Holotype

MEXICO: 5, CNAC009245, ex Tillandsia schiedeana at 2.4 m on Sapium macrocarpum. Collected

from the type locality on 12 Mar. 2014, O. Cortes and R. Paredes coll. (RPL1232).

Paratypes

MEXICO: 1 5, CIUM, ex T. hubertiana at 3.2 m on S. macrocarpum , 30 Apr. 2014 (RPL1267); 1 $,

CNAC009246, ex T. hubertiana at 2.6 m on S. macrocarpum , 22 Aug. 2014, S. Gomez and R. Paredes

coll. (RPL1303). Both specimens collected from the type locality, with same data as holotype except

where noted.

Type locality

MEXICO: Morelos, Tepoztlan, 1 km S of San Andres de la Cal, 18.94408° N, 99.11924° W, 1478 m a.s.l.

Description

Female (n = 3) (Fig. 5A-F)

Holotype female (followed in parentheses by range of holotype and two paratype females).

Gnathosoma. 68 (68-79) long and 77 (71-82) wide. Subcapitulum with subcapitular setae m 20 (20-24)

and two pairs of adoral setae Orl 9 (9) and Or2 10 (8-10); these three pairs nude and slender, m longest

(Fig. 5B); distance m-m 20 (20-21). Chelicerae 40 (27—40) long. Palp, five-segmented with following

setal distribution: trochanter without setae; femur with two serrated setae, d 19 (17-19) and V 31 (27-

31); genu with one long, slender, nude dorsal (d) seta 31 (30-32); tibia with three tactile setae (/ ’, 1” and

d) and one claw (sword-like seta); tarsus with two eupathidia (acmC and sulQ, two simple setae (ba and

va) and one small solenidion (co) (Fig. 5A). Setae elcp present.

Idiosoma. Longer than wide, 285 (225-285) long (gnathosoma excluded), 230 (200-235) wide. Cuticle

striated, except on coxae and attenuated between setal pair el (Fig. 5C-D). Dorsum. With 15 pairs of

serrated setae set on small tubercles; each setal pair just reaching next setal pairs (in transversal row),

c2 longest and h2 shortest. Two pairs of eyes are positioned between setae sci and see. Length of setae:

vi 48 (48-53), vc 46 (46^18), sci 43 (43-50), see 46 (46M8), pdx 46 (46-53), cl 50 (50-54), c2 63

(53-63), dl 51 (50-56), d2 40 (40M4), el 53 (53-56), c2 46 (40^16), fl 50 (50-54), f2 30 (30-32),

hi 34 (32-34), h2 28 (27-28). Distances between setae: vi-vi 40 (40-50), ve-ve 82 (64-94), vi-ve 28

(24-30), sci-sci 98 (80-110), scc-scc 130 (115-140), sci-sce 40 (34M0), cl-dl 46 (42-52), dl-dl

15 (14-15), dl-d2 70 (62-78), dl-el 40 (38M3), ve-sci 19 (19-21), pdx-pdx 12 (11-12), pdx-cl 31

(29-31), cl-cl 15 (14-15), cl-c2 77 (70-86), c2-c2 160 (140-170), el-el 12 (10-12), el-fl 42 (34-

42), fl-hl 56 (38-56), el-e2 54 (52-62), d2-e2 34 (34-37), fl-fl 11 (9-12), fl-f2 53 (43-53), f2-f2

91 (83-94), e2-f2 50 (30-50), hl-hl 8 (8-11), hl-h2 25 (23-25), h2-h2 57 (54-57), J2-h2 42 (26^12).

Venter. Coxal setae slenderer than dorsal setae. Setal pairs la, 2a, 4a, ag, g,psl,ps2 and ps2 nude and

short. Setae lb, lc, 2c, 3b, 3c, 4b and 4c serrated. Setae lc longest and g shortest. Setal pairs 4a and ag

located on individual platelets. Genito-anal valves with one pair of genital setae (g). Coxa I grouped

with coxa II, and coxa III with IV but not completely fused (Fig. 5D). Length of setae: la 32 (27-32), lb

22 (21-22), lc 50 (42-50), 2c 42 (35-42), 3a 37 (34-37), 3b 30 (25-30), 3c 31 (30-31), 4a 19 (19-30),

4b 18 (14-18), 4c 20 (19-20), ag 16 (12-16), g 9 (9-12), psl 14 (9-15), ps2 11 (10-12), ps3 12 (10-14).

European Journal of Taxonomy 202 : 1-25 ( 2016 )

Fig. 5. Neophyllobius tepoztlanensis sp. nov., holotype. A. Palp. B. Subcapitulum. C. Dorsal idiosoma.

D. Ventral idiosoma. E. Trochanter-tibia of leg I. F. Tarsus I.

PAREDES-LEON R. et al.. New species of camerobiid mites and setal homology in Neophyllobins

Legs. Slender and long, leg IV longest. Lengths (excluding ambulacra): leg I 500 (420-510), leg II 415

(390-450), leg III 455 (417-485), leg IV 505 (470-530). Podomere lengths: femur I 160 (160-165),

femur II 120 (110-130), femur III 125 (115-135), femur IV 140 (130-155), tibia I 145 (135-160),

tibia II 115 (110-125), tibia III 130 (130-165), tibia IV 165 (155-175), tarsus I 59 (54-61), tarsus II 58

(56-61), tarsus III 60 (57-66), tarsus IV 66 (62-66). Leg setae as indicated in Fig. 5E-F. Tarsus I with

five setal pairs: vs serrated, tc, a , p and u slender and nude, u bifurcated at tip. Setae tc longest. Setae

lengths: v ’ on femur I 34 (30-34), v ” on femur I 34 (30-37), /’ on femur I 38 (31-38), d on femur 131

(30-31), dorsal seta on genu I 54 (48-54), dorsal seta on genu II 45 (39—45), dorsal seta on genu III 45

(42-47), dorsal seta on genu IV 71 (68-71). All tarsi with ambulacrum bearing a pair of claws and an

empodium with two rows of tenent hairs. Counts of setae on legs I-IV, indicating additional solenidia

in parenthesis: coxae: 3-1-2-2, trochanters 1—1—1—1, femora 4-3-2-2, genua 1 (/e)— 1 (a:)— 1—1, tibiae

9(^)-8(y?)-8(y?)-7(^), tarsi 10(m)-10(m)-8-8.

Remarks

Neophyllobius tepoztlanensis sp. nov. resembles N. marginatus De Leon, 1958 collected from an

undetermined composite plant and Quercus sp. in Nayarit, Mexico. In both species, the dorsal setae

on genua I-IV are short, less than half as long as tibiae I-IV; the setae on femur I are short (about a

quarter of the length of the podomere). These species can be differentiated by the following characters:

in N. tepoztlanensis sp. nov., dorsal setae cl as long as setae dl , and setae v” on femora I—II positioned

distinctly in front of v ’. In N. marginatus , dorsal setae cl about half as long as seta dl , and setae v ’ and

v” on femora I—II are positioned horizontally on nearly the same level.

Ecology

Neophyllobius tepoztlanensis sp. nov. is deemed to be an inhabitant of the tree canopy and was found on

Tillandsia schiedeana and T. hubertiana at 2.7 m (± 0.4) on Sapium macrocarpum.

Distribution

This species is only known from San Andres de la Cal, Morelos, Mexico. The type locality of

N. tepoztlanensis sp. nov. is about 600 km from Ixtlan del Rio, Nayarit, Mexico and 660 km from Tepic,

Nayarit, the two known localities of N. marginatus.

Neophyllobius tescalicola sp. nov.

um:lsid:zoobank.org:act:C58F130B-B7EC-423F-8AB5-EF5C0216EC0C

Fig. 6A-F

Diagnosis

This species is unique due to a combination of following characters: dorsal setae reaches setae

immediately behind, femur II with three setae (d, v ’ and v ”), femur III with two setae (d and v ’), setae

d on genua I-IV passing tibiae, long setae (about V 3 of podomere length) on femur I; setae d on femur

I positioned behind setae / ’, dorsal idiosomal setae pdx and cl grouped on small and finely-striated

platelet.

Etymology

The specific name refers to the parent rock of the soil (lava flow) in which soil litter, the habitat of this

mite species, is deposited. The word “tescal” in the nahuatl language means covered basalt stone from

ancient volcanic eruptions or volcanic lava field, and the latin suffix “cola” means inhabitant (one who

inhabits). So, tescalicola means “inhabitant of volcanic lava fields”.

European Journal of Taxonomy 202 : 1-25 ( 2016 )

Type material

Holotype

MEXICO: 5, CNAC009247, on soil litter of Quercus obtusata. Collected from the type locality on

12 Mar. 2014, O. Cortes and R. Paredes coll. (RPL1224).

Paratypes

MEXICO: 2 $ §, CNAC009248, 1 CIUM, on soil litter of Quercus obtusata, 23 Oct. 2014, R. Reyes

and R. Paredes coll. (RPL1317). All collected from the type locality, with same data as holotype except

where noted.

Type locality

MEXICO: Morelos, Tepoztlan, 1 km S of San Andres de la Cal, 18.94329° N, 99.11784° W, 1485 m a.s.l.

Description

Female (n = 3) (Fig. 6A-F)

Holotype female (followed in parentheses by range of holotype and two paratype females).

Gnathosoma. 88 (85-90) long and 82 (80-85) wide. Subcapitulum with subcapitular setae m 24 (24-27)

and two pairs of adoral setae Orl 15 (12-15) and Or2 8 (8); these three pairs nude and slender, m longest

(Fig. 6B); distance m—m 22 (20—23). Chelicerae 24 (23-25) long. Palp, five-segmented with following

setal distribution: trochanter without setae; femur with two serrated setae, d 16 (15-16) and l’ 27 (25-

28); genu with one long, slender, nude dorsal (d) seta 37 (35-38); tibia with three tactile setae (/’, 1” and

d) and one claw (sword-like seta); tarsus with two eupathidia (acmC and sulQ, two simple setae (ba and

va) and one small solenidion (co) (Fig. 6A). Setae elcp present.

Idiosoma. Longer than wide, 340 (330-345) long (gnathosoma excluded), 305 (300-310) wide. Cuticle

striated, except on coxae and attenuated between setal pairs pdx, cl and dl (Fig. 6C-D). Dorsum.

With 15 pairs of serrated setae set on small tubercles; all setal pairs longer than distance to setal pairs

immediately behind, e2 longest and h2 shortest. Two pairs of eyes are positioned between setae sci and

see. Setal pairs pdx and cl associated in a small and weakly striated platelet (Fig. 6C). Length of setae: vi

81 (80-81), vc 77 (76-77), sci 70 (70), see 77 (75-80), pdx 71 (70-73), cl 80 (80-83), c2 84 (84-85), dl

11 (75-77), d2 74 (74-75), el 84 (84-85), c2 87 (85-90), fl 84 (83-84), f2 40 (39M1), hi 45 (43^15),

h2 36 (35-36). Distances between setae: vi-vi 53 (50-53), ve-ve 99 (99-100), vi-ve 36 (35-36), sci-

sci 120 (120-125), sce-sce 160 (160-165), sci-sce 42 (40M3), cl-dl 53 (50-53), dl-dl 19 (18-19),

dl-d2 88 (85-90), dl-el 54 (53-54), ve-sci 20 (18-20), pdx-pdx 20 (20), pdx-cl 26 (25-28), cl-cl

19 (18-20), cl-c2 91 (90-95), c2-c2 185 (180-185), el-el 12 (11-12), el-fl 37 (36-38), fl-hl 53

(53-55), el-e2 67 (65-67), d2-e2 54 (54-55), fl-fl 12 ( \2),fl-f2 56 (55-56 \f2-f2 70 (70-73), e2-f2

68 (65-70), hl-hl 9 (9), hl-h2 23 (20-25), h2-h2 42 (40M3), f2-h2 28 (27-28). Venter. Coxal setae

slenderer than dorsal setae. Setal pairs la, 2a, 4a, ag, g, psl, ps2 and ps3 nude and short. Setae lb,

lc, 2c, 2b, 3c, 4b and 4c serrated. Setae lc longest and ps2 shortest. Setal pairs 3a and 4a located on

individual platelets. Genito-anal valves with one pair of genital setae (g). Coxa I grouped with coxa II,

and coxa III with IV but not completely fused (Fig. 6D). Length of setae: la 29 (28-29), lb 19 (15-20),

lc 58 (55-58), 2c 51 (50-55), 3a 21 (20-21), 3b 27 (27-30), 3c 30 (30), 4a 22 (20-22), 4b 26 (25-27),

4c 30 (30), ag 20 (20), g 14 (13-14), psl 14 (13-14), ps2 12 (11-12 ),ps3 14 (13-14).

Legs. Slender and long, leg IV longest. Lengths (excluding ambulacra): leg I 660 (650-665), leg II 570

(565-580), leg III 660 (660-675), leg IV 750 (745-765). Podomere lengths: femur I 215 (210-215),

femur II 170 (170-180), femur III 185 (185), femur IV 225 (220-230), tibia I 215 (210-220), tibia II

185 (180-190), tibia III 235 (230-235), tibia IV 265 (260-270), tarsus I 80 (80), tarsus II 82 (80-83),

tarsus III 83 (81-84), tarsus IV 86 (86-90). Leg setae as indicated in Fig. 6E-F. Tarsi I with five setal

_ f

PAREDES-LEON R. et al ., New species of camerobiid mites and setal homology in Neophyllobins

Fig. 6. Neophyllobiiis tescalicola sp. nov., $, holotype. A. Palp. B. Subcapitulum. C. Dorsal idiosoma.

D. Ventral idiosoma. E. Trochanter-tibia of leg I. F. Tarsus I.

European Journal of Taxonomy 202 : 1-25 ( 2016 )

pairs: vs serrated, tc , a,p and u slender and nude, u bifurcated at tip. Setae tc longest. Setae lengths: v’

on femur I 80 (75-85), v” on femur I 67 (65-70), /’ on femur I 94 (94-98), d on femur I 68 (65-70),

dorsal seta on genu I 250 (245-250), dorsal seta on genu II250 (250), dorsal seta on genu III 285 (280-

290), dorsal seta on genu IV 341 (340-345). All tarsi with ambulacrum bearing a pair of claws and an

empodium with two rows of tenent hairs. Counts of setae on legs I-IV, indicating additional solenidia

in parenthesis: coxae: 3-1-2-2, trochanters 1—1—1—1, femora 4—3-2-2, genua l(/c)-l(/c)-l-l, tibiae

9(^)-8(^)-8(^)-7(^), tarsi 10(m)-10(m)-8-8.

Remarks

Neophyllobius tescalicola sp. nov. resembles N. farrieri and N. cibyci sp. nov. In these three species, the

dorsal setae on genua I-IV are very long, extending beyond the tip of the tarsus. These species can be

differentiated by the following characters: in N. tescalicola sp. nov., setae on femur I are long, about A

of podomere length, setae d on femur I is positioned behind setae / ’, dorsal idiosomal setae pdx and cl

are grouped on a small and finely-striated platelet. In N. farrieri and N. cibyci sp. nov., setae on femur

I are short, about a sixth of the length of the podomere, setae / ’ and d on femur I are nearly positioned

horizontally at same level, and dorsal idiosomal setae pdx and cl are not joined on a platelet.

Ecology

Neophyllobius tescalicola sp. nov. inhabits soil litter of Quercus obtusata , and was collected in dry

(March) and in rainy seasons (October).

Distribution

This species is only known from the type locality.

Key to the species of the genus Neophyllobius in Mexico

Based on females, updated from De Teon (1958).

1. Dorsal setae c 1, dl, el and fl not reaching setae immediately behind.

. Neophyllobius mexicanus McGregor, 1950

- At least one of these dorsal setae cl, dl, el and fl reaching or extending beyond the seta immediately

be hin d.2

2. Femur II with four setae. N. quadrisetosus De Teon, 1958

- Femur II with three setae.3

3. Femur III with three setae. N. trisetosus De Teon, 1958

- Femur III with two setae.4

4. Seta d on genu IV longer than or about as long as tibia.5

- Seta d of genu IV distinctly shorter than tibia IV.10

5. Setae d of genua III-IV extending beyond ends of respective legs.6

- Setae d of genua III-IV not extending beyond ends of respective legs.8

6. Femur I with long setae, about A of podomere length; setae / ’ on femur I positioned before setae d,

dorsal idiosomal setae pdx and cl grouped on a small and finely striated platelet.

. Neophyllobius tescalicola sp. nov.

- F emur I with short setae, about Ve of podomere length; setae / ’ and d on femur I positioned horizontally

nearly at same level; dorsal idiosomal setae pdx and cl not joined on a platelet.7

PAREDES-LEON R. et al.. New species of camerobiid mites and setal homology in Neophyllobins

7. Length of dorsal setae cl and dl same as distance between setae cl-dl and dl-el respectively; setae

el longest of dorsal setae; setae d and / ’on palpal femur, weakly spinos e... JV. farrieri De Leon, 1958

- Dorsal setae cl and dl distinctly longer than distance between setae cl-dl and dl-el respectively;

setae dl longest of dorsal setae; setae d and / ’on palpal femur, heavily spinose.

. Neophyllobius cibyci sp. nov.

8. Seta d of genu III distinctly shorter than tibia III . N. inequalis De Leon, 1958

- Seta d of genu III longer than or about as long as tibia III.9

9. Seta d on genu II shorter than tibia II, seta of genu III about as long as tibia III.

. N. equalis De Leon, 1958

- Setae d on genua II and III distinctly longer than respective tibiae... N. consobrinus De Leon, 1958

10. Striae of dorsum consisting chiefly of broken striae; posterior margins of femora I-IV, highly

serrulate. N. lobatus De Leon, 1958

- Striae of dorsum consisting chiefly of unbroken striae; posterior margins of femora, not highly

serrulate.11

11. Dorsal setae dl more than half as long as width of body; seta d on genu III about two-thirds as long

as tibia III. N. deleoni Bolland, 1991

- Dorsal setae dl less than half as long as width of body; seta d on genu III about one-half (or less) as

long as tibia III.12

12. Seta d on genu II distinctly longer than genu II.13

- Seta of genu II about as long as or shorter than genu II.15

13. Seta d on genu II nearly as long as seta of genu III and about half as long as seta of genu IV.14

- Seta d on genu II about half as long as seta of genu III and about one-third as long as seta d on

genu IV. N. longulus De Leon, 1958

14. Dorsal setae cl as long as setae dl\ setae v” on femora I—II distinctly in front of v’.

. Neophyllobius tepoztlcinensis sp. nov.

- Dorsal setae cl about half as long as setae dl\ setae v on femora I—II positioned horizontally on

nearly same level. N. marginatus De Leon, 1958

15. Genual setae coarse, setiform, distinctly spinose. N. curtipilus De Leon, 1958

- Genual setae linear (seta of genu II is slightly obovate), very faintly spinose.

. N. spatulus De Leon, 1958

Discussion

Some patterns of setal ontogeny found in Neophyllobius cibyci sp. nov. have also been found in other

Raphignathina. Lor example, the presence of setae v on trochanters I—III, retarded until the protonymphal

instar, was also reported in Caligonellidae (Swift 2001) and Pterygosomatidae (Paredes-Leon &

Guzman-Cornejo 2015).

The larva of N. cibyci sp. nov. bears the setal pair p (i.e., p ’ and p ”) on tarsus I instead of only p ’ as in

Neophyllobius sp. (Swift 2001). Also, tarsus IV in females ofV cibyci sp. nov. has one more seta (i.e.,

eight instead of seven like Neophyllobius sp.); however, Swift (2001) didn’t mention the nomenclature

of these seven setae, making the comparison with N. cibyci sp. nov. difficult. Another difference between

both species is the presence of iteral setae (it) on tarsi I-IV located anterodorsal to ( tc ) (Swift 2001).

Based on the location (anterodorsal to tc), we consider setae it to be absent in N. cibyci sp. nov., but setal

European Journal of Taxonomy 202 : 1-25 ( 2016 )

pair p to be present instead. The same condition (i.e., presence of setal pair p on tarsi I-IV instead of setal

pair it) was reported by Fan & Walter (2011) for Acamerobia inflatus (Camerobiidae).

The presence of setal pair it has been reported anterodorsal to setal pair p and a for some pterygosomatid

mites (Pterygosomatidae) only in tarsus I (Bochkov et al. 2008; Paredes-Leon et al. 2012; Paredes-

Leon & Guzman-Cornejo 2015). According to Khanjani et al. (2014), setal pair it is also absent in

Neophyllobius ostovani Khanjani, Hoseini, Yazdanpanah & Masoudian, 2014, and in N. lorestanicus

Khanjani, Hoseini, Yazdanpanah & Masoudian, 2014 adults. Compared with these latter species,

Neophyllobius cibyci sp. nov. has both setae of pair p (i.e.,/? ’ and p”) on tarsi I—II, whereas N. ostovani

has only one (p ’) on tarsi I—II. N. lorestanicus has both setae p on tarsus I but only p ’ on tarsus II.

In many other species of the genus Neophyllobius the counts of setae in adults, from coxa to tarsus,

appears to be consistent with that found in Neophyllobius cibyci sp. nov. (coxae: 3-1-2-2, trochanters

1—1—1—1, femora 4—3-2-2, genua l(/c)-l(/c)-l-l, tibiae 9(y?)—8(^>)—8(y?)—7(y?), tarsi 10(m)-10(m)-8-8),

such as N. lachishensis Bolland, 1998, N. piniphilus , N. demirsoyi Akyol & Kog, 2006 and N. quercus

Ulugay & Kog, 2014, among others (Bolland 2001; Akyol & Kog 2006; Ulugay & Kog 2014).

Teg setation teratologies similar to that shown by N. cibyci sp. nov. has also been reported in other

camerobiid mites, for example in N. consobrinus (Bolland & Swift 2000), Tycherobius stramenticola

Bolland, 1986 and T. polonicus Bolland, 1986 (Kog & Akyol 2007). In the last two species, abnormal

disposition of setae are also found in dorsal and ventral idiosoma (Kog & Akyol 2007).

Due to the lack of setal notations, and because many descriptions of camerobiid mites are based on single

specimens, it is very difficult to establish a homology hypothesis among instars and even among species.

Notwithstanding this fact, our data suggest a hypothesis to how leg setae appear at different life stages on

the podomeres of legs I-IV of larva, protonymph and adult. However, a detailed examination of the leg

chaetotaxy of taxa in Camerobiidae will have to be conducted to establish homologies within the family

and with the other raphignathoids.

The results presented here clearly show that the species richness of the family Camerobiidae in Mexico is

under-represented, and that systematic surveys sampling different components of vegetation are needed.

The species recorded had different habitat preferences, two of them inhabiting epiphytic bromeliads on

two tree species at an average height of 2.8 m (± 0.4), and one inhabiting soil litter of one tree species. This

suggests that habitat diversity determines the richness of the group but it needs to be studied in greater detail.

Acknowledgements

We are grateful to O. Cortes-Anzures, S. Gomez-Rosas and R. Reyes (CIByC) for field assistance.

Thanks to T.M. Perez and G. Montiel-Parra (CNAC) for allowing us to carry out acarological

procedures in their laboratory. Thanks to J.R. Ramirez-Rodriguez and G. Flores-Franco (CIByC) for

lending us their optic microscope. Thanks to D.E. Walter and Q-H. Fan (Ministry for Primary Industries,

Auckland, New Zealand) for their co mm ents on leg chaetotaxy in camerobiid mites. Thanks to J.A.

Ballesteros (George Washington University) and E.A. Ueckermann (Agricultural Research Council,

South Africa) for providing us with literature on camerobiid mites. Thanks to D. Ortiz and F. Chiang

(Instituto de Biologia, UNAM) for corroborating the correct formation of specific epithets in Latin.

Thanks to Ian Gardner (The seven seas translations agency) for improving the English language. We

are grateful to the editorial board of European Journal of Taxonomy , Koen Martens (Editor in Chief)

and two anonymous reviewers for useful comments which improved this manuscript. This work was

supported by the “Programa para el Desarrollo Profesional Docente para el tipo Superior-Secretaria de

Educacion Publica (PRODEP-SEP)” through a post-doctoral fellowship to the first author under grant

PRODEP/103.5/13/9465; and “Red Tematica de Colaboracion de Cuerpos Academicos: Sistematica y

ecologia en comunidades forestales y cultivos” under grant PRODEP/103.5/09/1187, Folio 13.

PAREDES-LEON R. et al.. New species of camerobiid mites and setal homology in Neophyllobius

References

Akyol M. & K 09 K. 2006. The camerobiid mites (Acari, Camerobiidae) of Turkey. Biologia, Bratislava

61 (2): 125-132. http://dx.doi.org/10.2478/sll756-006-002Q-l

Akyol M. 2013. Two new species of the genus Neophyllobius Berlese (Acari: Camerobiidae) from Turkey.

International journal ofAcarology 39 (7): 542-546. http://dx.doi.org/10.1080/01647954.2Q13.839741

Arizaga S., Martinez-Cruz J., Salcedo-Cabrales M. & Bello-Gonzalez M.A. 2009. Manual de la

biodiversidad de encinos michoacanos. Secretaria de Medio Ambiente y Recursos Naturales and

Instituto Nacional de Ecologia, Mexico City.

Bochkov A. V., OConnor B.M. & Wauthy G. 2008. Phylogenetic position of the mite family Myobiidae

within the infraorder Eleutherengona (Acariformes) and origins of parasitism in eleutherengone mites.

Zoologischer Anzeiger 247 (1): 15-45. http://dx.doi.Org/10.1016/i.icz.2006.12.003

Bolland H.R. 1983. A description of Neophyllobius aesculi n. sp. and its developmental stages (Acari:

Camerobiidae). Entomologische Berichten 43 (3): 42M7.

Bolland H.R. 1986. Review of the systematics of the family Camerobiidae (Acari, Raphignathoidea). I.

The genera Camerobia , Decaphyllobius, Tillandsobius , and Tycherobius. Tijdschrift Voor Entomologie

129: 191-215.

Bolland H.R. 1991. Review of the systematic of the family Camerobiidae, II. The genus Neophyllobius

Berlese, 1886 (Acari: Raphignathoidea). Genus 2: 59-226.

Bolland H.R. & Swift S.F. 2000. Hawaiian Raphignathoidea: Family Camerobiidae (Acariformes:

Prostigmata), with descriptions of three new species. International Journal of Acarology 26 (4): 347-

356. http://dx.doi.org/10.1080/016479500086842Q9

Bolland H.R. 2001. Observations and description of Neophyllobius piniphilus n. sp. (Acari: Camerobiidae)

from pine trees in The Netherlands. International Journal of Acarology 27 (2): 129-134. http://dx.doi.

org/10.1080/01647950108684241

Cortes-Anzures B.O. 2015. Distribucion de eplfitas entre forofitos de un bosque tropical caducifolio

sobre suelo volcanico de Tepoztldn, Morelos, asi como el efecto de Quercus obtusata y Sapium

macrocarpum sobre la germinacion de Encyclia spatella y Guarianthe aurantiaca. Master’s thesis,

Universidad Autonoma del Estado de Morelos, Cuernavaca.

De Leon D. 1958. The genus Neophyllobius in Mexico (Acarina: Neophyllobiidae). The Florida

Entomologist 41 (4): 173-181. http://dx.doi.org/10.2307/3492958

De Leon D. 1967. Some Mites of the Caribbean Area. Part I. Acarina on Plants in Trinidad, West Indies.

Allen Press Inc., Lawrence.

Espejo-Serna A., Lopez-Ferrari A.R., Ramirez-Morillo I., Holst B.K., Luther H E. & Till W. 2004.

Checklist of Mexican Bromeliaceae with notes on species distribution and levels of endemism. Selbyana

25: 33-86.

Fan Q.-H. & Walter D.E. 2011. Acamerobia inflatus gen. n. & sp. n. from Australia (Acari: Prostigmata:

Raphignathoidea: Camerobiidae) with notes on the idiosomal chaetotaxy. Zootaxa 3045: 45-56.

Gerson U., Smiley R.L. & Ochoa R. 2003. Mites (Acari) for pest control. Wiley-Blackwell, London.

Grandjean F. 1941. La chaetotaxie comparee des pattes chez les oribates (Ire serie). Bulletin de la

Societe Zoologique de France 66 : 33-50.

Grandjean F. 1944. Observations sur les Acariens de la famille des Stigmaeidae. Archives des Sciences

physiques et naturelles 26: 103-131.

European Journal of Taxonomy 202 : 1-25 ( 2016 )

Grandjean F. 1946. Au sujet de l’organe de Claparede, des eupathides multiples et des taenidies

mandibulaires chez les Acariens actinochitineux. Archives des Sciences physiques et naturelles 28: 63-87.

Hoffmann A. & Lopez-Campos G. 2000. Biodiversidad de los acaros en Mexico. Comision Nacional para

el Conocimiento y Uso de la Biodiversidad and Universidad Nacional Autonoma de Mexico, Mexico City.

Kethley J. 1990. Acarina: Prostigmata (Actinedida). In: Dindal D.L. (ed.) Soil biology guide: 667-756.

Wiley, New York.

Khanjani M. & Ueckermann E.A. 2006. A new species of the genus Neophyllobius Berlese (Acari:

Camerobiidae) from Iran. International Journal of Acarology 32 (3): 277-281. http://dx.doi.

org/10.1080/01647950608684468

Khanjani M., Fayaz B.A. & Ghanbalani G.N. 2010. Two new species of the genus Neophyllobius Berlese

(Acari: Camerobiidae) from Iran. Zootaxa 2521: 53-64.

Khanjani M., Asadabadi A., Izadi H. & Dogan S. 2012a. A new species of Neophyllobius (Acari:

Raphignathoidea, Camerobiidae) from southeast Iran. Systematic and Applied Acarology 17(1): 67-73.

http://dx.doi.Org/10.11158/saa.17.l.12

Khanjani M., Yazyanpanah S., Ostovan H. & Fayaz B.A. 2012b. Three new species of the genus

Tycherobius Bolland (Acari: Camerobiidae) from Iran. Zootaxa 3266: 23M0.

Khanjani M., Hajizadeh J., Hoseini M.A. & Jalili M. 2013. Two new species of the genus Tycherobius

Bolland (Acari: Camerobiidae) from north of Iran. International Journal of Acarology 39 (2): 130-139.

http://dx.doi.org/10.1080/01647954.2012.7524Q5

Khanjani M. & Hoseini M.A. 2013. Three new species of the genus Neophyllobius Berlese (Acari:

Camerobiidae) from southern and southwestern Iran. Zootaxa 3666: 510-522. http://dx.doi.org/10.11646/

zootaxa.3666.4.6

Khanjani M., Hoseini M.A., Yazdanpanah S. & Masoudian F. 2014. Neophyllobius lorestanicus sp.

nov. and N. ostovani sp. nov. (Acari: Camerobiidae) from Iran. Zootaxa 3764: 441-454. http://dx.doi.

org/10.11646/zootaxa.3764.4.4

K 09 K. & Ayyildiz N. 1996. Anew species of Neophyllobius Berlese (Acari: Camerobiidae) from Turkey.

International Journal of Acarology 22 (4): 291-294. http://dx.doi.org/10.1080/016479596086841Q7

K 09 K. & Akyol M. 2007. Morphometric & intraspecific variations between specimens of Tycherobius

stramenticola and Tycherobius polonicus (Acari: Camerobiidae) from Turkey. Acarologia 47 (3—4):

139-142.

McGregor E.A. 1950. Mites of the genus Neophyllobius. Bulletin of the Southern California Academy

of Sciences 49 (2): 55-70.

Norton R.A. 1977. A review of F. Grandjean’s system of leg chaetotaxy in the Oribatei and its

application to the Damaeidae. In: Dindal D.L. (ed.) Biology of oribatid mites : 33-62. SUNY College of

Environmental Science and Forestry, New York.

Palacios-Vargas J.G. 1982. Microartropodos asociados a bromeliaceas. In: Salinas P.J. (ed.) Zoologia

Neotropical Adas del VIII Congreso Latinoamericano de Zoologia. Vol. 1: 535-545. Consejo Nacional

de Investigaciones Cientificas y Tecnologicas, Merida.

Paredes-Leon R. & Guzman-Cornejo C. 2015. Anew species of pterygosomatid mite and its phylogenetic

position within the genus Geckobiella (Acariformes: Prostigmata: Pterygosomatidae). International

Journal of Acarology 41 (1): 19-30. http://dx.doi.org/10.1080/01647954.2Q14.988642

PAREDES-LEON R. et al.. New species of camerobiid mites and setal homology in Neophyllobius

Paredes-Leon R., Klompen H. & Perez T.M. 2012. Systematic revision of the genera Geckobiella Hirst,

1917 and Hirstiella Berlese, 1920 (Acari: Prostigmata: Pterygosomatidae) with description of a new

genus for American species parasites on geckos formerly placed in Hirstiella. Zootaxa 3510: 1—40.

Ruiz-Cordova J.P, Toledo-Hernandez V.H. & Flores-Palacios A. 2014. The effect of substrate abundance

in the vertical stratification of bromeliad epiphytes in a tropical dry forest (Mexico). Flora 209 ( 8 ):

375-384. http://dx.doi.Org/10.1016/i.flora.2014.06.003

Swift S.F. 2001. The leg chaetotaxy of Caligonellidae (Prostigmata: Raphignathoidea). In: Halliday

R.B., Walter D.E., Proctor H.C. & Colloff M.J. (eds) Acarology: Proceedings of the 10 lh International

Congress : 242-249. CSIRO Publishing, Melbourne.

The GIMP team. 2014. GIMP 2.8.14. Available from http://www.gimp.org [accessed 9 Aug. 2015]

Ulu 9 ay I. & K 09 K. 2014. A new species of Neophyllobius and description of male of Neophyllobius

yunusi (Acari: Camerobiidae) from Turkey. International Journal of Acarology 40 (1): 15-22. http://

dx.doi.org/10.1080/01647954.2013.870226

Vazquez M.M., Pereira A., Fragoso P. & Rodriguez J.A. 2011. Riqueza y biodiversidad de organismos

edaficos del Corredor Biologico Mesoamericano (Mexico, Belice y Guatemala). In: Delgado Bias V.H.,

Ortegon Aguilar J.S., Vazquez Gonzalez M.M., Gonzalez Damian A. & Hernandez-Rodriguez J. (eds)

Avances de cienciay tecnologia en Quintana Roo: 37-70. Plaza y Valdes, Mexico City.

Vazquez-Gonzalez M.M. 2008. Microartropodos edaficos litorales. Dugesiana 15: 7-15.

Vergara-Torres C.A., Pacheco-Alvarez M.C. & Flores-Palacios A. 2010. Host preference and host

limitation of vascular epiphytes in a tropical dry forest of central Mexico. Journal of Tropical Ecology

26 ( 6 ): 563-570. http://dx.doi.org/10.1017/S026646741000Q349

Walter D.E., Findquist E.E., Smith I.M., Cook D.R. & Krantz G.W. 2009. Order Trombidiformes. In:

Krantz G.W. & Walter D.E. (eds) A manual of Acarology, 3 rd edition : 233M20. Texas Tech University

Press, Texas.

Zeity M. & Gowda C.C. 2013. A new species of Neophyllobius (Acari: Camerobiidae) from India.

International Journal of Acarology 39 (7): 547-550. http://dx.doi.org/10.1080/01647954.2Q13.844726

Zhang Z.-Q., Fan Q.-H., Pesic V., Smit H., Bochkov A.V., Khaustov A.A., Baker A., Wohltmann A.,

Wen T., Amrine J.W., Beron P, Fin J., Gabrys G. & Husband R. 2011. Order Trombidiformes Reuter,

1909. In: Zhang Z.-Q. (ed.) Animal biodiversity: An outline of higher-level classification and survey of

taxonomic richness. Zootaxa 3148: 129-138.

Manuscript received: 16 October 2015

Manuscript accepted: 27 January 2016

Published on: 2 June 2016

Topic editor: Koen Martens

Desk editor: Charlotte Thionois

Printed versions of all papers are also deposited in the libraries of the institutes that are members of the

EJT consortium: Museum national d’Histoire naturelle, Paris, France; Botanic Garden Meise, Belgium;

Royal Museum for Central Africa, Tervuren, Belgium; Natural History Museum, Fondon, United

Kingdom; Royal Belgian Institute of Natural Sciences, Brussels, Belgium; Natural History Museum of

Denmark, Copenhagen, De nm ark