External morphology and systematics of the family Lobalgidae (Acari: Astigmata) with resolution of...

© Koninklijke Brill NV, Leiden, 2010 DOI 10.1163/187631210X496831

Insect Systematics & Evolution 41 (2010) 187–229 brill.nl/ise

External morphology and systematics of the family Lobalgidae (Acari: Astigmata) with resolution

of the question of family monophyly

Andre V. Bochkov a,* and Barry M. OConnor b a Zoological Institute, Russian Academy of Sciences, Universitetskaya

Embankment 1, St. Petersburg 199034, Russia b Museum of Zoology, University of Michigan, 1109 Geddes Avenue, Ann Arbor, MI 48109, USA

*Corresponding author, e-mail: [email protected]

Abstract Th e monophyly of the family Lobalgidae comprising the genera Lobalges , Echimytricalges and Coendalges is tested using morphological characters and the maximum parsimony approach. Representatives of all subfamilies of the family Psoroptidae, the families Pyroglyphidae ( Paralgopsis ) and Psoroptoididae ( Psoroptoides ), and most families of the superfamily Sarcoptoidea were used as close outgroups, with Ptiloxenus (Ptiloxenidae) and Acarus (Acaridae) selected as distant outgroups. Th e analysis revealed polyphyly of the family: the two lobalgid genera Lobalges and Echimytricalges appear as sister groups, whereas the genus Coendalges is a member of a clade comprising genera of the psoroptid subfamily Psoralginae. Th e composition of the family Lobalgidae is here limited to these two genera, and the genus Coendalges is placed in the subfamily Psoralginae (Psoroptidae). A taxonomic revision of the family Lobalgidae is given, including descriptions of two new species, Echimytricalges lonchothrix sp.n. and E. proechimys sp.n. We also validate the species Echimytricalges whitakeri Fain and Ritzi, which had previ-ously been a nomen nudum .

Keywords Phylogeny , Sarcoptoidea , parasites

Introduction

Th e family Lobalgidae Fain, 1965 (Acari: Astigmata) includes permanent, highly specialized symbionts of neotropical edentates and rodents (Fain 1965 ; Fain & Mendez 1979 ; OConnor 1982 ). To date, this relatively small family consists of eight species in three genera belonging respectively to three monobasic subfamilies, Lobalginae (1 species from sloths), Echimytricalginae Fain, 1970 (6 species from spiny rats of the family Echimyidae) and Coendalginae Fain and Mendez, 1979 (1 species from porcu-pines of the family Erethizontidae). Mites of the two monobasic subfamilies live on the host skin, whereas species of the subfamily Echimytricalginae inhabit the concave

188 A.V. Bochkov and B.M. OConnor / Insect Systematics & Evolution 41 (2010) 187–229

surfaces of the widened, spine-like hairs of certain Echimyidae (Fain & Lukoschus 1970 ; OConnor 1982 ).

Th e monophyly of Lobalgidae has recently been questioned (OConnor 2009 ) due to the strong morphological and host distinctions between its subfamilies, but no phy-logenetic analyses have been conducted to date. Th e external morphology of these mites has also not been adequately described, and the developmental homology of leg and idiosomal setae has never been established.

In the present study, we describe the external morphology of lobalgid mites in detail and establish homologies of the leg and idiosomal setae with those of the other Astigmata for the fi rst time. In order to document the synapomorphies supporting the monophyly of the family Lobalgidae, we compared these mites with representatives of most subfamilies belonging to the diverse, and possibly artifi cial, family Psoroptidae, which is considered to be closely related to Lobalgidae (OConnor 1982 , 2009 ), and conducted a phylogenetic analysis using maximum parsimony. We also provide a taxo-nomic revision of the Lobalgidae, including descriptions of two new species, making a third species that was a nomen nudum available, and present keys to all known species. We also provide emended familial and subfamilial diagnoses. Finally, we update the nomenclature of the hosts of these mites to refl ect recent revisions and taxonomic changes.

Historical remarks

Th e fi rst member of the family Lobalgidae, Lobalges trouessarti Fonseca, 1954 was described from Bradypus variegatus Schinz, 1825 ssp. brasiliensis Blainville, 1840 (= B. tridactylus brasiliensis) (Pilosa: Bradypodidae) in Brazil (Fonseca 1954 ). Fonseca ( 1954 ) established for this species the monobasic genus Lobalges and placed it in the family Epidermoptidae, which included at that time a heterogenous assemblage of mites associated with bird skin and also some genera with unclear phylogenetic links, such as Yunkeracarus, parasites inhabiting the nasal cavities of mammals, and Dermatophagoides, bird nest inhabitants , now placed in the families Gastronyssidae and Pyroglyphidae, respectively. Fain ( 1965 ) re-described L. trouessarti and created the monobasic family Lobalgidae noting several important features of the genus Lobalges that separated it from members of the families Epidermoptidae and Psoroptidae: coxal apodemes I fused into a sternum, strongly shortened coxal apodemes III, presence of idiosomal lobes in both sexes, and absence of tibial setae kT III.

Later, Fain ( 1970 ) described the second genus of the family, Echimytricalges Fain, 1970 , with two new species from other South American hosts, E. brasiliensis Fain, 1970 from Phyllomys pattoni Emmons et al., 2002 (= Echimys brasiliensis sensu Fain, 1970 ) (Rodentia: Echimyidae) and E. guyanensis Fain, 1970 from Caluromys philander (L., 1758) (Didelphomorphia: Didelphidae). Mites of this genus clearly diff er from L. trouessarti by idiosomal and leg setation, vulvar structure, hysteronotal sclerotization in females, and by the structure of legs IV in males (see generic key), and, therefore, a new monobasic subfamily Echimytricalginae was created for them (Fain 1970 ).

A.V. Bochkov and B.M. OConnor / Insect Systematics & Evolution 41 (2010) 187–229 189

To date, Fain and co-authors (Fain & Lukoschus 1970 ; Fain et al. 1982 ; Fain & Ritzi 2001 ) have added four more species described from diff erent spiny rats (see Table 6).

Th e third subfamily Coendalginae was established for the monotypic genus Coendalges Fain and Mendez, 1979 (Fain & Mendez 1979 ). Th e single species of this subfamily, C. panamensis Fain and Mendez, 1979 associated with Coendou rothschildi Th omas, 1902 (Rodentia: Erethizontidae) from Panama, diff ers from all the other lobalgids mainly by the broadly oval idiosoma, the presence of setae vi , widely sepa-rated coxal apodemes I, the presence of distinctly developed dorso-apical projections on tarsi I-II, and by the elongated tarsi III–IV in females.

Materials and methods

Material Most lobalgid specimens examined in this study were collected by one of us (B.M.OC.) from dried or fl uid preserved host specimens in museum collections. Type series of Coendalges panamensis was loaned from the Institut royal des Sciences naturelles de Belgique, Brussels, Belgium. Comparative material for the phylogenetic analysis is deposited in three institutions: Institut royal des Sciences naturelles de Belgique, Brussels, Belgium; Museum of Zoology, University of Michigan, Ann Arbor, MI, USA; Zoological Institute of the Russian Academy of Sciences, Saint-Petersburg, Russia. List of all outgroup species used in the phylogenetic analysis is given in Appendix A.

Mites were cleared in lactophenol and mounted in Hoyer’s medium (Krantz 1978 ). Drawings were made with a Zeiss microscope with a camera lucida using phase-contrast optics. Specimens were also studied using a Leica DMLB microscope equipped with diff erential interference contrast optics.

In the descriptions below, the idiosomal chaetotaxy follows Griffi ths et al. ( 1990 ) with modifi cations of Norton ( 1998 ) for coxal-genital setae. Th e leg chaetotaxy follows Grandjean ( 1939 ). All measurements are given in micrometers and were taken as fol-lows: body length=the total length from the anterior extremity of the prescapular shield to the posterior border of the body; body width=maximum width taken at whatever level it occurs; length of dorsal shields=maximum length, measured in the median line of the shields; length of the posterior legs=length from the most basal point of the trochanter to the apex of the tarsus, excluding pretarsal ambulacrum. In the collection records, names of hosts follow Wilson & Reeder ( 2005 ). All lobalgid records are sum-marized in Table 6.

Specimen depositories and reference numbers are cited using the following abbre-viations: BMOC, B.M. OConnor reference number; BMNH, Th e Natural History Museum, London, UK; FMNH, Field Museum of Natural History, Chicago, IL, USA; IRSNB, Institut royal des Sciences naturelles de Belgique, Brussels, Belgium; LPB, Laboratorio de Parasitologia do Instituto Butantan, Sao Paulo, Brazil; MCZ, Museum of Comparative Zoology, Harvard University, Cambridge, MA, USA; MUSM, Museo de Historia Natural de la Universidad Nacional Mayor de San Marcos, Lima, Peru; MVZ, Museum of Vertebrate Zoology, University of California, Berkeley, CA, USA; RMNH, Naturalis, Nationaal Natuurhistorisch Museum, Leiden, the Netherlands;


TTU, Th e Museum of Texas Tech University, Lubbock, TX, USA; UMMZ, Museum of Zoology, University of Michigan, Ann Arbor, MI, USA; USNM, National Museum of Natural History, Smithsonian Institution, Washington, DC, USA.

Phylogenetic analysis

Th e family Lobalgidae has been considered to be closely related to the possibly non-monophyletic family Psoroptidae, which includes seven subfamilies (OConnor 1982 , 1984 ). Among “higher” Psoroptidia (see Mironov & Bochkov 2009 ) we selected for analyses nine genera belonging to all psoroptid subfamilies in order to test lobalgid monophyly (see Table 5). Additionally, representatives of most other families of the superfamily Sarcoptoidea were included in this analysis (see Table 5). We did not include representatives of the family Chirorhynchobiidae and endoparasitic mites (Gastronyssidae, Pneumocoptidae, and Lemurnyssidae), whose external structures are highly reduced or lost. Th eir inclusion in the analysis leads many obviously false homologies. Th e families Rhyncoptidae and Sarcoptidae are sister (Klompen 1993 ) and, therefore, we selected for the analysis only representatives of the most archaic genus Caenolestocoptes (Rhyncoptidae). Th e monophyly of the family Myocoptidae is questionable (Bochkov & OConnor 2008 ), for this reason representatives of both sub-families, Myocoptinae and Dromiciocoptinae were selected. Two analgoid genera Paralgopsis (Analgoidea: Pyroglyphidae) and Psoroptoides (Psoroptoididae) were also selected as close outgroups. One pterolichoid genus belonging to “low” Psoroptidia and one free-living astigmatic genus, Ptiloxenus (Pterolichoidea: Ptiloxenidae) and Acarus (Acaroidea: Acaridae), respectively were chosen as distant outgroups.

Only qualitative characters, such as presence/absence of a morphological structure or the form of some morphological elements were selected. Th e autapomophies of the lobalgid taxa were also included for taxonomic-diagnostic purposes. In total, 23 taxa, 42 phylogenetically informative and 9 autapomorphies were included to the data matrix (Table 5). Preparing and editing of the data matrix were done using NEXUS Data Editor 0.5.0 (Page 2001 ). All characters were unordered and initially unweighted. Th e list of characters is given in Appendix B. Maximum parsimony analysis was used for testing lobalgid monophyly.

Th e reconstruction of phylogenetic relationships, analysis of character distributions, drawing, and editing of the trees were performed with PAUP 4.0b.10 for Windows (Swoff ord 2001 ) and WinClada 0.9.99m24 (Nixon 2000 ). Th e heuristic search option was used with random addition of taxa, 10 000 replications, and TBR branch swap-ping. Support for branches was estimated by Bremer support indices calculated with the program TreeRot for Macintosh (Sorenson 1999 ).

Th e external morphology of the family Lobalgidae

Gnathosoma (Figs 4C, 14A, B) Th e gnathosoma of lobalgid mites has the general structure typical for the Psoroptidia and is devoid of ventral apophyses. In Echimytricalginae, the subcapitular base bears a


pair of large dorsal lobes. Th e palps have two podomeres, tibia and tarsus (Norton 1998 ). Th e basal podomere or tibia bears two setae, d and l” , tarsus bears a dorsal seta, d, one solenidion ( ω ), two eupathidia ( u’ and u” ), and is covered dorsally by a membrane (OConnor 1981 ). Th e fi xed digit of the chelicera has four distinct teeth; the teeth of the movable digit are weakly discernible. Most cheliceral structures, such as the cheli-ceral paraxial hood, the cheliceral spur, and the seta-like apophysis are present, and only the antiaxial apophysis is indiscernible because of the strong sclerotization of this part of the chelicerae, but is possibly present as well. A pair of subcapitular setae ( subc ) and a pair of supracoxal setae of the palps ( elc. p .) are present.

Idiosoma (Figs 3-6, 9, 10)

In Lobalginae and Echimytricalginae , the idiosoma is dorso-ventrally fl attened and more or less elongated, whereas in Coendalginae, the idiosoma is broadly oval. In all representatives of the family, the sejugal furrow is not observable. In both sexes of Lobalginae and Coendalginae, as well as in males of Echimytricalginae, the dorsal sur-face of the idiosoma is covered by two shields, the propodonotal shield and the hyster-onotal shield. In females of Echimytricalginae, the hysteronotal shield is subdivided onto an unpaired central hysteronotal shield and paired opisthonotal shields. Th e openings of the supracoxal glands connecting with the podocephalic canals, and supraxocal setae of legs I, scx, are distinctly developed. Openings of the opisthonotal glands are situated dorso-laterally, anterior to the level of setae e1 .

Th e legs of lobalgids are inserted laterally, and the coxal fi elds are striated. In Lobalginae, coxal apodemes I are fused into a sternum; in Echimytricalginae, they are connected to each other by a strongly sclerotized bridge, but their posterior ends are free. In Coendalginae, coxal apodemes I are free and situated far from each other. Apodemes II are distinctly developed in all lobalgids. Apodemes III are distinctly devel-oped in Echimytricalginae and Coendalginae, whereas in Lobalginae, these apodemes are very short. In all lobalgids apodemes IV are moderately developed.

As in most ectoparasitic sarcoptoid mites, the posterior end of the lobalgid male idi-osoma bears a pair of terminal lobes bordered by lobar membranes.

In females of Coendalges, the posterior margin of the opisthosoma has a distinct median cleft (Fig. 6). In females of the subfamilies Lobalginae and Echimytricalginae, possessing the “true” opisthosomal lobes, these lobes are fl attened, distinctly sclero-tized, delineated from the remaining part of the opisthosomal surface, and widely sepa-rated medially from each other occupying the lateral margins of the opisthosoma. Moreover, the posterior margin of the opisthosoma between the lateral lobes bears an interlobar membrane (=median lobe) connecting these lobes. Fain & Mendez ( 1979 , p. 271) considered the posterior median cleft of the female opisthosoma of C. pana-mensis as an incision separating the opisthosomal lobes. In our opinion, the “true” opisthosomal lobes are absent in females of this species because its opisthosomal end is not diff erentiated from the anterior part of the opisthosoma, is covered by the same striations as the remaining part of the idiosomal cuticle, and is devoid of membranous structures.


Th e idiosomal setation diff ers slightly among three lobalgid subfamilies. Th e sub-family Coendalginae exhibits the most complete idiosomal setation: vi, si, se, c2-c3, cp, d1-2, e1-2, h2, 3, ps1-3, 1a, 3a, 4a, 4b , g, and the distinctly developed paranal suckers in males. In species of the other two subfamilies, setae vi are absent, and, in addition, in Lobalginae, setae c3 are absent, but setae c1 are present. All setae, as a rule, are seti-form and sometimes thickened. In both sexes of Echimytricalginae, setae cp are sword-like, and in males setae h3 bear membranous enlargements. Th e idiosomal setation of lobalgids is summarized in Table 1. Th e idiosomal setation of psoroptid genera included in the analyses is also included in Table 1.

Th e female copulatory system is similar to that described for Listrophoridae (Bochkov & OConnor 2006 ): the opening of the female bursa copulatrix is situated terminally and connected to the sacculiform spermatheca via the inseminatory canal; at the junction with the inseminatory canal, the spermatheca forms a globose, sclero-tised structure termed the basal cap; the paired eff erent sperm ducts are sclerotized for a short distance extending from the basal cap (Fig. 14F).

Th e ovipore is situated ventrally, between coxal fi elds III and IV in Lobalginae and Echimytricalginae, and between coxal fi elds II and III in Coendalginae. Th e epigynum is distinctly developed, arch-like. In Echimytricalginae, the epigynum is fused with apodemes III; in Coendalginae it is situated between apodemes I and II. In all lobal-gids, the undulate lamina of the ovipore is covered laterally by a pair of the progenital folds; in Lobalginae, these folds are almost indistinct. Two pairs of genital papillae fl anking the ovipore are situated near progenital folds.

Th e male genital organ (Figs 5C, 13D) is situated between the levels of coxal fi elds III and IV. It is covered by the genital valves. Two pairs of genital papillae fl ank the genital organ and are covered by the genital valves. Th e basal part of the aedeagus is enclosed within the genital capsule.

Legs (Figs 3C-G, 4D-E, 5D, E, 6C-I, 9C, D, 13A-C, E-J)

Th e legs of lobalgid females are relatively weakly modifi ed and devoid of retrorse spurs. Th ey include the full set of articulating segments plus a pretarsus consisting of an ambulacral stalk and ambulacral disc. In Lobalginae and Echimytricalginae, tarsi I and II each have an indistinct dorso-apical projection (Fig. 13E), whereas in Coendalginae, these processes are distinctly developed and curved. In males of Echimytricalginae, legs IV are strongly reduced and their femora and genua are completely fused (Figs 9D, 13C). In males of Coendalginae, legs III are hypertrophied; legs IV, consisting of fi ve articulated segments, are reduced in size, but not so strongly as those in echimytrical-gines. Tarsi III of echimytricalgine and coendalgine males bear a small ventral projec-tion situated in the apical part of the tarsus (Fig. 13B); tarsi IV of lobalgine males bear a short dorso-apical projection.

Th e structure of the lobalgid pretarsi corresponds well to those of the Listrophoridae (Wurst 1993 , Bochkov & OConnor 2006 ) and has the following structures described using the terminology of Atyeo ( 1978 ). Th e ambulacral disc is attached to the ambu-lacral stalk. Th e condylophores within the ambulacral stalk are straight. Th e ambulacral


Tabl

e 1.

Idi

osom

al se

tatio

n of

Lob

algi

dae

(L) a

nd se

lect

ed P

soro

ptid

ae (P

)

Taxo

n scx

vi

Si

se

c1

c2

cp

c3

d1

d2

e1

e2

f2

h1

H3

h2

g 1a

3a

4a

4b

ps

1 ps

2 ps

3

Proc

ebal

ges (

P)+

++

+–

++

++

++

+–/

+–

++

++

++

++

++

Mak

ialg

es (P

)–

++

++

++

++

++

++

–+

++

++

++

++

+ M

arsu

pial

ges (

P)–

–+

+–

++

++

++

++

–+

++

++

++

++

+ N

asal

ialg

es (P

)–

++

++

++

++

++

++

–+

++

++

++

++

+ Pa

ngor

illal

ges (

P)–

++

++

++

++

++

++

–+

++

++

++

++

+ Li

strop

sora

lges

(P)

–+

++

–+

++

–+

++

++

++

++

++

++

++

Psor

alge

s (P)

++

++

–+

++

++

++

+–

++

++

++

++

++

Eden

talg

es (P

)+

++

+–

++

++

++

++

–+

++

++

++

++

+ Tr

ouess

alge

s (P)

+–

++

++

++

++

++

+–

++

++

++

++

++

Hyr

acop

tes (

P)+

–+

++

++

++

++

++

–+

++

++

++

++

+ Ec

him

ytric

alge

s (L)

+–

++

–+

++

++

++

+–

++

++

++

++

++

Loba

lges

(L)

+–

++

++

+–

++

++

+–

++

++

++

++

++

Coe

ndal

ges (

L)+

++

+–

++

++

++

++

–+

++

++

++

++

+

–, a

bsen

t; +,

pre

sent

; /, f

emal

e/m

ale.


discs are fl attened dorso-ventrally. Th e unguiform sclerites are situated posterior to a massive central sclerite represented by the basilar piece and modifi ed empodial claw fl anking the condylophore guide. In Coendalginae a pair of strongly reduced lateral sclerites fl anks the basilar piece; empodial claws are poorly developed and are either reduced or completely fused with the basilar piece.

Th e leg setation of Lobalgidae slightly diff ers in all three subfamilies. In Echimytri-calginae, legs I-IV bear the following setae, solenidia and famulus: pR I-II, sR III, vF I- II, cG I-II , mG I-II , gT I-II , kT III-IV, σ I-II, φ I-IV, wa I-II , ra I-II , w III-IV , r III-IV , la I-II , s I-III , d I-IV , e I-IV , f I-IV , ε I , ω1 I-II, and ω3 I. All setae are setiform, sometimes thickened. In males, setae d and e IV are modifi ed into suckers. In Coendalginae, setae ba I–II are present. In Lobalginae, solenidia σ III are present, setae e III, s III, and kT IV are absent, in males, setae d IV and e IV are fi liform. Th e leg setation of lobalgids and psoroptids included in the analysis is given in Tables 2–4.

Immature instars (Figs 7, 8)

Only adults and tritonymphs of L. trouessarti and C. panamensis were available to us, whereas among Echimytricalginae, we studied all life stages in Echimytricalges hylomys Fain, Lukoschus and Mendez, 1982 and separate immature stages in several other spe-cies of the genus. In larvae of Echimytricalginae, the gnathosoma is as in adults, and the idiosoma is as in females, but the hysteronotal shield is entire (Fig. 7). Anterior apodemes I are not fused to each other. Th e idiosoma bears the following setae: scx, si, se, c2, c3, cp, d1, d2, e1, e2, h2, 1a, and 3a . Th e legs are similar to those in females, and their setation is typical for sarcoptoid larvae: I trochanter 0, femur 1 ( vF ), genu 3 ( cG, mG, σ1 ), tibia 2 ( gT, φ ), tarsus 9 ( wa, ra, la, s, d, e, f, ε, ω1 ); ΙΙ trochanter 0, femur 1 ( vF ), genu 3 ( cG, mG, σ ), tibia 2 ( gT, φ ), tarsus 8 ( wa, ra, la, s, d, e, f, ω1 ); ΙΙΙ trochanter 0, femur 0, genu 0, tibia 2 ( kT, φ ), tarsus 6 ( w, r, s, d, e, f ). In the prot-onymph, setae f 2, h3, ps1-3, and g are added, and one pair of genital papillae is present between coxae IV (Fig. 8A-C). Apodemes I are fused into a sternum. Th e tarsal setae d, r , and w are added on legs IV. In the tritonymph, setae 4a and 4b are added on the idiosoma (Fig. 8D-E), setae pR I-II and sR III are added on trochanters I-III, setae kT and solenidion φ are added on tibia IV, and setae e, f are added on tarsus IV. No diff er-ences between pharate tritonymphs containing males and females were observed. Setae h1 are absent not only in all lobalgids, but also in pyroglyphids and most psoroptids.

Pre-copulatory mate-guarding behavior

Physical attachment between males and nymphs is widely observed among psoroptid-ian mites and has been termed pre-copulatory mate guarding behavior (Witalinski et al. 1992 ). In most psoroptidians, males are positioned above the posterior part of the nymphal idiosoma, and the long axes of the bodies of the partners are orientated in opposite directions. Our results based on examination of dead specimens. Among Lobalgidae, the same position has been observed in Echimytricalges spp. and C. pana-mensis , whereas males of L. trouessarti , as in the case with Hemigalichus chrotogale Bochkov and OConnor, 2006 (Listrophoridae), are positioned above the anterior part


Tabl

e 2.

Leg

I se

tatio

n of

Lob

algi

dae

(L) a

nd se

lect

ed P

soro

ptid

ae (P

)

Taxo

n d

ba

e f

s la

ω1

ω3

σ1

σ2

ε

φ w

r

gT

mG

cG

vF

pR

Proc

ebal

ges (

P)F

–F

FF

F+

++

–+

+F

FF

FF

FF

Mak

ialg

es (P

)F

CF

FF

F+

++

–+

+F

FF

FF

FF

Mar

supi

alge

s (P)

F–

FF

–F

++

+–

++

FF

FF

FF

F N

asal

ialg

es (P

)F

FF

F–

–+

++

–+

++

++

++

++

Pang

orill

alge

s (P)

F–

FF

CF

++

+–

++

FF

FF

FF

F Li

strop

sora

lges

(P)

F–

FF

+–

++

+–

++

FF

FF

FF

F Ps

oral

ges (

P)F

FF

FF

F+

++

–+

+F

FF

FF

FF

Eden

talg

es (P

)F

FF

FF

F+

++

–+

+F

FF

FF

FF

Trou

essal

ges (

P)F

FF

FF

F+

++

–+

+F

FF

FF

FF

Hyr

acop

tes (

P)F

FF

FF

F+

++

++

+F

FF

FF

FF

Echi

myt

rical

ges (

L)F

–F

FF

F+

++

–+

+F

FF

FF

FF

Loba

lges

(L)

F–

FF

FF

++

+–

++

FF

FF

FF

F C

oend

alge

s (L)

FF

FF

FF

++

+–

++

FF

FF

FF

F

+, p

rese

nt; –

, abs

ent;

C, c

law

like

; F, fi

lifo

rm.


Tabl

e 3.

Leg

II se

tatio

n of

Lo

balg

idae

(L) a

nd se

lect

ed P

soro

ptid

ae (P

)

Taxo

n d

ba

e f

s la

ω 1

Σ φ

w

r gT

m

G

cG

vF

pR

Proc

ebal

ges (

P)F

–F

FF

F+

++

FF

FF

FF

F M

akia

lges

(P)

FC

FF

FF

++

+F

FF

FF

FF

Mar

supi

alge

s (P)

F–

FF

––

++

+F

FF

FF

FF

Nas

alia

lges

(P)

FF

FF

––

++

+F

FF

FF

FF

Pang

orill

alge

s (P)

F–

FF

CF

++

+F

FF

FF

FF

Listr

opso

ralg

es (P

)F

–F

F+

–+

++

FF

FF

FF

F Ps

oral

ges (

P)F

FF

FF

F+

++

FF

FF

FF

F Ed

enta

lges

(P)

FF

FF

FF

++

+F

FF

FF

FF

Trou

essal

ges (

P)F

FF

FF

F+

++

FF

FF

FF

F H

yrac

opte

s (P)

FF

FF

FF

++

+F

FF

FF

FF

Echi

myt

rical

ges (

L)F

–F

FF

F+

++

FF

FF

FF

F Lo

balg

es (L

)F

–F

FF

F+

++

FF

FF

FF

F C

oend

alge

s (L)

F+

FF

FF

++

+F

FF

FF

FF

+, p

rese

nt; –

, abs

ent;

C, c

law

like

; F, fi

lifo

rm.


Tabl

e 4.

Leg

III–

IV se

tatio

n of

Lob

algi

dae

(L) a

nd se

lect

ed P

soro

ptid

ae (P

)

Taxo

n d I

II d I

V e I

II e I

V f I

II f I

V s II

I s IV

sR II

I φ I

II φ I

V σ I

II w

III

w IV

r III

r IV

kT II

I kT

IV

Proc

ebal

ges (

P)F

F/S

FF/

SF

FF

–F

++

–F

FF

FF

F M

akia

lges

(P)

FF/

SF

F/S

FF

F–

F+

++

FF

FF

FF

Mar

supi

alge

s (P)

FF/

SF

F/S

FF

––

F+

+–

FF

FF

FF

Nas

alia

lges

(P)

FF

FF

FF

––

F+

+–

FF

–F

FF

Pang

orill

alge

s (P)

FF/

SF

F/S

FF

F–

F+

++

FF

FF

FF

Listr

opso

ralg

es (P

)F

FF

F/–

FF/

––

–F

++

+F

FF

FF

F Ps

oral

ges (

P)F

F/S

FF/

SF

FF

–F

++

–F

FF

FF

F Ed

enta

lges

(P)

FF/

SF

F/S

FF

–/F

–F

++

–F

FF

FF

F Tr

ouess

alge

s (P)

FF/

SF

F/S

FF

F–

F+

++

FF

FF

FF

Hyr

acop

tes (

P)F

F/S

FF/

SF

F–

–F

++

+F

FF

FF

F Ec

him

ytric

alge

s (L)

FF/

SF

F/S

FF

F–

F+

+–

FF

FF

FF

Loba

lges

(L)

FF

–F

FF

––

F+

++

FF

FF

F–

Coe

ndal

ges (

L)F

F/S

FF/

SF

FF

–F

++

–F

FF

FF

F

+, p

rese

nt; –

, abs

ent;

F, fi

lifor

m; S

, suc

ker l

ike;

/, fe

mal

e/m

ale.

198 A.V. Bochkov and B.M. OConnor / Insect Systematics & Evolution 41 (2010) 187–229 Ta

ble

5. D

ata

mat

rix

Taxa

Cha

ract

er st

ates

1

1

1111

1111

2

22

2222

2223

333

3333

334

4444

4444

45

555

1234

5678

90

1

2345

6789

0

12

3456

7890

123

4567

890

1234

5678

90

123

Acar

us ( A

cari

dae )

0000

–000

00

0

0000

0000

0

00

0000

0000

000

0000

000

00–0

0000

00

000

Ptilo

xenu

s ( Pt

iloxe

nida

e )00

1100

0010

110

0000

000

0100

0001

00

0

0000

0000

0

11

0000

0000

00

0 Ps

orop

toid

es ( P

soro

ptoi

dida

e )00

1100

0010

101

0000

001

0001

1110

00

0

0100

0001

0

11

1000

0000

10

0 Pa

ralg

opsis

( Pyr

ogly

phid

ae )

0010

–000

10

1

0001

0000

1

00

0110

0000

001

0100

010

1110

0100

00

000

Hyr

acop

tes (

Psor

opti

dae :

Pso

ropt

inae

)00

1100

1010

100

1100

001

0001

1010

00

0

0101

0001

0

11

1001

0000

00

0 Tr

ouess

alge

s ( Ps

orop

tida

e : P

soro

ptin

ae)

0011

0010

11

1

0011

0000

1

00

0110

1000

001

0111

010

1111

1101

00

000

Eden

talg

es ( P

soro

ptid

ae : P

sora

lgin

ae)

0010

–010

11

1

0001

0100

1

00

0110

1000

001

0001

110

1111

1101

00

000

Psor

alge

s ( Ps

orop

tida

e : P

sora

lgin

ae)

0010

–010

10

1

0001

0100

1

00

0110

1000

001

0001

110

1110

0101

00

000

Listr

opso

ralg

es ( P

soro

ptid

ae : L

istro

psor

algi

nae)

1011

0110

11

1

0101

0100

0

10

1111

1101

011

0001

010

1111

0001

01

110

Mak

ialg

es ( P

soro

ptid

ae : M

akia

lgin

ae)

1011

0010

11

1

0101

0000

1

00

0110

1000

001

0001

010

1111

0100

10

111

Mar

supi

alge

s ( Ps

orop

tida

e : M

arsu

pial

gina

e)00

10–0

1011

101

1100

001

0001

1112

00

1

1101

0111

0

11

1011

0000

10

0 N

asal

ialg

es ( P

soro

ptid

ae : N

asal

ialg

inae

)00

10–0

1010

101

0100

001

0001

1011

00

1

1101

0111

0

11

1001

0000

00

0 Pa

ngor

illal

ges (

Psor

opti

dae :

Par

acor

optin

ae)

1011

0010

11

1

0101

0000

1

00

0111

1000

001

0011

010

1110

0000

00

011

Proc

ebal

ges (

Psor

opti

dae :

Ceb

algi

nae)

1011

0010

11

1

0001

0100

1

00

0111

1000

001

0111

110

1110

0100

01

000

Coe

ndal

ges (

Loba

lgid

ae : C

oend

algi

nae)

0011

0010

11

1

0001

0100

1

00

0110

1000

001

0001

110

1110

0110

00

000

Echi

myt

rical

ges (

Loba

lgid

ae : E

chim

ytric

algi

nae)

0111

1001

10

1

1011

1110

1

01

0111

0000

001

0001

111

1110

0210

00

000

Loba

lges

( Lob

algi

dae :

Lob

algi

nae)

0011

0000

10

1

1011

1001

1

00

0111

0110

011

1001

000

1110

0200

00

000

Aplo

dont

ochi

rus (

List

roph

orid

ae )

0011

0000

10

1

0101

0000

0

11

0111

0100

011

0011

000

1110

0000

00

000

Cae

noles

toco

ptes

( Rhy

ncop

tida

e )00

11–0

0000

101

1100

100

1011

1111

10

0

1100

0110

1

11

1110

0000

10

0 Sc

hizo

copt

es ( C

hir o

disc

idae

)00

10–0

0010

101

1101

000

1011

1–11

00

0

010–

–10

00

1–10

0000

00

000

Myo

copt

es ( M

yoco

ptid

ae : M

yoco

ptin

ae)

0011

0010

00

1

0001

0000

0

00

0110

0000

011

0111

000

111–

–00

–00

000

Dro

mici

ocop

tes (

Myo

copt

idae

: Dro

mic

ioco

ptin

ae)

0011

0100

10

1

0011

0001

0

10

1110

0101

001

0111

000

111–

–00

–11

0

– –

Tenr

ecob

ia ( A

topo

mel

idae

)00

10–0

0000

100

1101

000

1011

1001

00

0

0101

1110

0

11

1000

0–00

00

0

0, 1

, 2, c

hara

cter

stat

es; –

, cha

ract

er in

appl

icab

le.


of the nymphal idiosoma, and the body axes of the partners are orientated in the same direction.

Phylogenetic analysis

Maximum parsimony analysis yielded seven shortest trees (tree length 137, CI exclud-ing uninformative characters 0.36, RI 0.55, RC 0.22). Th e strict consensus of these trees (Fig. 1) shows a basal dichotomy of “higher” Psoroptidia: cluster I – Echimytricalges - Lobalges ( Psoroptoides )- Paralgopsis (most psoroptids and the genus Coendalges ); cluster II

Acaridae (Acarus)

Ptiloxenidae (Ptiloxenus) «low» Psoroptidia

Psoroptoididae (Psoroptoides)

Pyroglyphidae (Paralgopsis)

Coendalges

Edentalges

Psoroptinae (Hyracoptes)

Listropsoralginae* (Listropsoralges)

Makialginae (Makialges)

Marsupialginae (Marsupialges)

Nasalialginae (Nasalialges)

Paracaroptinae (Pangorillalges)

Cebalginae (Procebalges)

Psoralges

Psoroptinae (Trouessalges)

Echimytricalginae (Echimytricalges)

Lobalginae (Lobalges)

Listrophoridae (Aplodontochirus)

Rhyncoptidae (Caenolestocoptes)

Chirodiscidae (Schizocoptes)

Myocoptidae (Myocoptes)

Dromiciocoptinae* (Dromiciocoptes)

Atopomelidae (Tenrecobia)

28

1

22

1

12

1

51

1

27

1

15

0

13

1

4

0

47

145

1

44

1

14

1

52

1

50

1

48

1

39

1

30

1

17

1

10

1

7

1

6

1

1

1

51

1

49

1

44

1

51

1

45

1

26

1

14

110

0

46

0

36

1

26

1

50

1

36

1

26

1

17

1

1

1

10

0

48

1

45

1

44

1

36

1

14

1

47

1

40

1

38

1

22

1

18

1

17

1

8

1

5

1

2

139

0

34

1

32

1

29

1

28

1

19

1

45

1

40

1

38

1

29

1

18

1

14

1

9

0

27

1

13

1

9

0

7

150

1

49

1

30

1

19

1

6

1

38

1

9

0

53

1

52

1

35

0

13

1

1

1

32

1

31

1

13

1

4

0

48

1

4

0

35

0

17

1

37

1

10

1

27

1

7

1

46

1

46

2

37

1

16

1

14

1

12

1

26

1

39

1

20

1

17

1

4

0

14

1

51

1

44

1

36

0

27

1

26

1

13

1

28

1

21

1

37

1

36

1

33

1

25

1

24

1

15

1

Psoralginae

Psoroptidae

Lobalgidae

cluster I

Sar

cop

toid

ea c

lust

er II

«higher Psoroptidia»

22

1

3

1

1

1

3

2

1

1

1

1

1

1

2

1

1

1

3

Fig. 1. Strict consensus of 7 shortest trees found for unordered and unweighted data set. Character opti-mization, unambiguous characters only; black circles, unique synapomorphies; white circles, homoplasies; numbers above circles, characters; number below circles, character states; bold numbers above branches, Bremer indices. *Subfamilies deserving familial status.


including a single psoroptid lineage represented by the genus Listropsoralges (Listropsoralginae) and all other sarcoptoids. Th is consensus demonstrates the polyphyly of the family Lobalgidae. Th e two lobalgid genera, Lobalges and Echimytricalges, form a monophyletic group which is the sister group of the genus Psoroptoides . Th e genus Coendalges is situated in the core of psoroptids and sister to genera of the subfamily Psoralginae ( Psoralges and Edentalges ). Th e subfamily Psoroptinae ( Hyracoptes and Trouessalges ) appears as polyphyletic in our analysis.

Th e position of the genus Listropsoralges in cluster II is not surprising. Th ese mites strongly diff er by their external morphology from other representatives of the family Psoroptidae (Bochkov & Wauthy 2009 ). Similarly, the monophyly of the family Myocoptidae was questioned earlier and unique synapomorphies sharing the sub-families Myocoptinae and Dromiciocoptinae were not established (Bochkov & OConnor 2008 ).

Th e obtained consensus tree well corresponds to the molecular phylogeny of Psoroptidia recently proposed by Klimov & OConnor ( 2008 ). According to their results, the superfamily Sarcoptoidea is not monophyletic: the families Psoroptidae and Lobalgidae are closely related with pyroglyphids, mites of the family Epider-moptidae, and some other analgoids, Psoroptoididae, Turbinoptidae etc. (psoroptid- epidermoptid complex); whereas other sarcoptoids are grouped with feather mites of the superfamily Analgoidea (sarcoptid-analgid complex) (Klimov & OConnor 2008 ).

As it is visible from the consensus tree (unambiguous characters only, characters demonstrated the same changes under ACCTRAN and DELTRAN optimizations), the branch Echimytricalges–Lobalges is marked by two unique and three non-unique synapomorphies: setae si situated distinctly anterior to level of setae se (character 16), the weakly developed (?reduced) dorso-apical projection of tarsi I and II (character 46.2), and the posterior end of the female opisthosoma bears a pair of distinctly scle-rotized lateral lobes connected medially by a membrane (character 12), setae vi and σ2 I are absent (characters 14 and 37, respectively).

Th e Lobalginae as well as the Echimytricalginae lineages are each supported by sev-eral characters. Th e subfamily Lobalginae is characterized by the absence of setae c3, e III, s III, kT IV (characters 19, 28, 29, 34), fi liform shape of setae d IV and e IV in males (character 28). In Echimytricalginae, the subcapitulum bears a pair of distinct dorsal lobes (character 2), the female hysteronotal shield is subdivided (character 5), female apodemes III are fused with the epigynum (character 8), setae cp are sword-like (character 18), setae h3 in males bear membranous lobe (character 22), solenidion σ III is absent (character 38), genu and femur IV in males are fused (character 40), and a ventral projection on tarsi III in males is present (character 47). All characters listed above are unambiguous. Only one non–unique, derived character, the absence of setae c1 (17), is shared between Coendalges with Echimytricalges . Th e absence of setae c1 is also shared by the psoroptid genera Edentalges , Procebalges and Psoralges , and disappear-ance of this seta shows a mosaic distribution among many phylogenetically distant Psoroptidia. Two non-unique characters are common to Coendalges and Echimytricalges , the absence of solenidion σ III (38) and the presence of a ventral projection on tarsi III in males (47). Among psoroptid taxa in this analysis, solenidion σ III is also absent in


the genera Marsupialges , Nasalialges , Procebalges and Psoralges . Like setae c1 , this solenid ion appears to be independently lost in many other Psoroptidia.

Due to the signifi cant number of homoplasies in our data, successive weighting (Farris 1969 ) was conducted. A single tree (CI excluding uninformative characters 0.63, RI 0.83, RC 0.61) was obtained after successive weighting (Fig. 2), this tree dif-fers from the consensus tree by fully resolved cluster II only.

Based on the results of this analysis, we restrict the composition of the family Lobalgidae to the two genera, Lobalges and Echimytricalges . Although continuing to recognize two monogeneric subfamilies is taxonomically redundant, we retain them due to the signifi cant morphological and ecological distinctions between these two genera as proposed by Fain ( 1970 ). Th e genus Coendalges is formally transferred to the family Psoroptidae as suggested by OConnor ( 2009 ) and included to the subfamily Psoralginae.

Ptiloxenidae (Ptiloxenus) «low» Psoroptidia

Psoroptoididae (Psoroptoides)

Pyroglyphidae (Paralgopsis)

Coendalges

Edentalges

Psoroptinae (Hyracoptes)

Listropsoralginae* (Listropsoralges)

Makialginae (Makialges)

Marsupialginae (Marsupialges)

Nasalialginae (Nasalialges)

Paracaroptinae (Pangorillalges)

Cebalginae (Procebalges)

Psoralges

Psoroptinae (Trouessalges)

Echimytricalginae (Echimytricalges)

Lobalginae (Lobalges)

Listrophoridae (Aplodontochirus)

Rhyncoptidae (Caenolestocoptes)

Chirodiscidae (Schizocoptes)

Myocoptidae (Myocoptes)

Dromiciocoptinae* (Dromiciocoptes)

Atopomelidae (Tenrecobia)

28

1

22

1

12

1

51

1

27

1

15

0

13

1

4

0

47

145

1

44

1

14

1

52

1

50

1

48

1

39

1

30

1

14

0

10

1

7

1

6

1

1

1

51

1

49

1

44

1

36

0

51

1

45

1

26

1

14

110

0

46

0

50

1

38

1

17

1

10

0

48

1

45

1

44

1

14

1

47

1

40

1

38

1

22

1

18

1

17

1

8

1

5

1

2

139

0

34

1

32

1

29

1

28

1

19

1

26

1

13

1

45

1

40

1

38

1

29

1

18

1

17

0

9

0

9

0

7

1

50

1

49

1

30

1

19

1

6

138

1

9

0

51

1

44

1

32

1

27

1

13

1

17

1

23

1

14

1

28

1

21

1

37

1

36

1

53

1

52

1

35

0

13

1

1

1

36

1

32

1

31

1

13

1

48

1

35

0

17

1

38

1

37

1

10

1

27

1

7

1

46

1

46

2

37

1

16

1

14

1

12

1

26

1

39

1

20

1

33

1

25

1

24

1

15

1

Sar

cop

toid

ea c

lust

er II

«higher» Psoroptidia

Psoralginae

Psoroptidae

22

1

cluster I

Fig. 2. A single most parsimonious tree obtained after successive weighting. Character optimisation, unambiguous characters only; black circles, unique synapomorphies; white circles, homoplasies; numbers above circles, characters; number below circles, character states. *Subfamilies deserving familial status.


Th e primary goal of this study was to test the lobalgid monophyly. Th erefore, despite in the fact that the subfamilies Listropsoralginae and Dromiciocoptinae deserve famil-ial status, the elucidation of relationships among the various sarcoptoid families (clus-ter II) as well as main lineages (subfamilies) within Psoroptidae requires many more taxa and characters than were involved in this analysis.

Systematics

Family Lobalgidae Fain, 1965 Fain 1965 : 30, Fain & Lukoschus 1970 : 49, Fain & Mendez 1979 : 271, Fain et al. 1982 : 121, OConnor 1982 : 166, Fain & Ritzi 2001 : 255. Type genus. Lobalges Fonseca, 1954

Diagnosis

Adults . Gnathosoma without ventral apophyses. Gnathosomal setae subc and elc. p. present. Chelicerae chelate; fi xed digit with 4 distinct teeth; movable digit with indiscernible teeth. Paraxial cheliceral hood, cheliceral spur, and seta-like apophysis present. Idiosoma dorso-ventrally fl attened, elongated. Sejugal furrow not developed. Propodonotal shield present. Openings of supracoxal glands and supraxocal setae scx present, distinctly developed. Openings of opisthonotal glands distinct. Legs inserted laterally. Coxal fi elds striated. Coxal apodemes I completely fused into sternum or con-nivient and connected by strongly sclerotized bridge. Genital papillae present. In both sexes, posterior end of opisthosoma bearing pair of lateral lobes and membranous interlobar membrane. Idiosomal setation: si, se, c1 (present or absent) , c2, cp, d1–2, e1–2, h2, 3, ps1–3, 1a, 3a, 4a, 4b , and g. Setae si situated distinctly anterior to level of setae se . Setae c3 present or absent. Legs without retrorse spurs. Legs I–III consist of fi ve articulating segments and a pretarsus consisting of ambulacral stalk and ambulac-ral disc. Tarsi I and II with weakly developed (? reduced) dorso-apical curved process. Pretarsal condylophores straight; unguiform sclerites, basilar piece, and condylophore guide present. Leg setation: pR I–II, sR III, vF I–II, cG I–II , mG I–II , gT I–II , kT III, σ I–II, φ I–IV, wa I–II , ra I–II , w III–IV , r III–IV , la I–II , s I–III , d I–IV , e I–II, IV , f I–IV , ε Ι , ω1 I–II and ω3 I. Setae e III, k TIV, s III, and solenidion σ III present or absent. Male . Hysteronotal shield present. Posterior end of idiosoma bearing pair of termi-nal lobes and lobar membranes. Paranal suckers present and distinctly developed. Aedeagus situated between levels of coxal fi elds III and IV. Legs IV consisting of 4–5 articulated segments (femur and genu IV separated or fused). Legs III and IV subequal or legs IV distinctly shorter. Pretarsi III and IV present. Setae d IV and e IV fi liform or modifi ed into suckers. Female . Hysteronotum with single median shield, or pair of opisthonotal shields present posterior to this shield. Bursa copulatrix terminal. Ovipore situated ventrally, between levels of coxal fi elds III and IV. Epigynum distinctly developed, arch-like. Legs III and IV with 5 articulated segments and pretarsi, subequal in length. Tarsi III–IV without apical process.


Other genera included . Echimytricalges Fain, 1970

Remarks . As mentioned above, mites of the family Lobalgidae parasitize hosts inhabit-ing the neotropical region but belonging to distantly related orders, Pilosa (sloths of the families Bradypodidae and Megalonychidae) and Rodentia (spiny rats of the family Echimyidae belonging to both tree and ground-dwelling genera) (Table 6). Th e record of a single male E. guyanensis on an opossum is certainly the result of accidental con-tamination, as indicated by our collection of several specimens of this species from spiny rats (Table 6). Th e strong morphological distinctions between the lobalgid sub-families suggests a long time since their divergence. On the other hand, their associa-tions with such phylogenetically distant hosts strongly suggests secondary switching from one host group to the other.

Key to genera of the family Lobalgidae Fain, 1965

1. Subcapitulum dorsally without lobes. Setae cp fi liform. Setae c1 present, c3 absent. Setae h3 whip-like. Posterior ends of apodemes I fused into sternum. Solenidion σ III present. Setae e III, s III, kT IV absent. In males legs III and IV subequal, well developed; femur and genu IV not fused; setae d IV and e IV fi liform. In females hysteronotum bearing one hysteronotal shield; apodemes III short, not fused with arch of epigy-num (parasites of sloths, Pilosa: Bradypodidae and Megalonychidae) ...................….. ................................................................................................ Lobalges Fonseca, 1954 – Subcapitulum dorsally with pair of large lobes. Setae cp sword-like. Setae c1 absent, c3 present. Setae h3 with foliate membrane. Posterior ends of apodemes I ends free. Solenidion σ III absent. Setae e III, s III and kT IV present. In males legs IV strongly reduced, distinctly shorter than legs III, with fused femur and genu; setae d IV and e IV sucker-like. In females hysteronotum bearing 3 shields, unpaired central hysteronotal shield, and pair of opisthonotal shields; apodemes III distinctly developed, fused with arch of epigynum (parasites of spiny rats, Rodentia: Echimyidae) ...........................… ........................................................................................... Echimytricalges Fain, 1970

Subfamily Lobalginae Fain, 1965

Diagnosis Setae cp fi liform. Setae c1 present, c3 absent. Apodemes I completely fused into ster-num. Solenidion σ III present. Setae e III, s III, and kT IV absent. Male . Setae h3 whip-like. Legs III and IV subequal, distinctly developed; femur and genu IV not fused; tarsi IV with short apicodorsal projection. Setae d IV and e IV fi liform. Female . Hysteronotum bearing 1 hysteronotal shield. Posterogynal folds not devel-oped. Apodemes III short, not fused with arch of epigynum. Type genus . Lobalges Fonseca, 1954 . Other genera. Type genus only.


Mite

spec

ies

Hos

t spe

cies

Hos

t fam

ilyLo

calit

y (ty

pe)

Refe

renc

e

Loba

lges

Fons

eca,

195

4 @ L

obal

ges t

roue

ssart

i Fon

seca

, 195

4 * ! B

rady

pus v

arieg

atus

Sch

inz,

1825

Brad

ypod

idae

Braz

il (S

aõ P

aulo

Sta

te),

Su

rinam

e,Fo

nsec

a 19

54

@ L

obal

ges t

roue

ssart

i Fon

seca

, 195

4 ! B

rady

pus v

arieg

atus

Sch

inz,

1825

Brad

ypod

idae

Pana

ma

Our

dat

a @ L

obal

ges t

roue

ssart

i Fon

seca

, 195

4 Br

adyp

us tr

idac

tylu

s trid

acty

lus L

., 17

58Br

adyp

odid

aeSu

rinam

eFa

in 1

965 ;

Fai

n &

Luko

schu

s 197

0 @ L

obal

ges t

roue

ssart

i Fon

seca

, 195

4 ! C

hloe

pus d

idac

tylu

s (L.

, 175

8)M

egal

onyc

hida

eU

nkno

wn

Fain

196

5 Ec

him

ytric

alge

s Fai

n, 1

970

@ Ec

him

ytric

alge

s bra

silien

sis

Fa

in a

nd L

ukos

chus

, 197

0 *! P

hyllo

mys

pat

toni

Em

mon

s, Le

ite, K

ock

&

C

osta

, 200

2Ec

him

yida

e

(Ech

imyi

nae)

Braz

il (R

io d

e Ja

neiro

)Fa

in 1

970

@ Ec

him

ytric

alge

s bra

silien

sis

Fa

in a

nd L

ukos

chus

, 197

0 & Tr

inom

ys a

lbisp

inus

(I. G

eoff r

oy, 1

838)

(Ech

imyi

nae)

(Eum

ysop

inae

)Br

azil

Our

dat

a

@ Ec

him

ytric

alge

s bra

silien

sis

Fa

in a

nd L

ukos

chus

, 197

0 & Tr

inom

ys ih

erin

gi (Th

om

as, 1

911)

(Ech

imyi

nae)

(Eum

ysop

inae

)Br

azil

Our

dat

a

Echi

myt

rical

ges s

urin

amen

sis

Fa

in a

nd L

ukos

chus

, 197

0 * P

roec

him

ys g

uyan

nens

is gu

yann

ensis

(E. G

eoff r

oy, 1

803)

(Ech

imyi

nae)

(Eum

ysop

inae

)Su

rinam

e (U

itkijk

),

Guy

ana

Fain

& L

ukos

chus

1970

, our

dat

a Ec

him

ytric

alge

s sur

inam

ensis

Fain

and

Luk

osch

us, 1

970

& Pr

oech

imys

oco

nnell

i Alle

n, 1

913

(Ech

imyi

nae)

(Eum

ysop

inae

)C

olom

bia

Our

dat

a

Echi

myt

rical

ges s

urin

amen

sis

Fa

in a

nd L

ukos

chus

, 197

0 & Pr

oech

imys

qua

drup

licat

us H

ersh

kovi

tz, 1

948

(Ech

imyi

nae)

(Eum

ysop

inae

)Pe

ruFa

in &

Ritz

i 200

1

Echi

myt

rical

ges s

urin

amen

sis

Fa

in a

nd L

ukos

chus

, 197

0 & Pr

oech

imys

cuvi

eri P

ette

r , 19

78(E

chim

yina

e)

(E

umys

opin

ae)

Peru

Fain

& R

itzi 2

001

Echi

myt

rical

ges h

oplo

mys

Fain

, Luk

osch

us a

nd M

ende

z, 19

82

* & H

oplo

mys

gym

nuru

s (Th

omas

, 189

7)(E

chim

yina

e)

(E

umys

opin

ae)

Pana

ma

(Moj

unga

Swam

p); E

cuad

orFa

in e

t al.

1982

; O

ur d

ata

Echi

myt

rical

ges h

oplo

mys

Fain

, Luk

osch

us a

nd M

ende

z, 19

82

& Pr

oech

imys

sem

ispin

osus

(Tom

es, 1

860)

(Ech

imyi

nae)

(Eum

ysop

inae

)C

osta

Ric

aO

ur d

ata

Echi

myt

rical

ges p

roec

him

ys

Bo

chko

v an

d O

Con

nor,

sp.n

. * &

Proe

chim

ys b

revi

caud

a (G

ünth

er, 1

877)

(Ech

imyi

nae)

(Eum

ysop

inae

)Ec

uado

r (R

io Ja

tun

Ya

cu)

Our

dat

a

Tabl

e 6.

Hos

t and

geo

grap

hica

l dist

ribut

ion

of th

e fa

mily

Lob

algi

dae

Fain

, 196

5


Echi

myt

rical

ges m

esom

ys

Fa

in, L

ukos

chus

and

Men

dez,

1982

! M

esom

ys h

ispid

us (D

esm

ares

t, 18

17)

(Ech

imyi

nae)

(Eum

ysop

inae

)C

olom

bia

(Rio

Apop

oris)

Fain

et a

l. 19

82

Echi

myt

rical

ges m

esom

ys

Fa

in, L

ukos

chus

and

Men

dez,

1982

! M

esom

ys h

ispid

us (D

esm

ares

t, 18

17)

(Ech

imyi

nae)

(Eum

ysop

inae

)Pe

ru; B

razil

Fain

& R

itzi 2

001 ,

our d

ata

Echi

myt

rical

ges l

onch

othr

ix

Bo

chko

v an

d O

Con

nor,

sp.n

. * ! L

onch

othr

ix em

iliae

Th o

mas

, 192

0(E

chim

yina

e)

(E

umys

opin

ae)

Braz

il (B

oim

)O

ur d

ata

Echi

myt

rical

ges w

hita

keri

Fa

in a

nd R

itzi s

p. n

. *&

Proe

chim

ys b

revi

caud

a (G

ünth

er, 1

877)

(Ech

imyi

nae)

(Eum

ysop

inae

)Pe

ru (M

ayna

s)Fa

in &

Ritz

i 200

1

Echi

myt

rical

ges w

hita

keri

Fa

in a

nd R

itzi s

p. n

. & Pr

oech

imys

cuvi

eri P

ette

r, 19

78(E

chim

yina

e)

(E

umys

opin

ae)

Peru

Fain

& R

itzi 2

001

Echi

myt

rical

ges g

uyan

ensis

Fain

, 197

0 * ? C

alur

omys

phi

land

er (L

., 17

58)

Did

elph

idae

Fren

ch G

uyan

a (?

)Fa

in 1

970

Echi

myt

rical

ges g

uyan

ensis

Fain

, 197

0 & Pr

oech

imys

sim

onsi

Th om

as, 1

900

Did

elph

idae

(Eum

ysop

inae

)Pe

ruO

ur d

ata

@ , t

ype

spec

ies;

* , ty

pe h

ost;

? , ac

cide

ntal

con

tam

inat

ion;

! , arb

orea

l hos

t spe

cies

; &, t

erre

stria

l hos

t spe

cies

.


Genus Lobalges Fonse ca, 1954

Lobalges Fonseca, 1954 : 86, Fain 1965 : 25; type species: Lobalges trouessarti Fonseca, 1954 ; by original designation. Type species . Lobalges trouessarti Fonseca, 1954 . Other species. Type species only.

Diagnosis

With characters of subfamily.

Lobalges trouessarti Fonseca, 1954 (Figs 3, 4)

Lobalges trouessarti Fonseca 1954 : 86, fi gs 1–4; Fain 1965 : 26, fi gs 19–22; Fain & Lukoschus 1970 : 58. Type deposition: holotype in LPIB.

Material examined

3♂, 10♀ and 3TN (BMOC 79-0115-006, UMMZ) from Bradypus variegatus Schinz, 1825 (MVZ 136053), PANAMA: Darien, Yaviza, 8°11’N, 77°41’W, 2.xi.1967, C. Cavalier.

Host ranges and distribution

Th is species was described from Bradypus variegatus brasiliensis Blainville, 1840 in Brazil (Fonseca 1954 ). Later on it was recorded from B. tridactylus tridactylus L., 1758 from Suriname (Fain 1965 , Fain & Lukoschus 1970 ) and from Chloepus didactylus (L., 1758) (Megalonychidae) from an unknown locality (Fain 1965 ).

Redescription

Male (3 specimens from type host; Fig. 3). Body, including gnathosoma, 330–350 long, 180–205 wide. Gnathosoma 37–45 long, 45–55 wide. Propodonotal shield 58–65 long in midline, 80–90 maximum wide, without ornamentation. Posterior margin of propodonotal shield widely concave. Setae si fl anking propodonotal shield; setae se situated far from lateral margins of propodonotal shield. Hysteronotal shield 110–125 long in midline, 105–110 wide, without ornamentation, its anterior margin slightly concave; setae d1, d2, e1 and e2 situated on this shield. Opisthosomal lobes 65–72 long (length of terminal cleft) and 29–33 wide at base of lobe. Lobar mem-branes very short. Setae ps1 serrate. Dorsal cuticle distinctly striated between shields. Ventral cuticle striated including coxal fi elds. Sternum about 60 long. Apodemes II about 65 long. Aedeagus 22–23 long. Diameter of paranal suckers 12–13. Legs I –IV, excluding pretarsi, about 130, 135, 175, and 165 long, respectively. Tarsi IV 40–45 long. Lengths of some setae and solenidia: si 2–3, se 95–105, cp 15–17, ps1 90–95, h2 300–320, h3 320–345, ω1 I 20–22, ω3 I 25–27, φ I 60–65, σI–II about 35, ω1 II 17–19, φ II 65–70, φ III–IV about 40, σIII 28–31.


Fig. 3. Lobalges trouessarti Fonseca, male: (A) dorsal view; (B) ventral view; (C) tarsus I in dorsal view; (D) same in ventral view; (E) tarsus III in dorsal view; (F) tarsus IV in dorsal view; (G) same in ventral view. Scale lines: 100 μm (A, B), 50 μm (C–G).

Female (10 specimens from type host, Fig. 4). Body, including gnathosoma, 435–450 long, 190–235 wide. Gnathosoma about 65 long and wide. Propodonotal shield 80–90 long in midline, 95–100 wide, without ornamentation. Posterior margin of propodonotal shield widely concave. Setae si fl anking propodonotal shield; setae se situated far from lateral margins of propodonotal shield. Hysteronotal shield 95–100 long, 65–69 wide; bearing only setae e1 . Lateral lobes about 40 long (length of termi-nal cleft) and 35 wide at base of lobe. Dorsal cuticle distinctly striated between shields. Ventral cuticle striated including coxal fi elds. Sternum about 75 long. Apodemes II about 75 long. Arch of epigynum about 65 wide. Legs I –IV, excluding pretarsi, about


Fig. 4. Lobalges trouessarti Fonseca, female: (A) dorsal view; (B) ventral view; (C) gnathosoma in dorsal view; (D) tarsus III in dorsal view; (E) tarsus IV in ventral view. Scale lines: 100 μm (A, B), 50 μm (C–E).

120, 125, 155, 160 long, respectively. Lengths of some setae and solenidia: si 2–3, se 100–110, cp 14–15, ps1 20–22, h2 490–520, h3 430–450, ω1 I 15–16, ω3 I 28–30, φI 60–65, σI–II 19–22, ω1 II 16–17, φ II 63–65, φ III–IV about 60, σIII 28–32.

Subfamily Echimytricalginae Fain, 1970

Fain 1970 : 272, Fain & Lukoschus 1970 : 49, Fain & Mendez 1979 : 271, Fain et al. 1982 : 121, OConnor 1982 : 166, Fain & Ritzi 2001 : 255. Type genus . Echimytricalges Fain, 1970.


Diagnosis

Subcapitulum dorsally with pair of large lobes. Setae cp present, sword-like. Setae c1 absent, c3 present. Apodemes I fused only in median part, their posterior ends free. Solenidion σ III absent. Setae e III and kT IV present. Male . Setae h3 with foliate membrane. Legs IV strongly reduced, distinctly shorter than legs III, femur and genu IV fused; tarsi IV with ventro-apical projection. Setae d IV and e IV sucker-like. Female . Hysteronotum bearing 3 shields, unpaired central hysteronotal shield, and pair of opisthonotal shields. Posterogynal folds distinctly developed. Apodemes III dis-tinctly developed, fused with arch of epigynum. Other genera included . Type genus only.

Genus Echimytricalges Fain, 1970

Echimytricalges Fain 1970 : 272, Fain & Lukoschus 1970 : 49, Fain & Mendez 1979 : 271, Fain et al. 1982 : 121, Fain & Ritzi 2001 : 255. Type species: Echimytricalges bra-siliensis Fain, 1970 ; by original designation. Other species included . E. guyanensis Fain, 1970 , E. hoplomys Fain, Lukoschus and Mendez, 1982 , E. lonchothrix Bochkov and OConnor, sp.n., E. mesomys Fain, Lukoschus and Mendez, 1982 , E. proechimys Bochkov and OConnor, sp.n., E. surina-mensis Fain and Lukoschus, 1970, E. whitakeri Fain and Ritzi, sp.n.

Diagnosis

With characters of subfamily.

Echimytricalges brasiliensis Fain, 1970

Echimytricalges brasiliensis Fain 1970 : 272, Fain et al. 1982 : 123, fi g. 1, Fain & Ritzi 2001 : 255, fi gs 6, 12, 19, 26, 27. Type deposition: holotype in BMNH.

Material examined

1♂ and 1♀ (BMOC 76-1201-024, UMMZ) from Trinomys albispinus (Geoff roy, 1838) (MCZ 38311), BRAZIL: Bahia, Macaco Secco near Anadarahy, 10.xi.1913, E.H. Becker; 1♀ and 1TN (BMOC 76-1201-024, UMMZ) from Trinomys iheringi (Th omas, 1911) (MCZ 25768), BRAZIL: Sao Paulo, Ilha de Sao Sebastiao, 2.i.1906, unknown.


Th is species was described from specimens collected from two host specimens in BMNH identifi ed as Echimys brasiliensis (Lund, 1840) from Brazil (BMNH 5.4.16.4 and 5.4.16.5) (Fain 1970 ). In their revision of the genus Phyllomys , Emmons et al.


Fig. 5. Coendalges panamensis Fain and Mendez, male: (A) dorsal view; (B) same in ventral view; (C) aedeagus; (D, E), tarsi III and IV in ventral view, respectively. Scale lines: 100 μm (A, B), 50 μm (C–E).

( 2002 ) examined these specimens and determined them to be their new species, Phyllomys pattoni Emmons et al. 2002 . T. albispinus and T. iheringi from Brazil represent new host records for this species. Th e genera Phyllomys and Trinomys are represented by arboreal and terrestrial rats, respectively, belonging to diff erent echimyid subfamilies (Table 6). Th us, the simultaneous parasitism of these hosts by E. brasiliensis seems quite unusual. Fain ( 1970 ) described this species from alcohol preserved specimens of P. pattoni housed in BMNH. Th ese records could have been the result of museum contamination.

Echimytricalges surinamensis Fain and Lukoschus, 1970

Echimytricalges surinamensis Fain & Lukoschus 1970 : 58, fi gs 5–8, Fain & Ritzi 2001 : 257, fi gs 5, 9A, 13, 18, 22, 23. Type deposition: holotype in RMNH.


Fig. 6. Coendalges panamensis Fain and Mendez, female: (A) dorsal view; (B) same in ventral view; (C) tarsus I in dorsal view; (D) same in ventral view; (E) tarsus II in dorsal view; (F) same in ventral view; (G) tarsus III in dorsal view; (H) same in ventral view; (I) tarsus IV in dorsal view; (J) in ventral view. Scale lines: 100 μm (A, B), 50 μm (C–J).

Type material examined

1♂ and 1♀ paratypes (UMMZ) from Proechimys guyannensis, SURINAME: Wanica, Lelydorp, 5°42’N, 55°14’W, 18.i.1969, F.S. Lukoschus.

Additional material examined

2♂ and 2♀ (BMOC 76-1201-020, UMMZ) from Proechimys oconnelli Allen, 1913 (MCZ 27564), COLOMBIA: Magdelena, Santa Ana, 1930, Maria & Niceforo; 1♂ and 2♀ (BMOC 76-1201-016, UMMZ) from Proechimys guyannensis (E. Geoff roy, 1803) (MCZ 17848), GUYANA: Demerara River, 6°48’N, 58°10’W, 6.viii.1897, Rosenberg.


Th is species was described from Proechimys guyannensis in Suriname (Fain & Lukoschus 1970 ). It was later re-collected from Proechimys quadruplicatus Hershkovitz, 1948 and P. cuvieri Petter, 1978 in Peru (Fain & Ritzi 2001 ). We record this species from the type host from Guyana and from P. oconnelli from Colombia for the fi rst time.


Echimytricalges hoplomys Fain, Lukoschus and Mendez, 1982 (Figs 7, 8)

Echimytricalges hoplomys Fain et al., 1982 : 125, fi g. 4; Fain & Ritzi 2001 : 258, fi gs 8, 11, 16, 28, 29. Type deposition: holotype in USNM.

Material examined

1♂, 7♀, and 1 TN (BMOC 76-1201-038, UMMZ, MCZ) from Hoplomys gymnurus (Th omas, 1897) (MCZ 17846), ECUADOR: (north Ecuador) San Javier, 3.viii.1900, unknown; 5♂, 12♀, 1TN containing male, 1TN containing female, 10TN, 2PN, and 1 larva (BMOC 87-0903-005, UMMZ, FMNH) from H. gymnurus (FMNH 128460), COSTA RICA: Heredia, Parque Nacional Braulio Carrillo, 11 km S. 14.5 km W Puerto Viejo, 280m, 28.iii.1986, R.M. Timm (RMT 3585); 3♂ and 3♀ (BMOC 76-0620-070, UMMZ), from Proechimys semispinosus (Tomes, 1860) (TTU 13497),

Fig. 7. Echimytricalges hoplomys Fain et al., larva: (A) dorsal view; (B) ventral view.


Fig. 8. Echimytricalges hoplomys Fain et al., nymphs: (A) protonymph in dorsal view; (B) same in ventral view; (C) tarsus I of protonymph in dorsal view; (D) opisthosoma of tritonymph in dorsal view; (E) same in ventral view. Scale lines: 100 μm (A, B), 50 μm (D, E), 25 μm (C).

COSTA RICA: Heredia, 7 mi S Puerto Viejo, 10°23’30”N, 83°56’27”W, 7.iv.1971, V.R. McDaniel


Th is species was described from Hoplomys gymnurus in Panama (Fain et al. 1982 ). We recorded this species from the type host from Ecuador and Costa Rica. Proechimys semispinosus from Costa Rica is a new host for this species.


Echimytricalges proechimys Bochkov and OConnor, sp.n. (Figs 9, 10, 14D, E)


♂ holotype (MCZ) and 1 ♀ paratype (BMOC 76-1201-025, UMMZ) from Proechimys brevicauda (Günther, 1877) (MCZ 37895), ECUADOR: Oriente, Rio Jatun Yacu, 16.xi.1937, W. Clarke-Macintyre. Type deposition: holotype in MCZ, paratype in UMMZ.


So far this species is known only from the type host and locality, Proechimys brevicauda from Ecuador.

Etymology

Th e species name is derived from the generic name of the host and is a noun in apposition.

Description

♂ (holotype; Fig. 9). Body, including gnathosoma, 350 long, 195 wide. Gnathosoma 55 long, 70 wide. Propodonotal shield 110 long, 65 wide, not fused with antero-lateral shields, bearing pair of longitudinal median furrows. Setae si and se situated on small plates jointed to propodonotal shield. Hysteronotal shield 115 long, 100 wide, orna-mented in median part; setae d1 and d2 situated on this shield. Opisthosomal lobes 45 long. Lobar membranes distinctly developed, terminal cleft about 30 long. Dorsal cuticle distinctly striated between shields. Ventral cuticle striated including coxal fi elds. Apodemes I about 90 long, fused to each other in median part; their posterior ends slightly divergent and not fused with apodemes II. Apodemes III 55 long. Aedeagus 23 long. Diameter of paranal suckers 11. Setae ps1 broken (these setae apparently should be relatively long, because of their bases are signifi cantly larger than bases of e1 and e2 and their remnants are strong and thickened). Legs I –IV, excluding pretarsi, 115, 122, 165, and 55 long, respectively. Tarsi III 42 long, with weakly developed ventro-apical projection, 3 long. Lengths of some setae and solenidia: si 2, se 105, cp broken, h2 145, h3 135 and about 17 maximum wide, ω1 I 16, ω3 I 27, φ I 32, cG I 25, d II 70, ω1 II 18, φ II 30, φ III 10, φ IV 13. ♀ (1 paratype; Figs 10, 14E, F). Body, including gnathosoma, 500 long, 185 wide. Gnathosoma 65 long, 82 wide. Propodonotal shield 115 long, 85 wide, not fused with antero-lateral shields. Setae se situated off this shield. Hysteronotal shield 30 long, 85 wide; length/width ratio of hysteronotal shield 1:2.8; setae d1 situated off this shield. Opisthonotal shields distinctly developed 90 long, 35 wide, connected with lobar scle-rites, bearing setae e2 . Setae e1 situated on lateral margins of these shields. Setae e1 situated at level of setae e2 . Lateral lobes 25 long (terminal cleft) and 35 wide at base of lobe, ventrally jointed by narrow transverse sclerotized bridge. Median lobe 13 long and 30 wide, its posterior margin straight. Dorsal cuticle distinctly striated between


shields. Ventral cuticle striated including coxal fi elds. Apodemes I 85 long, fused in median part; their posterior ends divergent and not fused with apodemes II. Apodemes III 55 long, fused with epigynum. Arch of epigynum 70 wide. Legs I –IV, excluding pretarsi, 122, 117, 110, 115 long, respectively. Lengths of some setae and solenidia: si 2, se 145, cp 60, h2 155, h3 broken, ω1 I 14, ω3 I 35, φ I 37, cG I 24, d II 75, ω1 II 20, φ II 38, φ III–IV about 7.

Remarks

Diff erential characters of this species are given in the key below.

Echimytricalges guyanensis Fain, 1970 (Fig. 14F)

Echimytricalges guyanensis Fain, 1970 : 272, Fain et al. 1982 : 125, fi g. 2, Fain & Ritzi 2001 : 257, fi g. 9. Type deposition: holotype in IRSNB.

Fig. 9. Echimytricalges proechimys sp.n., male: (A) dorsal view; (B) ventral view; (C) tarsus III in dorsal view; (D) leg IV in ventral view. Scale lines: 100 μm (A, B), 50 μm (D, E).


Material examined

5♂, 4♀, 2TN each containing male, 1TN containing female, and 2TN (BMOC 01-0101-253, UMMZ, FMNH, MUSM) from Proechimys simonsi Th omas, 1900 (FMNH 170724), PERU: Madre de Dios, Quebrada Aguas Calientes, 2.75 km E. Shintuya, 12°88’S, 71°26’W, 12.x.1999, P.M. Velasco (PMV 450).


Th is species was described from a single male specimen from Caluromys philander (L., 1758) (Didelphimorphia: Didelphidae) from French Guyana (Fain 1970 ). Examination

Fig. 10. Echimytricalges proechimys sp.n., female: (A) dorsal view; (B) ventral view.


of additional specimens of this host by BMOC has not yielded additional specimens of this species, and we believe that this record was the result of accidental contamina-tion. We record this species from Proechimys simonsi from Peru.

Redescription

Male (5 specimens from P. simonsi ). Body, including gnathosoma, 350–365 long, 150–180 wide. Gnathosoma 55–60 long, 67–70 wide. Propodonotal shield 105–120 long, 75–80 wide, bearing pair of longitudinal median crests, fused with antero-lateral shields with striated weakly sclerotized bridges. Setae se situated on small plates joining to propodonotal shield. Hysteronotal shield 95–100 long, 85–90 maximum wide, devoid of ornamentation, without furrows; setae d1 and d2 situated on and off this shield, respectively. Terminal cleft of opisthosomal lobes about 30 long. Lobar mem-branes short, about 17 long. Dorsal cuticle distinctly striated between shields. Ventral cuticle striated including coxal fi elds. Apodemes I about 85 long, fused to each other in median part; their posterior ends slightly divergent and not fused with apodemes II. Apodemes III about 55 long. Aedeagus about 18 long. Diameter of paranal suckers about 9. Legs I –IV, excluding pretarsi, about 105, 110, 155 and 45 long, respectively. Tarsi III about 40 long, with moderately developed apicoventral projection, 4–5 long. Lengths of some setae and solenidia: si 1–2, se 140–150, cp 80–85, h2 190–210, h3 120–135 and 18 maximum width, ω1 I 10–11, ω3 I 32–33, φ I 38–40, cG I 40–45, d II 100–110, ω1 II 13–15, φ II 33–35, φ III 7–8, φ IV 12–14. Female (4 specimens from P. simonsi ; Fig. 14D). Body, including gnathosoma, 410–470 long, 150–165 wide. Gnathosoma 57–60 long, 75–78 wide. Propodonotal shield 110–122 long, 80–85 wide, not fused with antero-lateral shields. Setae se situated on small plates adjoining propodonotal shield. Hysteronotal shield 33–35 long, 84–90 wide; length/width ratio of hysteronotal shield about 1:2.7; setae d1 situated off this shield. Opisthonotal shields distinctly developed 75–80 long, 30–40 wide, bearing setae e2 , connected with lobar sclerites. Setae e1 situated slightly posterior to setae e2 . Lateral lobes about 25 long (terminal cleft) and 35 wide at base of lobe; median lobe about 10 long and 45 wide, its posterior margin almost straight. Lateral lobes ventrally joined by narrow transverse sclerotized bridge. Setae h3 about 1.4 times shorter than h2 . Dorsal cuticle distinctly striated between shields. Ventral cuticle striated including coxal fi elds. Apodemes I about 90 long, fused in median part; their posterior ends slightly divergent and not fused with apodemes II. Apodemes III about 70 long, fused with epigynum. Arch of epigynum about 70 wide. Legs I–IV, excluding pretarsi, about 115, 125, 95, 100 long, respectively. Lengths of some setae and solenidia: si 2–3, se 148–155, cp 70–75, h2 155–160, h3 110–120, ω1 I 11–13, ω3 I 34–36, φ I 38–40, cG I 34–38, d II 88–92, ω1 II 17–18, φ II 39–41, φ III–IV about 8.

Echimytricalges mesomys Fain, Lukoschus and Mendez, 1982

Echimytricalges mesomys Fain et al., 1982 : 127, fi gs 3, 5–6, Fain & Ritzi 2001 : 257, fi gs 7, 10, 15, 20, 21. Type deposition: holotype in FMNH.



1♂ and 1♀ paratypes (UMMZ) from Mesomys hispidus (Desmarest, 1817) (FMNH 57242), COLOMBIA: Amazonas, Ino Goje, Rio Apaporis, 1°23'1"S, 69°25'24"W, Cabrera.

Additional material examined

1♂, 7♀, and 1TN (BMOC 76-1201-030, UMMZ, MCZ) from Mesomys hispidus (Desmarest, 1817) (MCZ 30871), BRAZIL: Para, Rio Tapajoz, Pinhy, 18.v.1933, A.M. Olalla.


Th is species was described from Mesomys sp. in Colombia (Fain et al. 1982 ). Th e host specimen was later determined to be Mesomys hispidus ferrugineus. It was subsequently collected from Mesomys hispidus from Peru (Fain & Ritzi 2001 ). We recorded this spe-cies from M. hispidus from Brazil.

Echimytricalges lonchothrix Bochkov and OConnor, sp.n. (Figs 11–14A–C)


♂ holotype (MCZ), 3♂ and 1♀ paratypes (BMOC 76-1201-039) from Lonchothrix emiliae Th omas, 1920 (MCZ 30907-8), BRAZIL: Para, Rio Tapajoz, Boim, 03°05’S, 55°18’W, 14.xii.1932, A.M. Olalla. Type deposition: holotype in MCZ, paratypes in MCZ, UMMZ.


So far this species is known only from the type host and locality, L. emiliae from Brazil (Boim).

Etymology

Th e species name is derived from the generic name of the host and is a noun in apposition.

Description

Male (holotype; Figs 11, 13A–D). Body, including gnathosoma, 380 long (370–390 in 3 paratypes), 235 wide (210–235). Gnathosoma 55 long (53–55), 80 wide (80–85). Propodonotal shield 145 long (140–150), 110 wide (100–110), not fused with antero–lateral shields, devoid of median furrows, and bearing setae se . Hysteronotal shield 165 long (160–165), 125 wide (110–125); without striations; setae d1 situated on and setae d2 off this shield. Terminal cleft of opisthosomal lobes about 25 long. Lobar membranes short, about 10 long. Dorsal cuticle weakly striated between shields. Ventral cuticle distinctly striated including coxal fi elds. Apodemes I about 100 long,


fused to each other in median part; their posterior ends widely divergent and not fused with apodemes II. Apodemes III about 70 long. Aedeagus about 50 long. Diameter of paranal suckers about 13. Legs I –IV, excluding pretarsi, about 135, 130, 170, and 65 long, respectively. Tarsi III 65 long, with distinctly developed apicoventral projection, 13 long. Lengths of some setae and solenidia: si 3 (2–3), se 6 (5–6), cp 30 (28–30), h2 155 (150–160), h3 165 and about 10 maximum wide (160–165), ω1 I 13 (12–13), ω3 I 19 (18–19), φ I 31 (29–31), cG I 16 (16–18), d II 75 (70–80), ω1 II 15 (13–15), φ II 28 (26–29), φ III 10 (9–10), φ IV 22 (21–22).

Fig. 11. Echimytricalges lonchothrix sp.n., male: (A) dorsal view; (B) ventral view.


Female (1 paratype; Figs 12, 13E–I, 14A–C). Body, including gnathosoma, 550 long, 215 wide. Gnathosoma 65 long, 95 wide. Propodonotal shield 175 long, 125 wide, not fused with antero-lateral shields, and bearing setae se . Hysteronotal shield 110 long, 70 wide; length/width ratio of hysteronotal shield 1.7:1; setae d1 situated on this shield. Opisthonotal shields distinctly developed 120 long, 65 wide, bearing setae e1 and e2 , connected with lobar sclerites by fi ne sclerotized bands. Setae e1 situated slightly anterior to level of setae e2 . Length (terminal cleft) and width (at base of lobe) of lateral lobes subequal, about 35 long; median lobe 45 long and 35 wide, its posterior margin with triangle median incision. Ventral sclerotized bridge between lateral lobes absent. Setae h3 and h2 subequal in length. Dorsal cuticle distinctly striated between

Fig. 12. Echimytricalges lonchothrix sp.n., female: (A) dorsal view; (B) ventral view.


shields. Ventral cuticle distinctly striated including coxal fi elds. Apodemes I 115 long, fused at level of setae 1a ; their posterior ends widely divergent and not fused with apo-demes II. Apodemes III 55 long, fused with epigynum. Arch of epigynum 77 wide. Legs I –IV, excluding pretarsi, 155, 150, 120, 125 long, respectively. Lengths of some setae and solenidia: si 3, se 5, cp 47, h2 155, h3 145, ω1 I 13, ω3 I 22, φ I 30, cG I 15–17, d II 120, ω1 II 12, φ II 35, φ III–IV about 10.

Remarks

Diff erential characters of this species are given in the key below.

Echimytricalges whitakeri Fain and Ritzi, sp. n.

Echimytricalges whitakeri Fain and Ritzi, 2001 : 258, fi gs 5, 14, 17, 24, 25 nomen nudum

Fig. 13. Echimytricalges lonchothrix sp.n., details: Male (A– D): (A) tarsus I in ventral view; (B) tarsus III in ventral view; (C) leg IV in ventral view; (D) aedeagus. Female (E–I): (E) tarsus I in dorsal view; (F) same in ventral view; (G) tarsus II in dorsal view; (H) tarsus III in ventral view; (I) same in dorsal view; (J) pretarsus IV.



Holotype ♀ (USNM), 1♀ (IRSNB) and 2♂ (USNM, IRSNB) paratypes from Proechimys brevicauda (host no. CLH 2654), PERU: Loreto, Maynas, Estacion Biologica Allpahuayo, 21 km (by road) S of Iquitos, 03°58’S, 73°25’W, 1.ix.1998, C.L. Hice. Type deposition: holotype in USNM, paratypes in IRSNB, USNM.


Th is species was collected from Proechimys brevicauda (type host) and P. cuvieri from Peru (Fain & Ritzi 2001 ).

Remarks

Fain & Ritzi ( 2001 ) provided a description of this species and designated a holotype but did not explicitly indicate where the holotype was deposited. Because Article 16.4.2 of the International Code of Zoological Nomenclature, 4th edition (1999) requires the name and location of the type depository as a criterion of availability after 1999, this name is a nomen nudum . In order to make this name available, we repeat the descrip-tion given by Fain and Ritzi ( 2001 ), updating setal nomenclature: “♀ (holotype). Body 450 long (in midline, gnathosoma included), 180 wide. In 2 paratypes 438×160 and 426×165. Dorsum: Propodonotal shield 96 long, 80 wide. Angterolateral punctate areas poorly developed and widely separated from propodono-tal shield. Setae se set on small punctate plates connected with shield. Mediodorsal

Fig. 14. Echimytricalges spp., details: E. lonchothrix (A–C): (A) gnathosoma in dorsal view; (B) same in ventral view; (C) female opisthosomal lobe in dorsal view. E. proechimys : (D) female opisthosomal lobe in dorsal view; (E) female opisthosomal lobe in dorsal view. E. guyanensis : (F) sclerotized parts of female copulatory system.


shield wider (93) than long (maximum length 93). Paired opisthonotal plates extending on posterior lobes, total length 135, maximum width 45. Distance l5–l5 (= h2–h2 ) 57. Posterior extremity ending into 2 sclerotized lobes aboaut 30 long and 12 wide in pos-terior half. Setae l5 (= h2 ) 80 long. Setae h (= cp ) very thick, 45 long. Venter: Sternum very thick, with posterior branches slightly divergent and not reaching epimera II. Epigynium ( sic ) regularly rounded, maximum width 80. Male (3 paratypes). Total length in midline and width in 3 paratypes: 330×180, 331×185 and 338×186. Cuticular striations well developed. Propodonotal shield 90 long, 63 wide, with lateral convex margins. Th e anterolateral punctate areas very small and widely separated from propodonotal shield. Hysteronotal shield 120 long, 90 wide (maximum). Diameter of adanal suckers 9 and 33 apart. Distance l5–l5 (= h2–h2 ) 90. Tarsus IV 42 long, normal in shape and slightly curved apically.

Key to species of the genus Echimytricalges Fain, 1970

Males 1. Propodonotal shield without median longitudinal crests ...................................... 5 – Propodonotal shield with pair of distinct median longitudinal crests .................... 2 2. Hysteronotal shield without ornamentation ........................................................ 4 – Hysteronotal shield distinctly striated in median part ........................................... 3 3. Setae si situated on propodonotal shield. Ventro-apical projection of tarsi III dis-tinctly developed, about 7 long. Setae h2 about 2 times longer than h3 …................. ......................................................... E. hoplomys Fain, Lukoschus and Mendez, 1982 – Setae si situated off propodonotal shield. Ventro-apical projection of tarsi III weakly developed, about 3 long. Setae h3 and h2 subequal in length …..................................................................................... E. proechimys Bochkov and OConnor, sp.n. (Fig. 9) 4. Propodonotal shield fused with antero-lateral shields. Bridges between propodono-tal shield and antero-lateral sclerites striate. Hysteronotal shield with pair of longitudi-nal median folds in posterior part …......... E. surinamensis Fain and Lukoschus, 1970 – Propodonotal shield not fused with antero-lateral shields. Hysteronotal shield with-out folds …........................................................................... E. guyanensis Fain, 1970 5. Setae si and se situated on propodonotal shield. Tarsi IV with distinct apicoventral projection ................................................................................................................ 6 – Setae si and se situated off propodonotal shield. Tarsi IV without distinct apicoventral projection ….......................................................... E. whitakeri Fain and Ritzi, sp. n. 6. Setae se macrosetae .............................................................................................. 7 – Setae se short, 5–6 long …...................................................................................... ...................................E. lonchothrix Bochkov and OConnor, sp.n. (Figs 11, 13A–D) 7. Apodemes I fused with apodemes II …............................................................................................................................ E. mesomys Fain, Lukoschus and Mendez, 1982 – Apodemes I separated from apodemes II …....................... E. brasiliensis Fain, 1970

Females (Females of E. guyanensis Fain, 1970 , E. surinamensis Fain and Lukoschus, 1970 , and E. proechimys Bochkov and OConnor, sp.n. are indistinguishable)


1. Hysterosomal shield at least 2.5 time wider than long ......................................... 3 – Hysteronotal shield longer than wide or its length and width subequal … ............. 2 2. Setae se macrosetae. Apodemes I fused with apodemes II. Median opisthosomal lobe widely rounded. Setae h3 about 1.7 times shorter than h2 …................................................................................................ E. mesomys Fain, Lukoschus and Mendez, 1982 – Setae se short, 5–6 long. Apodemes I separated from apodemes II. Median opistho-somal lobe with triangular incision. Setae h3 and h2 subequal …....................................................... E. lonchothrix Bochkov and OConnor, sp.n. (Figs 12, 13E–I, 14A–C) 3. Setae se situated off propodonotal shield. Dorsal striations numerous. Setae e1 situ-ated at level or posterior to setae e2 .......................................................................... 4 – Setae se situated on propodonotal shield. Dorsal striation widely separated. Setae e1 situated anterior to level of setae e2 …................................. E. brasiliensis Fain, 1970 4. Setae si situated on propodonotal shield … .......................................................... 5 – Setae si situated off propodonotal shield .............. E . whitakeri Fain and Ritzi, sp. n. 5. Lateral lobes without distinctly developed triangular projection near bases of setae h3 …............................................... E. guyanensis Fain, 1970 , E. surinamensis Fain and Lukoschus, 1970 , and E. proechimys Bochkov and OConnor, sp.n. (Figs 10, 14D–E) – Lateral lobes with distinctly developed triangular projection near bases of setae h3 … ......................................................... E. hoplomys Fain, Lukoschus and Mendez, 1982

Acknowledgements

Th e authors thank Dr. Serge Mironov (ZISP) for his critical review of the manuscript. We would like to thank the various curators and institutions who provided material for this study, especially, the late Dr. Alex Fain and Dr. Georges Wauthy (IRSNB). Th is research was supported by a grant from the US National Science Foundation DEB-0118766 (PEET) to B.M.OC. and by a grant from the Russian Foundation for Basic Research (07-04-00426a) to A.V.B.

References

Atyeo , W.T. ( 1978 ) Th e pretarsi of astigmatid mites . Acarologia 20 : 245 – 269 . Bochkov , A.V. & OConnor , B.M. ( 2006 ) Th e life-cycle of Hemigalichus chrotogale sp. nov. (Acari:

Listrophoridae), with comparative observations on listrophorid morphology . Journal of Natural History 39 : 3811 – 3832 .

Bochkov , A.V. & OConnor , B.M. ( 2008 ) Morphology and systematics of the mite subfamily Dromiciocoptinae (Acari: Myocoptidae), with description of two new species . Annals of the Entomological Society of America 101 : 289 – 296 .

Bochkov , A.V. & Wauthy , G. ( 2009 ) A review of the genus Listropsoralges (Acari, Psoroptidae) with the descriptions of two new species . Acta Parasitologica 54 : 269 – 275 .

Emmons , L.H , Leite , Y.L.R. , Kock , D. & Costa , L.P. ( 2002 ) A review of the named forms of Phyllomys (Rodentia: Echimyidae) with the description of a new species from coastal Brazil . American Museum Novitates 3380 : 1 – 40 .

Fain , A. ( 1965 ) Les acariens producteurs de gale chez les edentes et les marsupiaux (Psoropridae et Lobalgidae: Sarcoptiformes) . Bulletin de Institut royal des Sciences Naturelles de Belgique 41 : 1 – 41 .


Fain , A. ( 1970 ) Diagnoses de nouveaux Lobalgides and Listrophorides (Acarina: Sarcoptiformes) . Revue de Zoologie et de Botanique africaines 81 : 271 – 300 .

Fain , A. & Lukoschus , F.S. ( 1970 ) Parasitic mites of Suriname II. Skin and fur mites of the families Psoroptidae and Lobalgidae . Acta Zoologica et Pathologica Antverpensia 51 : 49 – 60 .

Fain , A. , Lukoschus , F.S. & Mendez , E. ( 1982 ) Two new species of the genus Echimytricalges Fain, 1970 (Acari, Astigmata, Lobalgidae) from American spiny rats (Echimyidae) . Bulletin and Annales de la Société royale Belge d’Entomologie 118 : 121 – 130 .

Fain , A. & Mendez , E. ( 1979 ) Coendalges panamensis g.n., sp.n. from the porcupine in Panama (Astigmata: Lobalgidae) . International Journal of Acarology 5 : 271 – 274 .

Fain , A. & Ritzi , C.M. ( 2001 ) Observations on some astigmatid mites (Acari) parasitic on rodents (Rodentia) from Peru, with description of three new species . International Journal of Acarology 27 : 251 – 259 .

Farris , J.S. ( 1969 ) A successive approximations approach to character weighting . Systematic Zoology 18 : 374 – 385 .

Fonseca , F. ( 1954 ) Notas de Acarologia. XXXVIII. Sarcoptiformes da preguica; Lobalges trouessarti gen. n., sp. n. (Acari. Epidermoptidae) . Memorias do Instituto Butantan 26 : 85 – 92 .

Grandjean , F. ( 1939 ) La chaetotaxie des pattes chez les Acaridiae . Bulletin de la Société Zoologique de France 64 : 50 – 60 .

Griffi ths , D.A. , Atyeo , W.T. , Norton , R.A. & Lynch , C.A. ( 1990 ) Th e idiosomal chaetotaxy of astigmatid mites . Journal of Zoology, London 220 : 1 – 32 .

Klimov , P.B. & OConnor , B.M. ( 2008 ). Origin and higher-level relationships of psoroptidian mites (Acari: Astigmata: Psoroptidia): evidence from three nuclear genes . Molecular Phylogenetics and Evolution 47 : 1135 – 1156 .

Klompen , J.S.H. ( 1993 ) Phylogenetic relationships in the skin-inhabiting Sarcoptoidea (Acari: Astigmata) . Systematic Parasitology 24 : 17 – 33 .

Krantz , G.W. ( 1978 ) A manual of Acarology , second edition. Oregon State University Book Stores , Corvallis, OR , 509 pp.

Mironov , S.V. & Bochkov , A.V. ( 2009 ) Modern conceptions of macrophylogeny in acariform mites (Chelicerata, Acariformes ) . Zoologicheskiy Zhurnal 88 : 922 – 937 . [In Russian with English summary]

Nixon , K.C. ( 2000 ) WinClada. Version 0.9. 0.9.99m24. Program and documentation in cladistic.com . Ithaca, NY.

Norton , R. ( 1998 ) Morphological evidence for the evolutionary origin of Astigmata (Acari: Acariformes) . Experimental & Applied Acarology 22 : 559 – 594 .

OConnor , B.M. ( 1981 ) A systematic revision of family-group taxa in the non-psoroptid Astigmata (Acari: Acariformes) . Ph. D. thesis, Cornell University , Ithaca, NY , 594 pp.

OConnor , B.M. ( 1982 ) Astigmata : In: Parker , S.B. (Ed.) Synopsis and Classifi cation of Living Organisms , Vol. 2 . McGraw-Hill , New York, NY , pp. 146 – 169 .

OConnor , B.M. ( 1984 ) Co-evolutionary patterns between astigmatid mites and primates . In: Griffi ths , D.A. & Bowman , C.E. (Eds) Acarology VI , Vol. 1 . Ellis Horwood , Chichester , pp. 186 – 195 .

OConnor , B.M. ( 2009 ) Cohort Astigmatina . In: Krantz , G.W. & Walter , D.E. (Eds) A Manual of Acarology , third edition. Texas Tech University Press , Lubbock, TX , pp. 565 – 657 .

Page , R.D.M. ( 2001 ) NDE: NEXUS Data Editor 0.5.0 . University of Glasgow , Glasgow . Sorenson , M.D. ( 1999 ) TreeRot, version 2 . Boston University , Boston, MA . Swoff ord , D.L. ( 2001 ) PAUP*: Phylogenetic Analysis Using Parsimony (* and Other Methods), Version 4 .

Sinauer Associates , Sunderland, MA . Wilson , D.E. & Reeder , D.M. ( 2005 ) Mammal species of the world. A taxonomic and geographic reference ,

3rd edn. Th e Johns Hopkins University Press , Baltimore, MD , 2142 pp. Witalinski , W. , Dabert , J. & Walzl , M.G. ( 1992 ) Morphological adaptation for precopulatory guarding

in astigmatic mites (Acari: Acaridida) . International Journal of Acarology 18 : 49 – 54 . Wurst , E. ( 1993 ) Investigations on the anatomy and the behaviour of the fur mite Listrophorus leuckarti

(Acari: Listrophoridae) . Stuttgarter Beiträge zur Naturkunde. Serie A (Biologie) 503 : 1 – 68 .


Appendix A.

List of comparative material used in cladistic analysis

Acaridae: Acarus siro (L., 1758) 5♂, 5♀ (BMOC 97-0221-001, UMMZ) in chicken feed, COSTA RICA: San Jose, san Pedro de Mt. de Oca, Universidad de Costa Rica, Lab. Arvicola, 9°56’N, 84°3’W, 14.xi.1996, G. Bonilla. Ptiloxenidae: Ptiloxenus elegans (Cerny, 1969) 1♂, 1♀ (University of Adam Mickiewicz, Poznan, Poland) from Tachybaptus domini-cus (L., 1766) (Podicipediformes: Podicipedidae) (USNM 390654), PARAGUAY, Anaha, Chao, ii.1938, unknown. Psoroptoididae: Psoroptoides truncatipes (Trouessart and Neumann, 1888) 1♂, 1♀ (ZISP) from Buceros hydrocorax L., 1766 (Coraciiformes: Bucerotidae,), THE PHILIPPINES: Leyte Island, Baybay, Mt. Pangasugan, 22.x.1984, A. Almoroda. Pyroglyphidae: Paralgopsis sp. 5♂, 5♀ (BMOC 87-0318-004, UMMZ) from Ara militaris (L., 1766) [subspecies mexicana ] (Psittaciformes: Psittacidae) (UMMZ 72164), MEXICO: Sinaloa, Rosario, 27.xii.1933, C.C. Lamb. Psoroptidae: Psoroptinae: Hyracoptes emersoni Fain and Lukoschus, 1981 1♂, 1♀ paratypes (UMMZ) from Dendrohyrax dorsalis (Fraser, 1855) (Hyracoidea: Procaviidae), ZAIRE: Tandala, Gemena, 21.vii.1979, Emerson. Trouessalges peccari Fonseca, 1954 2♂, 1♀ (UMMZ) from Pecari tajacu (L., 1758) (Tayassuidae), Hildago Co., 24-25.ii.1964. Psoralginae: Edentalges choloepi Fain, 1964 1♂, 1♀ paratypes (IRSNB) from Choloepus didactylus L., 1758 (Pilosa: Megalonichidae), other data unknown. Psoralges libertus Trouessart, 1896 5♂, 1♀ (BMOC 76-0620-022, UMMZ) ex Tamandua tetradactyla (L., 1758) (Pilosa: Myrmicophagidae), COSTA RICA: San Jose, 8.8 mi SW San Isidro del General, 11.viii.1971, R.S. Baker. Listropsoralginae: Listropsoralges marmosa Fain, 1965 4♂, 1♀ paratypes (IRSNB) from Marmosa robinsoni fulviventer Bangs, 1901 (Dedelphimorphia: Didelphidae), PANAMA: Perlas archipelago, San Miguel Island, 08°27’00˝N, 78°51’00˝W, 05.viii.1970, Ritte. Makialginae: Makialges lepilemuri (Gaud and Till, 1957) 1♂, 1♀ (ZISP) from Lepilemur mustelinus Geoff roy, 1851 (Primates: Lemuridae), MADAGASCAR: Ambatolampy, other data unknown. Marsupialginae: Marsupialges missonei Fain, 1963 1♂, 1♀ paratypes (IRSNB) from Caluromys philander (L., 1758) (Didelphimorphia: Didelphidae), FRENCH GUIANA: other data unknown. Nasalialginae: Nasalialges borneensis Fain and Nadchatram, 1979 1♂, 1♀ paratypes (IRSNB) from Nasalis larvatus (Wumb, 1787) (Primates: Cercopithecidae) (IRSNB), Malaysia, Sabah, Kuala Penyu District (N.E. Borneo), vi.1977, M. Nadchatram.


Paracoroptinae: Pangorillalges gorillae (Gaud and Till, 1957) 1♂, 1♀ paratype (ZISP) from Gorilla gorilla (Savage and Wyman, 1847) (Primates: Hominidae) (UMMZ), CAMEROON: Yaounde, iv.1956, unknown. Cebalginae: Procebalges pitheciae Fain, 1963 10♂, 10♀ (BMOC 81-0809-009 UMMZ), Pithecia monachus (Geoff roy, 1812) (Primates: Cebidae) (FMNH 122796), PERU: Loreto, Nauta, Rio Samiria, 30.xi.1980, P. Hershkovitz (PH 9088). Lobalgidae: Coendalginae: Coendalges panamensis Fain and Mendez, 1979 1♂, 1♀ paratypes (IRSNB) from Coendou rothschildi Th omas, 1902 (Rodentia: Erethizontidae), PANAMA: Panama Province, Aguacate, Capira, 4.i.1979, H. Montenegro. Listrophoridae: Aplodontochirus borealis Fain and Hyland, 1972 2♂, 1♀ (BMOC 79-1020-001, UMMZ) from Aplodontia rufa (Rafi nesque, 1817) (Rodentia: Aplodontidae), USA: Oregon, Benton Co., Mary’s Peak, 44˚30’N, 123˚33’W, 20.x.1979, B. M. OConnor. Rhyncoptidae: Caenolestocoptes inca Fain and Lukoschus, 1976 1♂, 1♀ paratypes (IRSNB) from Lestoros inca Th omas, 1917 (Paucituberculata: Caenolestidae), PERU: Matchu Pichu Valley, Torontoy, iv.1915, E. Heller [mites removed by F.S. Lukoschus] Chirodiscidae: Schizocoptes chrysochloris Fain, 1970 1♂, 1♀ paratypes (ZISP) from Calcochloris leucorhinus (Huet, 1885) (Afrosoricida: Chrysochloridae) , DEMOCRATIC REPUBLIC OF CONGO: Luluabourg, 15.i.1964, A. Fain Myocoptidae: Myocoptinae : Myocoptes pitymys Fain and Bochkov, 2004 1♂, 1♀ paratypes (IRSNB) from Microtus pinetorum (Le Conte, 1830) (Rodentia: Cricetidae), USA: Illinois, Clark Co., 6 mi ME Marshall, 27.xi.1968, D.D. Pascal (DDP # 458). Dromiciocoptinae: Dromiciocoptes marmosops Bochkov and OConnor, 2008 1♂ paratype (BMOC 01-0101-181, ZISP) from Marmosops parvidens Tate, 1931 (Didelphimorphia: Didelphidae) (FMNH 169800), PERU: Madre de Dios, Manu, Quebrada Aguas Calientes, left bank, Rio Alto Madre de Dios, 2.75 km E Shintuya, 450 m. 30.viii.1999, S. Solari (SS 1796). Dromiciocoptes caenolestes Bochkov and OConnor, 2008 1♀ paratype (BMOC 81-0129-11, UMMZ) from Caenolestes`fuliginosus (Tomes, 1863) (Paucituberculata: Caenolestidae) (UMMZ 127157), ECUADOR: Napo, 6.9 km W. Papallacto, 12.vii.1978, R. S. Voss. Atopomelidae: Tenrecobia tenrec Fain, 1970 9♂, 4♀ (BMOC 99-0620-021, UMMZ) from Tenrec ecaudatus (Schreber, 1778) (Afrosoricida: Tenrecidae) (SMG 10647), MADAGASCAR: Fianarantsoa, Foret d’Ianasana, 7 km W Itremo, along Atsirakambaity River, 1630 m, 20°36’6”S, 48°25’24”E, 3.ii.1999, S.M. Goodman (SMG 10647).


Appendix B

Characters used in cladistic analysis

(0), ancestral state; (1 or 2), derived states.

1. Ventral retrorse apophyses of gnathosoma: absent (0); present (1) 2. Dorsal lobes of subcapitulum: absent (0); present (1) 3. Hysteronotal shield in males: absent (0); present (1) 4. Hysteronotal shield in females: absent (0); present (1) 5. Female hysteronotal shield: entire (0); subdivided onto 3 shields, median and 2

opisthonotal (1); subdivided onto 3 shields, 2 hysteronotal and pygidial (2) 6. Opisthogastric shield in female: absent (0); present (1) 7. Apodemes of coxae I: fused (0); separated (1) 8. Development of apodemes III in females: not fused with epigynum (0); fused with

epigynum (1) 9. Epigynum: represented by indistinct, weakly sclerotized plate (0); distinctly

developed, strongly sclerotized, arch-like (1) 10. Position of epigynum: situated between levels of coxal fi elds II and III (0); situated

between levels of coxal fi elds I and II (1) 11. Opisthosomal lobes in males: absent (0); present (1) 12. Opisthosomal lobes and interlobar membrane in females: absent (0); present (1) 13. Setae scx : present (0); absent (1) 14. Setae vi : present (0); absent (1) 15. Setae ve : present (0); absent (1) 16. Position of setae si : at level or slightly anterior or posterior to setae se (0); distinctly

anterior to setae se (1) 17. Setae c1 : present (0); absent (1) 18. Setae cp : fi liform (0); sword-like (1) 19. Setae c3 : present (0); absent (1) 20. Setae h1 : present (0); absent (1) 21. Setae h3 in males: whip-like (0); as short as ps1 (1) 22. Shape of setae h3 in males: without membranous lobe (0); with membranous

lobe (1) 23. Setae h3 in females: as long as h2 (0); much shorter than h2 (1) 24. Setae ad in females: at least one pair present (0); absent (1) 25. Setae p and q of tarsi I, II: present (0); absent (1) 26. Setae ba I–II: present (0); absent (1) 27. Setae d of posterior legs: not longer or only slightly longer than tibia–tarsus (0);

distinctly longer than tibio–tarsus at least on one pair of legs (1) 28. Setae d IV and e IV (if present) in males: sucker-like (0); fi liform (1) 29. Setae e III: present (0); absent (1) 30. Setae eIV in males: present (0); absent or alveolar (1) 31. Setae s I, II: present (0); absent (1) 32. Setae s III: present (0); absent (1)


33. Setae s IV: present (0); absent (1) 34. Setae kT IV: present (0); absent (1) 35. Solenidion ω1 I: situated in basal or median part of tarsus (0); situated in apical

part of tarsus (1) 36. Solenidion ω1 II: situated in basal or median part of tarsus (0); situated in apical

part of tarsus (1) 37. Solenidion σ2 I: present (0); absent (1) 38. Solenidion σ III: present (0); absent (1) 39. Legs III in males: subequal to legs IV (0); longer and thickened than legs IV (1) 40. Genu and femur IV in males: separated (0); fused (1) 41. Pretarsi: not modifi ed into discs (0); modifi ed into discs (1) 42. Empodium of tarsi I–II: present (0); absent or strongly reduced (1) 43. Condylophore guide: absent (0); present (1) 44. Pretarsi III in males: present, distinctly developed (0); absent or strongly reduced

to small stalk (1) 45. Pretarsi III in females: present (0); absent (1) 46. Dorso-apical projection on tarsi I–II: absent (0); large (1); very small

(?reduced) (2) 47. Ventro-subapical projection on tarsi III in males: absent (0); present (1) 48. Tarsi III–IV in females: longer than tibiae (0); shorter than tibiae (1) 49. Ventral spur of coxae I–II: absent (0); present (1) 50. Ventral spur of coxae III: absent (0); present (1) 51. Ventral retrorse spurs on tarsi, tibiae and genua I–II: absent (0); present (1) 52. Ventral retrorse spurs on tarsi and tibia III in females: absent (0); present (1) 53. Ventral retrorse spur on tarsi IV in females: absent (0); present (1)

External morphology and systematics of the family Lobalgidae (Acari: Astigmata) with resolution of...

Documents

Transcript of External morphology and systematics of the family Lobalgidae (Acari: Astigmata) with resolution of...