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Bulletin of the University of Nebraska State Museum Museum, University of Nebraska State

3-1971

A Generic Revision of the Pterodedinae, a New Subfamily of Feather Mites (Sarcoptiformes: Analgoidea)

Chong K. Park

Warren T. Atyeo

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This Article is brought to you for free and open access by the Museum, University of Nebraska State at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Bulletin of the University of Nebraska State Museum by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. BULLETIN OF VOLUME 9, NUMBER 3 The University of Nebraska State Museum MARCH. 1971

Chong K. Park and Warren T. Atyeo

A Generic Revision of the Pterodedinae, a New Subfamily of Feather Mites (Sarcoptiformes: Analgoidea) Chong K. Park

and

Warren T. Atyeo

A Generic Revision of the Pterodectinae, a New Subfamily of Feather Mites (Sarcoptiformes: Analgoidea)

BULLETIN OF The University of Nebraska State Museum VOLUME 9, NUMBER 3 MARCH, 1971 BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM VOLUME 9, NUMBER 3 Pp. 39-88, Figs. 1-68 MARCH, 1971

ABSTRACT

A Generic Revision of the Pterodectinae, a New Subfamily of Feather Mites (Sarcoptiformes: Analgoidea)

Chong K. Park Warren T. Atyeo

The generic rev1s1on of the Pterodectinae, new subfamily of the Proctophyllodidae, is based on 88 named and over 160 new species. Included are morphology, host-parasite relationships, diagnoses of four named and eight new genera, and illustrations of each type species. The named genera are: Anisodiscus Gaud and Mouchet, 1957; Montesauria Oudemans, 1905; Proterothrix Gaud, 1968; Pterodectes Robin, 1877. The new genera and their type species are: Dolichodectes, Proctophyllodes (Pterocolus) edwardsi Trouessart, 1885; Megalodectes, Proctophyllodes (Pterodectes) major Trouessart, 1885; Neodectes, Proctophyl/odes (Pterodectes) securiclatus Trouessart and Neumann, 1888; Pedanodectes, Pterodectes hologaster Gaud, 1953; Synto­ modectes, Proctophyllodes (Pterodectes) selenurus Trouessart, 1885; Toxerodectes, Ptero­ dectes gladiger hastifolia Trouessart, 1899; Trochilodectes, Proctophyllodes (Pterodectes) trochilidarum Trouessart, 1885; Xynonodectes, Proctophyllodes (Pterodectes) gracilior Trouessart, 1885. The subgenus Pterodectes (Proterothrix) Gaud, 1968, is elevated to genus. Eighty-eight species are assigned to the appropriate genera, six species are unassigned, Pterodectes rotifer (Trouessart and Neumann), 1888, is re-assigned to the genus Trouessartia, and Pterodectes armatus Banks, 1909, is assigned to the genus Proctophyllodes. New synonymy: Montesauria cylindrica (Robin), 1877 = Pterodectes corvincola Oudemans, 1905. Park1 and Atyeo2

A Generic Revision of the Pterodectinae, a New Subfamily of Feather Mites (Sarcoptiformes: Analgoidea)3

INTRODUCTION he demonstrated that tritonymph and adult fe­ males laid viable bisexual eggs. The genus Pterodectes Robin, 1877 (Procto­ The present study, based on over 250 new phyllodidae) as previously defined (Trouessart, and described species, is the first concerted 1885, 1899; Gaud, 1952, 1953; Gaud and Mou­ effort to recognize and diagnose supraspecific chet, 1957) is one of the largest groups of categories within the genus Pterodectes (s.l.). feather mites, but includes a heterogenous as­ The genera Pterodectes (s.s.), Montesauria, An­ semblage of species. Numerous species groups isodiscus, Proterothrix and eight new genera are evident for the described species, but only are defined; the type species for each genus is the genera Anisodiscus Gaud and Mouchet, illustrated, and described species are re-as­ 1957, Montesauria Oudemans, 1905, and Pro­ signed. Descriptions and redescriptions of the terothrix Gaud, 1968, have been recognized as more than 250 species will appear in future supraspecific taxa. The bionomics of these aca­ studies. rines, broadly classified as epizoic scavengers, The material for this investigation is part of are virtually unknown. Popp (1967) in conjunc­ an extensive feather mite collection now housed tion with studies of the morphology of the repro­ at the University of Georgia. The collection con­ ductive systems, conducted mating experiments sists of approximately 16,000 vials and 35,000 with two species of Pterodectes (s.l.) in which slides acquired through examination of 1,900 field collected birds and 20,000 museum study skins, and through loans and exchanges with 1 Chong K. Park: Research Assistant, Department of Entomology, University of Nebraska, Lincoln, Nebraska. various persons and museums. In addition, Present Position: Post-doctoral Research Associate, De­ through the cooperation of Ors. Jean Gaud and partment of Entomology, University of Georgia, Athens, Max Vachon, types have been made available Georgia. for most of the described species. 2 Warren T. Atyeo: Professor of Entomology, Department of Entomology, University of Nebraska, and Curator of The collection and preparation of the feather Entomology, University of Nebraska State Museum, Lin­ mite specimens follow the procedures of Atyeo coln, Nebraska. Present Position: Professor of Entomology, Department and Braasch (1966). The optical equipment used of Entomology, University of Georgia, Athens, Georgia, in this study included a Wild-Heerbrugg phase­ and Research Associate, Division of Entomology, Univer­ contrast microscope with drawing attachment sity of Nebraska State Museum, Lincoln, Nebraska. and an AO Spencer phase-contrast microscope • The research was supported by the National Science Foundation (GB-7943, GB-8606, GB-15105). equipped with an ocular micrometer. 39 40 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

MORPHOLOGY tures encountered in studying this group of The general idiosomal conformation of both acarines (Figs. 1-4). An analysis of the gnatho­ the males and females is relatively simple. Modi­ soma is not included; it is similar to that of fications of the male genital region, the idioso­ Proctophyllodes species described by Donald E. mal termini of the male and female, and the Johnston (in Atyeo and Braasch, 1966). The ter­ female spermatheca are some of the more use­ minology used for the descriptive morphology ful characteristics for species differentiation. follows Atyeo and Braasch (1966) and the chae­ For illustrative purposes, a hypothetical species totaxal signatures are those of Atyeo and Gaud has been created to include morphological fea- (1966).

----- pore

13 ____ Sac

Figs. 1 and 2. Hypothetical pterodectine male. A, anus; AD, adanal discs; AS, anal shields; EP1-••• epimerites; GD, genital discs; GO, genital organ; MS, metapodosomal shields; Sac, supranal concavity; TC, terminal cleft; VOS, ventrolateral shields. SETAE: a, anal; Ci-a, centrals; ex., coxal Ill; d1-•, dorsal hysterosomals; h, humeral; '1-s, lateral hysterosomals; pae, pai, external and internal postanals; s, coxal I; see, sci, external and internal scapulars; sh, subhumeral; ve, external vertical. A GENERIC REVISION OF THE PTERODECTINAE / 41

ldiosoma gladiform appendage supported internally by Dorsal idiosoma (Figs. 1 and 3). The propodo­ a sclerotized rod. If these appendages are somal shield bears two or three pairs of setae; absent, setae d5 are extremely long. the scapular setae (sci, see) are always pres­ Ventral idiosoma (Figs. 2 and 4). An impor­ ent and the external vertical setae (ve) may tant diagnostic feature of the ventral idiosoma be present or absent. The external margins of is the pattern formed by the epimerites, apo­ this shield may be indistinct as the sclerotiza­ demes, and associated sclerites (sternocoxal tion gradually diminishes and blends with the skeleton). The various conditions of epimerites surrounding striated integument. Nevertheless, 1-111 are usually consistent within a genus; the the approximate shape of the shield can be posterior epimerites (llla-lVa) vary between taxa described as approximately rectangular, trian­ and between sexes. Without exception there are gular, or trapezoidal. The shape of the propod­ surface sclerotizations (surface shields) closely osomal shield is generally constant within a associated with one or more of the epimerites. genus and as such, has little value for species Epimerites I are basically Y-, V-, or ir-shaped differentiation. with or without posterolateral extensions (com­ Two scapular shields, when present, are im­ pare Figs. 2, 22, 36, and 38). Occasionally, the mediately posterior to legs II and the shields posterolateral extensions are sufficiently devel­ complete the complement of sclerotized regions oped to extend to epimerites 11 thereby enclos­ of the dorsal (and lateral) propodosoma. The ing coxal fields I (Fig. 54). Epimerites II usually scapular shields never bear setae. curve toward the midline and end free, rarely The shields of the hysterosoma consist of the are they connected with epimerites Ila to form large median shield, two lateral humeral shields closed coxal fields. The latter epimerites of this anterior to legs 111, and rarely two small meta­ coxa (Ila) are always associated with the pos­ podosomal shields between legs Ill and IV. Ex­ teroventral edge of the scapular shields and cept for setae 11, the hysterosomal shield usually usually terminate as bluntly rounded apodemes bears all of the posterior setae, the openings of before reaching the meson. There may be a pair the dorsal hysterosomal glands, the supranal of small internal structures that are presumably concavity, and the various ornamentations asso­ remnants of these epimerites mesal to the major ciated with the termini (Figs. 1, 3). The humeral terminations. shiel~s often bear the lateral pair of the first row Epimerites Ill are ventrolateral apodemes as­ of hysterosomal setae (/1), the long humeral sociated with the humeral shields and the ante­ setae (h), and the short, often bladelike sub­ rior articulations of legs Ill; those of the male humeral setae (sh). and female are similar (Figs. 2 and 4). In a fow With the exception of the new genus Toxero­ genera, extensions of the epimerites form closed dectes, the male hysterosoma usually tapers coxal fields (Fig. 30). gradually from the posterior articulations of legs A complex arrangement of the posterior epim­ IV to the terminus. The terminus is weakly to erites, surface shields, and apodemes is illus­ distinctly bilobed, the lobes being separated by trated in a hypothetical male (Fig. 2). Epimerites a terminal cleft of various configurations which Illa + IV and IVa are joined on each side may be unique to certain genera. through a fusion with a median Y-shaped scler­ The female hysterosoma is primitively divided ite. Epimerites IVa of each side connect ante­ into a large anterior portion and a smaller termi­ riorly to the genital arch and surface shields of nal region which may bear ensiform append­ IVa are present posterior to the genital arch. ages. The terminus is bilobed and the dorsal Commonly the median Y-shaped sclerite is ab­ surface is usually separated from the idiosoma sent and epimerites IVa and associated sclero­ proper by a distinct suture. In a few genera tizations are weakly developed. For additional this suture is absent or partially developed. As information, see the section on the male genital in the male, the terminal cleft may be variously region. formed. Each of the posterior lobes may bear a In the region between the male genital arch 42 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

h

3 4 Figs. 3 and 4. Hypothetical pterodectine female. A, anus; Ep,-•• , epimerites; GD, genital discs; IS, interlobar shield; PgA, pregenital apodeme; Sac, supranal concavity; TC, terminal cleft. SETAE: a, anal; c,-,, centrals; cxs, coxal Ill; d,-,, dorsal hysterosomals; h, humeral; Ii-,, lateral hysterosomals; pae, pai, external and inter­ nal postanals; s, coxal I; see, sci, external and internal scapulars; sh, subhumeral; ve, external vertical. and the adanal discs, there may be one or more discs there may be sclerotized areas, termed sclerites. The more anterior, mentioned above, the ventrolateral shields (VOS); thE·se areas may connect epimerites IVa across the venter may bridge the lateral margins and the anterior of the mite (Fig. 2) or may appear as two small cleft (Fig. 2), may appear as extensive lateral shields connected with the terminations of the shields (Fig. 30), or be absent. In a few heavily genital arch. These latter are simply expansions sclerotized species (not figured), the entire ven­ of the genital arch. Anterior to the adanal discs tral surface posterior to the adanal discs may and posterior to setae c3 are the small adanal be sclerotized. shields (AS) that may or may not be connected In the female (Fig. 4), the pregenital apodeme to each other and may or may not bear the anal and epimerites IV are fused into an omega­ setae (a). Posterior and/or lateral to the adanal shaped arch characteristic for the related gen- A GENERIC REVISION OF THE PTERODECTINAE / 43 era cited in this study. The pregenital apodeme positioned posterior to the genital arch, the is basically in the shape of an inverted U with discs may not be evident (Fig. 30). rounded or square corners or in the shape of an The various apodemes adjacent to or con­ arc of a circle. The epimerites connected to the nected with the genital arch are usually very pregenital apodeme may be short to long and conspicuous. Although elements of the posterior have variously shaped shields associated with epimerites of coxae IV may extend in front of them. Not associated with the epimerites or pre­ the genital apparatus, a pregenital apodeme per genital apodeme are ventral sclerotized areas se occurs only in species of Anisodiscus. The joining the idiosomal margins to the top of the apodemal configurations result primarily from interlobar cleft, the interlobar shields US). These modifications of the (antero)mesally directed shields may be weakly to well developed and posterior epimerites of coxae IV (epimerites may bear setae pae. IVa). These epimerites may be absent, weakly Male genital region (Fig. 2). The structures of developed, united across the venter of the idio­ the male genital region and their relationships soma, or united with the supporting structures to each other and to other components of the of the genital organ, i.e., the genital arch. ventral hysterosoma provide important criteria In the genera Pterodectes, Proterothrix, Neo­ for the differentiation of species and genera. dectes, Megalodectes, Toxerodectes and Xyn­ The length of the genital organ, the presence or onodectes and many species of Montesauria, absence of a pregenital apodeme, the develop­ the genital arch is positioned between coxae ment of the shields, and the positioning of the IV and with few exceptions, is independent of genital discs and ventral setae are examples of the weakly developed epimerites IVa (Figs. 22 these characters. and 38). The exceptions are species in which The obvious structures of the male reproduc­ epimerites IVa are well developed and directed tive system are the genital arch and the stylet­ anteromesal from trochanters IV to anterior to like genital organ. The primary reproductive the genital arch where they end free (not fig­ system, however, consists of paired testes, ured). In a few species of Montesauria and in paired vas defferens uniting into an annulated Anisodiscus megadiscus, the genital arch is common duct, a seminal vesicle and an acces­ positioned between the anterior articulations of sory gland each leading to the common duct trochanters IV and epimerites IVa are weakly (internal ejaculatory duct), and a three-cham­ joined to the posterolateral terminations of the bered sperm pump (Popp, 1967). Our method of arch (not figured). The species of Trochilodec­ slide preparation destroys the testes, vas deffer­ tes, new genus, have epimerites IVa well devel­ ens, seminal receptacle, and accessory gland; oped and connected anterior to the genital arch the first visible internal structures of the system to form a prominent inverted U-shaped apo­ are the annulated duct and sperm pump. Two deme (Fig. 54). This apodeme is independent of variations of Popp's illustration of the pump are the genital arch and bears the insertions of the noted: the third chamber (Blase 1//) may be second pair of central setae (c2). The heavily wanting and the small internal plate dorsal to sclerotized species of Dolichodectes, new ge­ the sperm pump (lnnenskeletales F/Ogelpaar) nus, have the genital arch surrounded by the may be circular or have the lateral tips directed many sclerites associated with the epimerites rostrad. The development of the latter structures (Fig. 30). varies considerably within the various taxa. With the exception of the Montesauria species Two pairs of atrophied genital discs are posi­ and Anisodiscus megadiscus previously men­ tioned either anterior or posterior to the genital tioned, the genital structures and the epimerites arch. The discs on each side are usually ap­ have been independent from each other. The proximate to each other and are often borne on species of Pedanodectes have epimerites IVa small sclerotizations; they are widely separated anastomosed with the anterior edge of the gen­ in only a few species (Fig. 50). In heavily sclero­ ital arch. The mesal ends of these epimerites tized species and especially if the discs are may end at the level of the apex of the arch (as 44 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

in Fig. 34), may curve rostrad and end free be­ of feather mites. Subfamilies of the Proctophyl­ fore reaching setae c1 (not figured), or may join lodidae, for example, can be separated by the the opposite member immediately anterior to presence or absence of solenidia cr1 on genua II, the genital organ (as in Trochilodectes, Fig. 54). the structure of the pretarsi, and the condition In Anisodiscus (except megadiscus), the gen­ of the articulations between the genua and ital arch is midway between coxae Ill and IV femora. and the posterolateral terminations of the arch Most genera in this study are characterized are joined with epimerites IVa. Short, mesally by five-segmented, subequal legs which have directed apodemes arise from the fused pos­ the genua and femora freely articulated. The terior epimerites of legs Ill (=Illa) and the ante­ only obvious hypertrophy occurs in species of rior epimerites of legs IV to connect a short and Montesauria, Pterodectes, Neodectes, and Pro­ distinct pregenital arch (Fig. 38). terothrix in which legs I may be enlarged. A Female genital region. There are two external few groups have males in which legs Ill-IV or IV openings to the reproductive system-an in­ are slightly stouter and thicker than legs 1-11, verted V- or Y-shaped oviporus (tocostome) im­ and a few groups have the articulation between mediately posterior to the crescentic pregenital the genu and femur partially fused. apodeme and a copulatory opening anterior to The pretarsi have rounded ambulacra in the terminal cleft which leads into the bursa which the condylophores are unguiform; apical copulatrix. Combining observations of the vis­ points may be present. The sizes of the ambu­ ible internal structures (after specimen prep­ lacra vary according to genus or species; they aration) with Popp's (1967) study, the female may be subequal (Figs. 17-20) or 1-11 may be reproductive system is as follows. The copula­ larger (Figs. 9-12) or smaller (Figs. 5-8) than tory opening is variously positioned along the Ill-IV. It is noted that the larger ambulacra may midline from anterior to the dorsal limits of the appear spade-shaped as there is a tendency for terminal cleft (often marked by the supranal the lateral margins to be folded. concavity) ventral to the posterior limits of the Chaetotaxy anal slit. Immediately internal to the opening there may be a small expanded bursa copulatrix Since Atyeo and Gaud (1966) proposed chae­ which is the terminal ending of the primary totaxal signatures for the sarcoptiform feather spermduct (bursa copulatrix of Popp). Before mites, the system has been successfully used entering the large seminal receptacle (receptac­ for various mite groups. The system is applied ulum seminalis), the sperm duct may be locally to the genera and species cited in this study. expanded and/or surrounded by a thinly granu­ The chaetotaxy of the idiosoma and legs is lated sheath. The receptacle, a voluminous, very similar to that described for the genus thin-walled sac, is connected to the ovaries by Proctophy/lodes by Atyeo and Braasch (1966). two secondary spermducts (our preparations Except for the marked deviations from Procto­ show only the basal portions of these ducts). phy/Jodes, only a resume and illustrations of the Popp illustrates two ovaries, two lateral ovi­ setae and their positions are included (Figs. ducts, a median oviduct (uterus or vagina of 1-20). Popp), and the oviporus (tocostome of authors). Dorsal idiosoma. The propodosomal shield The oviporus is flanked by two lightly sclero­ bears the short internal and the long external tized latigynial plates and the apex is marked scapular setae (sci, see) and, if present, the by a minute, often hexagonal, sclerite. The lati­ external vertical setae (ve). As in all proctophyl­ gynial plates are usually connected to the junc­ lodine mites, the internal vertical setae (vertical tions of the pregenital apodeme and epimerites setae of authors) are always absent. IV. Two pairs of atrophied genital discs and two The dorsal hystersoma theoretically has five transverse rows of four setae per row (d1_5, l1_5); pairs of setae (c1 , c2) are lateral to the oviporus. the anterior four rows are microsetae and the Legs fifth row is composed of variously modified mac­

Leg morphology is important in the taxonomy rosetae. In males, setae 13 are simple and setae A GENERIC REVISION OF THE PTERODECTINAE / 45

vF 5

9

gT cG

gT

vF ra 6

mG e vF 10

sR

7

1 1

8 w 12 Figs. 5-12. Legs I-IV of Anisodiscus megacau/us (Trouessart) female (5-8) and Proterothrix phy/lura (Troues­ sart) male (9-12). 46 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

13

vF 17

e ~.!, • vF 14

e vF 18

d

e

15

19

16

Figs. 13-20. Legs I-IV of Xynonodectes species male (13-16) and Mega/odectes major (Trouessart) male (17-20).

d 5 may be setiform, ovate, lanceolate, or spic- appendages are small unless the appendages

uliform. In females, setae /5, positioned on the are absent, in which case the lengths of the lateral expansions of the terminus are simple or setae are greatly exaggerated. bladelike, with or without a terminal filament; Two additional pairs of setae may appear on

setae d 5 , positioned at the base of the terminal the dorsal or lateral surfaces of the terminus, A GENERIC REVISION OF THE PTERODECTINAE / 47 the postanal setae (pai, pae). In males, the inter­ a1 are subequal to or larger than w3 in length nal postanals are usually on the mesal margins (Fig. 13). Finally, setae cG and mG on genua I of the hysterosomal lobes and are setiform or or 1-11 may be modified into spinelike setae in spiculiform. The external postanals are conspic­ some genera (Figs. 9, 17, and 18). uous and ventrolateral to setae 14 and antero­ ventral to setae 15 • In females, setae pai are on HOST-PARASITE RELATIONSHIPS the cleft margins or middorsal on the lobes; Limitations of the Study setae pae are ventral and inserted between The University of Georgia feather mite collec­ setae 15 and the anus. tion is probably the largest in the world. Al­ Lateral idiosoma. Anterior to legs 111 are the though the collection contains at least a few long humeral setae (h) and the more posteriorly samples from every avian order except the large positioned subhumeral setae (sh). The humerals ratites and penguins, limitations do exist that are always long and setiform; the subhumerals may introduce bias into observations on host­ are short and vary in shape from spiculiform to parasite relationships. In amassing material, bladelike. large numbers of field collected samples have Ventral idiosoma. The usual six pairs are pres­ been acquired from the United States, South ent: two pairs of coxal setae, three pairs of cen­ Africa, and southeastern Asia. The African and tral setae, and one pair of anal setae; in females, North American collections are general while a seventh pair is present, the external postanals those from the remaining area are primarily (mentioned above). In characterizing males, the from the Apodiformes, Trogoniformes, Pici­ relative positions of the posterior pair of central formes (Picidae only), and the Passeriformes setae (c3), the anal setae (a), and the adanal (Passeres only). Over a nine-year period, we discs are useful. For example, members of the have collected from a wide range of host spe­ pair c3 may be closer to each other than are the cies at various museums (see Acknowledge­ members of the pair of anals, and the anal setae ments) either by attempting to find mites on may be positioned anterior, anterolateral, lat­ every species or many representative species eral, or posterolateral to the adanal discs. The within every family. To date we have had insuffi­ external postanal setae and setae 14 may be in cient time to examine study skins from major a ventral position due to the encrouchment of groups of birds (except hosts for named feather the dorsal hysterosomal shield onto the ventral mite species), namely, Falconiformes, Columbi­ region. formes, Psittaciformes, Musophagiformes, Cu­ The ventral chaetotaxy of the female is con­ culiformes, Caprimulgiformes, and numerous sistent with other proctophyllodine genera. With families of the Passeriformes. With the excep­ the coalescence of the pregenital apodeme and tion of the latter order, we do not believe these the epimerites, the two anterior pairs of central orders to be important hosts for the Ptero­ setae are within the top of the omega-shaped dectinae. pregenital apodeme and lateral to the oviporus. Relationships Legs. There are several differences between Based on our collection containing an esti­ the leg chaetotaxy of Proctophyllodes and cer­ mated 250 species of pterodectine mites from tain genera in this study. Setae ba, la, wa on over 500 species of birds, it can be stated that tarsi 1-11 may be in a whorl as in Proctophyllodes the genera and species of the Pterodectinae (Figs. 13 and 14) or the ventral member (wa) occur primarily on the Passeriformes and Apodi­ may be subapical, that is, distant from the other formes. Additionally, from our collection and members of the whorl (Figs. 9, 10, 17, and 18). from literature records, we know that a limited Setae sR on trochanters Ill and solenidia a1 on number of species have adapted to birds from genua Ill may be lacking, or only a1 may be ab­ other than two primary host orders. sent (Fig. 7); in Proctophyl/odes, both structures The common relationship of the birds and are always present. Solenidia a1 are usually mites is a one parasite-one host association; smaller than w3 on legs I, but in certain genera the next most common is one parasite on two 48 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM or more closely related host species. As in­ sufficient for major adaptations as have species formation accumulates, the number of single of the Proctophyllodinae. In the latter group the host-single parasite associations will probably taxa described from the Tyranni are unique to decrease as it is thought that a pterodectine that avian suborder. species is able to live on a number of related Only one species is known to be associated host species. It should also be emphasized that with the suborder Menurae. An extremely large one mite species often shares the same host mite, Megalodectes major (Trouessart), 1885, species (or genus) with other pterodectine and occurs on the lyrebird Menura superba. Ptero­ non-pterodectine species. dectine mites are not known from the second The species of the four new genera of the species of Menura or from the two species of Trochilodectes group are restricted to the Tro­ the family Atrichornithidae. chilidae, but there is no demonstrable host For the suborder Passeres (Oscines of au­ specificity. It appears that any pterodectine thors), sixty percent of the families contain species of this group is able to subsist on any known hosts of pterodectine mites. This per­ species of hummingbird {although two species centage is expected to be much higher when of Trochilodectes have been collected only from all the host families have been adequately species of Ag/aeactis). The apparent lack of host studied. As would be expected, records indicate specificity is peculiar; obviously there has been that the larger genera of mites occur on more geographical isolation of the hosts and multiple families of birds than do the smaller genera. invasions of the ectoparastiic arthropods, but it However, it should be emphasized that there is not uncommon to find numerous species of are within these larger genera loosely defined mites on one bird specimen. Additional records species groups that may eventually be restricted may show that certain mites are associated with to certain host groups. The smaller genera have certain species or groups of species of hum­ a more restricted host list: Anisodiscus from mingbirds and/or that some mite species are Nectariniidae and Sylviidae; Pedanodectes from restricted in their geographical distribution. The Dicaeidae, Laniidae, Muscicapidae, Nectarini­ only certainty has been stated-the distinct idae, and Pycnonotidae; and Dolichodectes from hummingbirds mites are found only on hum­ Muscicapidae, Sylviidae, and Turdidae. mingbirds and do not occur even on the other Considering next those mites from birds other families of the Apodiformes. than the orders Apodiformes and Passeriformes, The majority of the pterodectine species are there are only limited numbers of samples, associated with birds of the order Passeri­ some of which are questionable associations. formes. The ectoparasites, although known to Falling into the suspect categories are the col­ occur on all of the passeriform suborders are lections from the Strigiformes (Tytonidae and not evenly distributed through these groups. Strigidae); each of the species appear to be From the suborder Eurylaimi, one species has conspecific with species known to occur on the been described: Proterothrix xiphiura (Troues­ Passeriformes. In other orders there are valid sart), 1885 from Psarisomus dalhousiae and an­ associations: one species from Musophagi­ other species, mentioned by Trouessart (1885) formes (Musophagidae), one from Trogoni­ as a variety of Pterodectes mainati from Eurylai­ formes (Trogonidae), one from Coraciiformes mus ochromelas, will eventually be described as {Alcedinidae), and one or more species from a species of Proterothrix. the Piciformes (Galbulidae, Ramphasitidae, Species of the larger genera Pterodectes and Picidae). Proterothrix are known to occur on families of Trouessart (1885) described Pterodectes trulla the Tyranni, namely, Furnariidae, Cotingidae, from Tauraco macrorhynchus (Musophagidae) Tyrannidae, and Phytotomidae. The mite species from Gabon. This species, the only one known are typical of the mentioned genera and either from the Musophagiformes, belongs to a Pro­ they have not evolved as rapidly or have not terothrix species group that is found only on the been associated with the hosts for a period Paradisaeidae of New Guinea. The ranges of A GENERIC REVISION OF THE PTERODECTINAE / 49 the two families, Musophagidae and Paradisae­ relates to the mite taxa has been general, but idae, do not overlap today, the former being from after critical evaluation of all species and rec­ Africa south of the Sahara (except Madagascar) ords has been completed, it is probable that and the latter being from the Moluccas, New more families and species will be added to the Guinea and adjacent islands, and north and host lists, but that the essential inter- and eastern Australia. The obvious implications of intragroup relationships will not be significantly past sympatry could be made. changed. One species of Pterodectes occurs on the TAXONOMY Trogonidae of the New World. Although many collections have been examined from the Old Historical Account World genera, especially Harpactes, pterodec­ The name Pterodectes (s./.) first appeared in tine species have never been discovered. The a footnote in Robin's "Memoire sur les Sarcop­ possibility that the New and Old World Trogoni­ tides avicoles et sur les metamorphoses des dae each supports a unique fauna will be ex­ Acariens" in 1868. In this footnote (pp. 786-7) plored after all of our information is collated. Robin stated that his investigation was based In the Piciformes, extensive collections have on species of Dermaleichus Koch and several been obtained from the Bucconidae, Ramphasti­ new genera-Ptero/ichus, Pteronyssus, Procto­ dae, and Jyngidae but there is a paucity of phyl/odes, and Pterodectes. However, the de­ material from the Galbulidae, Capitonidae, ln­ scriptions of the new genera and species were dicatoridae, and Picidae. A few species of not published until 1877 when Robin (in Robin Pterodectes occur on the Ramphastidae, Galbu­ and Megnin) described several taxa including lidae and Picidae and one species of Protero­ the subgenus Pterodectes. Each of his new thrix has been recovered from the latter family. taxa was footnoted by a reference to the 1868 It is doubtful that pterodectine mites will be paper-a paper which in essence contained only nomina nuda. found on the puffbirds or wrynecks, but it is probable that when sufficient representatives of Trouessart (1885) and Trouessart and Neu­ the 208 species of woodpeckers have been ex­ mann (1888), in addition to describing many amined, a number of new species will be found. new species of Pterodectes, divided the genus As concerns the remaining families, it may be Proctophyllodes into five subgenera: Procto­ phyl/odes, Trouessartia (=Pteroco/us), Alloptes, that a limited number of additional species and/ Pterodectes and Pterophagus. These subgenera or host records will be forthcoming. were recognized as genera by Canestrini and The Alcedinidae is the only family in the Kramer (1899) and Trouessart (1915). The only Coraciiformes known to harbor pterodectine other genera erected for pterodectine species mites. One species of Proterothrix has been have been Montesauria Oudemans, 1905, Anisa­ found associated with the kingfishers. Other discus Gaud and Mouchet, 1957, and Protero­ families of this avian order are infested by thrix Gaud, 1968. species of the proctophyllodid subfamily Troues­ Other than the descriptions of Montesauria sartinae, but species from neither the Procto­ and a few new species, there was little activity phyllodinae nor Alloptinae have been recovered. on the Pterodectinae from 1900 until Gaud In summary, the orders Apodiformes, Trogoni­ (1952, 1953), Gaud and Mouchet (1957) and Till formes, Coraciiformes and Piciformes appear to (1954, 1957), described many new species dis­ have been invaded on numerous occasions by covered in studies of the African fauna. Gaud members of the Pterodectinae, probably by spe­ (1962, 1964) has since described species from cies normally found on the Passeriformes. Only other regions as have Berla (1958, 1959, 1960), those mites associated with the Apodiformes Cerny (1963), and Vassilev (1958). (Trochilidae) have had sufficient time to evolve into a distinctive fauna; species from the other Synonymies orders are related to or conspecific with species In this and future investigations on the Ptero­ from the Passeriformes. The discussion as it dectinae, we intend to give only the pertinent 50 i BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM synonymies for the genera and species. Many ZSZM Zoologisches Staatsinstitut und Zoo­ of the works published have been faunal lists logisches Museum, Von-Melle-Park or host-parasite lists compiled from various lit­ 10, 2000 Hamburg 13, Germany. ertaure sources, and as such have added little Descriptive Terminology-Genera new information. Publications of this nature (e.g., Canestrini and Kramer, 1899; Gaud and In addition to the familial and subfamilial Till, 1961; Radford, 1953, 1958; Poppe, 1888; features, each genus is defined by thirty-two Turk, 1953) unless they contain new records or characters. To facilitate comparisons, each nomenclatural changes are not cited, but are character is numbered and the same sequence listed in the bibliography. is used in each definition. The few terms that are not in common usage or are not self-ex­ Deposition of Type Material planatory are defined below. The character In the descriptive sections, the names of in­ number is given in parentheses. stitutions and persons receiving primary and Metapodosomal shields (6): A pair of small secondary types are denoted by the following plates on males situated on the dorsal hystero­ abbreviations: soma lateral to the hysterosomal shield between BAS Zoological Institute, Bulgarian Acad­ legs Ill and IV (Fig. 53). These structures are emy of Sciences, Boulevard Ruski 1, unique to species of Trochilodectes, new genus. Sofia, Bulgaria. Ventrolateral shields, male (7): These sclero­ BMNH British Museum (Natural History), tizations, if present, are connected to, or exten­ Cromwell Road, London S.W. 7, sions of the dorsal hysterosomal shield and United Kingdom. serve as strengthening devices for the ventro­ lateral opisthosoma. The shields are variously GAUD Dr. J. Gaud, Laboratoire de Parasit­ shaped and may be confined to the lateral mar­ ologie, Faculte de Medecine, Rennes gins (Figs. 34 and 38), may extend from the (llle-et-Vilaine), France. margins to the anterior limits of the terminal LAS Zoological Institute, Academy of Sci­ cleft (Fig. 54), or may connect across the venter ences of the U.S.S.R., Leningrad posterior to the adanal discs (Fig. 30). 8-164, U.S.S.R. Seta/ arrangements (14): The positions of two NU University of Nebraska, Lincoln, Ne­ pairs of setae in relation to each other are said braska 68503. to be arranged in a square, a rectangle, or a RNH Rikjmuseum van Natuurlikje Historie, trapezoid. Raamsteeg 2, Leiden, Netherlands. Hysterosomal terminus, female (21): The dis­ SAIMR South African Institute for Medical tinct posterior section of the hysterosoma, the Research, Hospital Street, Post Of­ terminus, usually bears two lobes, various setae, fice Box 1038, Johannesburg, South and terminal appendages. The terminus may be Africa. freely articulated, or partially or completely SEA Stazione Entomologica Agraria, via fused with the anterior hysterosoma, the degree Romana 15-17, Florence, Italy. is reflected by the completeness of the con­ junctiva separating the terminus from the re­ TC Trouessart Collection, c/o Dr. Max mainder of the idiosoma. Vachon, 61 rue de Bufton, 75 Paris, Genitocoxal apodemes, female (23): This France. complex is composed of the pregenital apo­ UGA University of Georgia, Athens, Geor­ deme and the epimerites of the posterior two gia 30601. pairs of legs. The pregenital apodeme has three USNM United States National Museum, basic shapes: 1) omega-shaped, oval (Fig. 24) Washington, D. C. 20560. or circular (Fig. 35) arc approximating 270°, 2) ZSBS Zoologische Sammlung des Bayer­ inverted U-shaped (Fig. 52), and 3) inverted U­ ischen Staates, Menzingstraase 67, shaped with angular corners (Fig. 60). The Munich 19, Germany. length of the pregenital apodeme is measured A GENERIC REVISION OF THE PTERODECTINAE / 51

as the vertical distance between the apex to the Within the genera of the Proctophyllodinae, level of the posterolateral limits (where the arch the females are more similar in form than the joins the posterior epimerites). The length of the males; often it is difficult to make species deter­ coxal apodemes is the vertical distance from the minations based on the females. However, using limits of the pregenital apodeme to the level of the two basic arrangements of the pregenital the posterior limits of epimerites IV. The gentio­ apodeme and epimerites IV, females can be coxal apodemes are considered as: short if the divided into two distinct groups. In one group length of the pregenital apodeme is greater than these structures are independent and in the the length of the coxal apodemes (Fig. 64), nor­ other, the posterolateral ends of the pregenital mal if the two measurements are approximately apodeme connect with epimerites IV to form a equal (Figs. 24, 52, and 60), or elongated if the Moresque arch-shaped structure (enlarged Q). coxal apodemes are more than 1 ½ times longer On the bases of these and other modifications in than the pregenital apodeme (Figs. 36 and 40). the males and females, the genera can be placed into two major groups which are desig­ Descriptive Terminology-Species nated as subfamilies. The format will be the same for each species to be described or redescribed in future papers Proctophyllodinae dealing with the Pterodectinae. Those charac­ (Apodemes independent) ters and descriptive methods which for clarity Allodectes Gaud and Berla, 1963 need to be defined are explained below; for Anisophyl/odes Atyeo, 1967 additional discussions refer to the Morphology Bradyphyllodes Atyeo and Gaud, 1970 section. Diproctophyllodes Atyeo and Gaud, 1968 Male Favettea Trouessart, 1915 Length of body. Distance between pedipalp Hemipterodectes Berla, 1959 apices and the terminus without considering the Monojoubertia Radford, 1950 terminal setae. Nycteridocaulus Atyeo, 1966 Length of hysterosomal shield. Distance be­ Philepittalges Atyeo, 1966 tween the most anterior point and the terminus. Proctophyllodes Robin, 1877 Length of genital organ. Distance between the Ptyctophyl/odes Atyeo, 1967 top of the genital arch and the apex of the Tanyphyllodes Atyeo, 1966 genital organ. Distance between adanal discs. Measurement Pterodectinae, new subfamily between the centers of the discs. (Apodemes joined) Female Anisodiscus Gaud & Mouchet, 1957 Length of body. Distance between pedipalp Dolichodectes, new genus apices and the end of the hysterosomal lobes at Megalodectes, new genus the level of setae d5, excluding the terminal Montesauria Oudemans, 1905 appendages. Neodectes, new genus Pedanodectes, new genus Family PROCTOPHYLLODIDAE Proterothrix Gaud, 1968 Trouessart and Megnin Pterodectes Robin, 1877 The family Proctophyllodidae includes forty­ Syntomodectes, new genus four named genera which have been separated Toxerodectes, new genus into three subfamilies: the Alloptinae (19 gen­ Trochilodectes, new genus era), the Proctophyllodinae (16 genera), and the Xynonodectes, new genus Trouessartiinae (9 genera). The Alloptinae and The genus Allodectes appears to be mis­ Trouessartiinae will not be discussed as each is placed; this monotypic genus, erected for Proc­ a distinct group and each will eventually be tophyllodes (Alloptes) norneri Trouessart, 1885, afforded familial rank. is restricted to the avian family Trochilidae. 52 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

Both sexes have the coxal fields heavily sclero­ Diagnosis: Proctophyllodid mites; males tized and all legs have the genua and femora strongly or weakly bilobed, with ensiform genital incompletely fused. The males have legs IV en­ organ, without terminal lamellae; females with larged and the hysterosomal terminus entire; pregenital apodeme and epimerites IV joined in they resemble the males of the genera Alloptes a broad Q-shape and usually with distinct termi­ Canestrini, 1879 (Alloptinae) or Monojoubertia. nal region bearing well-developed lobes and The females resemble the Pterodectes group ensiform appendages. only in the connections of the pregenital apo­ ldiosoma with dorsal shields; propodosoma deme to the posterior epimerites. These con­ with internal vertical setae (vi) absent, external nections are very weak and probably are a vertical setae (ve) present or absent; hystero­ reflection of the extremely dense sclerotizations soma usually with five pairs of dorsal (d1-r.) and of coxal fields Ill and IV. five pairs of lateral U1-a) setae; setae d1-s, l4 and/ A few species occurring on the Tyranni ap­ or pai may be absent; setae d1_4 and 11-4 are usu­ pear to be intermediate between the two sub­ ally microsetae, setae d 5 and 15 are macrosetae families. The males have small terminal lamellae and may be variously modified. ldiosomal ven­ and a genital organ similar to species of the ter with or without shields; epimerites I are Proctophyllodinae, but the general impression V-, Y-, or :u-shaped, with or without posterolat­ of the idiosoma, ventral apodemes, anal shields, eral extensions. Legs five segmented, usually and positions of the central and anal setae im­ subequal, femorogenual articulations free to mediately suggest the genus Proterothrix. The partially fused; solenidion cr2 absent on genu I; females of these species are typical of the Proc­ solenidia cr1_2 absent on genu 11; solenidion cr1 tophyllodinae except the pregenital apodeme and seta sR may be absent on legs Ill; setae ba, extends almost to, or weakly connects the pos­ s, p, q, absent from tarsi HI; ambulacra usually terior epimerites. The species of this group are: ovoid with triangular apotele and unguiform Pterodectes minor Berla, 1959, P. ocelatus condylophores. Berla, 1960, both from the Furnariidae; possibly Type genus: Pterodectes Robin, 1877. P. intermedius (Trouessart), 1885, from the Eurylaimidae; and a new species from the Key to the genera of the Pterodectinae

Dendrocolaptidae. 1. Both sexes with solenidion cr1 smaller PTERODECTINAE, new subfamily than solenidion w3 on legs I and seta The new taxon is based on the comparative wa distant from setae ra and la on studies of approximately 250 species, most of legs 1-11 (Figs. 5 and 9); on non-apodi­ which have not been described. It will be noted form birds; the Pterodectes group...... 2 that a few of the larger genera can be divided Both sexes with solenidion cr1 subequal into species groups that may be recognized as or larger than w3 on legs I and setae genera after additional material has been wa, ra, and la approximate and ar­ studied from key hosts. ranged in a whorl on legs HI (Fig. 13); lntergeneric relationships among the ptero­ only on Trochilidae; the Trochilo- dectine mites are based on comparative morph­ dectes group ...... 9 ology and when possible, host preference, 2. Male with anal setae (a) anterior to ada- realizing that adaptations to the various hosts nal discs and positioned mesal to disc may be strongly reflected in the morphological centers; setae a and c 3 in rectangular modifications of the ectoparasites. The four new arrangement ...... 3 genera from hummingbirds (the Trochilodectes Male with anal setae lateral or posterior group) and the remaining genera (the Ptero­ to adanal discs and setae a and c3 in dectes group) are separated by the positions of trapezoidal arrangement or setae a setae wa relative to setae la and ra on tarsi HI, positioned posterior to adanal discs.... 5 the relative development of solenidia

Male with genital discs posterior to gen- 9. Male with epimerites !Va not joining an­ ital arch, often impossible to discern terior to genital arch, with setae a ...... Neodectes, new genus, p. 69 lateral to adanal discs, without meta­ 4. Male with genital discs widely separated podosomal shields. Female with setae from each other and from genital /5 long, either setiform or lanceolate, arch; on Menuridae ...... with terminal filament...... 10 ...... Megalodectes, new genus, p. 71 Male with epimerites IVa forming a mas­ Male with genital discs approximate to sive arch in front of genital apparatus, each other and to genital arch ...... with setae a anterior to adanal discs, ...... Proterothrix Gaud, p. 66 with metapodosomal shields. Female 5. Male with setae a positioned lateral to with setae /5 short and bladelike with-

adanal discs; setae a and c3 in trap­ out terminal filament...... ezoidal arrangement...... 6 ...... Trochilodectes, new genus, p. 73 Male with setae a posterior to adanal 10. Both sexes with epimerites I either Y- or

discs; setae a and c3 in long rectangu- 1t-shaped with posterolateral exten­ lar arrangement...... sions. Male broad, without distinct ...... Dolichodectes, new genus, p. 60 bilobation and without a pronounced 6. Male with genital discs posterior to gen- terminal cleft. Female with terminus ital arch or not evident (i.e., invisible) 7 fused to anterior hysterosoma, with Male with genital discs anterior to gen- pregenital apodeme U- or Q-shaped .... 11 ital arch ...... Pterodectes Robin, p. 54 Both sexes with epimerites I Y-shaped without posterolateral extensions. 7. Both sexes with solenidion cr1 on legs Ill and pai present; epimerites I variously Male narrow, distinctly bilobed with shaped, with or without posterolateral V-shaped cleft. Female with terminus extensions. Male with genital discs freely articulated to anterior hystero­ embedded in heavy sclerotizations soma, with pregenital apodeme al- and often not apparent. Female with most square (Fig. 60) ...... ambulacra of legs Ill-IV equal to or ...... Xynonodectes, new genus, p. 75 smaller than on legs 1-11...... 8 11. Male with terminus broadly arched. Fe­ male with terminal lobes attenuated .... Both sexes with solenidion cr 1 on legs Ill and pai absent; epimerites I Y-shaped ...... Toxerodectes, new genus, p. 75 without posterolateral extensions. Male with terminus weakly bilobed, with Male with genital discs independent small U-shaped cleft. Female wtih ter- of sclerotizations and visible. Female minal lobes abbreviated ...... with ambulacra of legs Ill-IV larger ...... Syntomodectes, new genus, p. 77 than legs 1-11; usually on Nectariniidae The Pterodectes Group .... Anisodiscus Gaud and Mouchet, p. 64 Eight of the twelve pterodectine genera have 8. Both sexes with all dorsal hysterosomal setae wa distant from /a and ra on the anterior setae present (i.e., d1.3 present); setae two pairs of tarsi and have solenidia cr1 smaller / 1 inserted off hysterosomal shield; than solenidia W3 on legs I; additionally, the legs I may be enlarged. Males with eight genera are not known to occur on the terminus distinctly bilobed ...... hummingbirds (Trochilidae). Subdivisions can ...... Montesauria, Oudemans, p. 58 be established for these genera either by the

Both sexes with setae d1.3 or d2 3 absent; positions of the male genital discs in relation to setae 11 inserted at anterolateral an­ the genital arch or by the positions of setae a gles of hysterosomal shield; legs 1-11 in relation to setae C3 and the adanal discs. The subequal. Male terminus truncated, two systems of division are not compatible and without distinct lobes ...... we will discuss only the former as it is the least ...... Pedanodectes, new genus, p. 62 complicated. 54 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

In analgoid protonymphs a pair of genital Montesauria Oudemans, discs is positioned between coxae IV and in the 1905 Neodectes, new genus tritonymphs a second pair of discs is added im­ Pedanodectes, new genus mediately posterior to the protonymphal pair. The Pterodectes subgroup is characterized by Thus, the configuration in the tritonymph is the genital discs anterior to the genital arch and four discs between coxae IV with the discs on in the Monetsauria subgroup these positions are each side of the midline closer to each other reversed. Within the latter complex, the genera than to their opposite members. These relative Anisodiscus and Pedanodectes are closely re­ positions are maintained in adult females, but lated as evidenced by the mutual loss and/or in males, the discs on each side of the idiosoma modifications of specific setae, the overall di­ are arranged side-by-side rather than anterior mensions of the females, and host preferences. and posterior; the two discs on each side may The genus Dolichodectes is unique, having a be connected by a small sclerotization. Con­ high degree of development of the ventral nected or not, the result is four discs arranged shields in the male although the female is not across the idiosoma with the left and right mem­ spectacular. The genus Neodectes, although the bers of each pair adjacent to each other and the discs are posterior to the genital arch, generally two left discs distant from the two right discs. resembles Proterothrix; if the positions of the In most genera the genital discs are near the discs are basic, then this apparent relationship apex of the genital arch. is an excellent example of parallel evolution. Sporadically in the Analgoidea the normal The remaining genera are large and have been positions of the discs are shifted, usually by the studied only enough to suggest that there may discs retaining the tritonymphal positions and be distinct species groups which may be ele­ the male genital structures developing anterior vated to generic rank after additional material or posterior to them. A second type of juxta­ from critical host groups is obtained. positioning is the shifting from the normal ante­ Genus Pterodectes Robin rior position to immediately posterior to the Pterodectes Robin, 1868, Compt. rend. Acad. Sci. Paris, arch with the discs retaining their side-by-side 66(16): 786 (no men nudum) ;-Proctophyllodes (Ptero­ relationship. Additional modifications do occur, dectes) Robin, in Robin and Megnin, 1877, J. Anat. one of which will be demonstrated in the new Physiol., 13: 392; Trouessart, 1885, Bull. Soc. Etud. sci. Angers, 14: 78;-Pterodectes: Oudemans, 1905, Entomol. monotypic genus Megalodectes. Ber., 1(24): 240; Trouessart, 1916, Bull. Soc. zool. The anteriorly positioned discs may become France, 40: 221; Till, 1954, Moc;:ambique doc. trim., (79): 85-6; Gaud, 1957, Bull. Soc. Sci. nat. Phys. Maroc, 37(2): separated from each other and from the genital 108. arch. Within the genus Proterothrix, most spe­ Type species: Proctophyllodes (Pterodectes) cies have the typical disc-genital arch arrange­ rutilus Robin, 1877 (by subsequent designa­ ment, but in a few species one pair of discs is tion). slightly separated from and anterior to the The definition of the genus Pterodectes is second pair. This tendency culminates in Me­ based on nine described and about ninety new galodectes major with the discs being widely species. Within the taxon, the type species is an separated from the genital arch and from each anomaly in respect to certain characters, so other (Fig. 50). much so that it is best to consider the genus as Eliminating Megalodectes, which is the only being composed of two species groups. The pterodectine group occurring on the passeri­ first, the rutilus group, has females with setae /5 form suborder Menurae, the remaining genera almost setiform with a long terminal filament can be divided into two subgroups based on the (Fig. 23), males with a broadly expanded genital positions of the discs relative to the genital arch. arch (Fig. 22), both sexes with setae /1 inserted on the hysterosomal shield, and the hosts are Pterodectes subgroup Montesauria subgroup in the family Hirundinidae. Only P. ruti/us is Pterodectes Robin, 1877 Anisodiscus Gaud & known in the group, but we believe that this Proterothrix Gaud, 1968 Mouchet, 1957 Dolichodectes, new genus species is in actuality a species complex. A GENERIC REVISION OF THE PTERODECTINAE / 55

200µ

Figs. 21-24. Pterodectes rutilus {Robin): dorsal and ventral aspects of male (21, 22) and female (23, 24). 56 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

The second, the graci/is group, has /5 lanceo­ Male and Female late without a terminal filament (as in Fig. 27), 26. Hysterosomal setae absent: none. has a narrower genital arch (as in Figs. 42 and 27. Setae Ii on hysterosomal shields (ruti/us) 62), has setae Ii inserted off the hysterosomal or off. shield, and the hosts are not in the Hirundinidae. 28. Solenidia a1 smaller than wa on legs I. The following definition of the genus includes 29. Setae wa distant from /a and ra on legs 1-11. the above conditions of the characters. 30. Setae cG and mG on legs 1-11 setiform or bladelike. Male 31. Solenidia a1 and setae sR present on 1. Epimerites I V-, U-, or ic-shaped with or legs Ill. without posterolateral extensions. 32. Found on birds of the passeriform subor­ 2. Coxal fields I-IV open. ders Tyranni and Passeres. 3. Legs I-IV subequal. 4. Hysterosomal lobes distinct forming V- The following described species are retained shaped cleft. in the genus Pterodectes; species re-assigned 5. Supranal concavity distinct or indistinct. to the genus are denoted by an asterisk: 6. Metapodosomal shields absent. Pterodectes bilineatus Berla 7. Ventrolateral shields may be present. Pterodectes bilineatus Baria, 1958. Bolm. Mus. Nae. Rio de 8. Pregenital apodeme absent. J., N.S., Zool. (186): 1-3. 9. Genital arch broad, massive (rutilus) or moderately developed (as in Figs. 42 and Pterodectes crassus Trouessart 62) and situated between coxae IV inde­ Proctophyllodes (Pterodectes) crassus Trouessart, 1885, pendent of epimerites IVa. Bull. Soc. Etud. sci. Angers, 14: 79; Pterodectes c.: Canestrini and Kramer, 1899, Tierreich, 7: 125. 10. Genital discs approximate and anterolat- eral to genital arch. Pterodectes gracilis Trouessart 11. Anal shields absent. Proctophyllodes (Pterodectes) gracilis Trouessart, 1885, 12. Adanal discs dentate or edentate. Bull. Soc. Etud. sci. Angers, 14: 79; Pterodectes g.: 13. Setae a lateral or posterolateral to adanal Canestrini and Kramer, 1899, Tierreich, 7: 125. discs. Pterodectes interifolia Trouessart 14. Setae a and ca in trapezoidal arrangement. Pterodectes interifolia Trouessart, 1899, Bull. Soc. Etud. 15. Setae d5 setiform. sci. Angers, 28: 61. 16. Setae pai minute and setiform. 17. Solendia 0 on legs Ill-IV subequal. Pterodectes muticus Banks Pterodectes muticus Banks, 1909, Proc. Entomol. Soc. Female Washington, 11(3): 141. 18. Epimerites I V- or Y-shaped without pos- Pterodectes nordestensis Berla terolateral extensions. Pterodectes nordestensis Baria, 1958. Bolm. Mus. Nae. Rio 19. Legs I-IV subequal. de J., N.S., Zoo I. (186): 4-6. 20. Ambulacra I-IV subequal. 21. Hysterosomal terminus articulated with an­ Pterodectes rutilus Robin terior idiosoma, with or without terminal Pterodectes rutilus Robin, 1868, Comp. rend. Acad. Sci. Paris, 66(16): 786 (nomun nudum); Proctophyllodes appendages. (Pterodectes) rutilus Robin, 1877, in Robin and Megnin, 22. Supranal concavity distinct. 1877, J. Anat. Physiol., 13: 644; Derma/eichus hirundinis 23. Genitocoxal apodemes normal with pregen­ Canestrini, 1878, Atti 1st. veneto Sci., Ser. 5, 5:66; Pterodectes r.: Canestrini, 1886, Prosp. Acarof., 2: ital adopeme Q-shaped. 305-6; Pterodectes rhodesiensis Till, 1954, Mogambique 24. Setae /5 almost setiform with terminal fila­ doc. trim., (79): 90. ment (rutilus) or bladelike without terminal filament. Pterodectes sialiarum (Stoll) Proctophyllodes sialiarum Stoll, 1893, Biol. Cent.-Amer., 25. Solenidia 0 on legs Ill longer than on 3: 42; Pterodectes s: Atyeo and Braasch, 1966, Bull. legs IV. Univ. Nebraska St. Mus., 5: 317. A GENERIC REVISION OF THE PTERODECTINAE / 57

200µ

Figs. 25-28. Montesauria cylindrica (Robin): dorsal and ventral aspects of male (25, 26) and female (27, 28). 58 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

Pterodectes turdinus Berla 5. Supranal concavity distinct and round or Pterodectes turdinus Berla, 1959, Bolm. Mus. Nae. Rio de indistinct. J., N.S., Zoo I. (209): 11. 6. Metapodosomal shields absent. 7. Ventrolateral shields present or absent. Genus Montesauria Oudemans 8. Pregenital apodeme absent. Montesauria Oudemans, 1905, Entomol. Ber., 1 (24): 240; 9. Genital arch expanded, massive, often ir­ Trouessart, 1916, Bull. Soc. zool. France, 40: 221; regular in outline (small in bilobatus group); -Pterodectes: Till, 1954, Mo9ambique doc. trim., (79): 86; Gaud, 1957, Bull. Soc. Sci. nat. Phys. Maroc, 37(2): epimerites IVa may be independent or in­ 108;-Montesauria: Vassilev, 1959, Bulg. Acad. Sci., corporated along anterior margin of arch. Proc. Zool. Inst., 8: 49, 50. 10. Genital discs approximate and posterior to Type species: Proctophyllodes (Pterodectes) genital arch, often not visible as they are cylindricus Robin, 1877 (by original desig­ incorporated in the expanded arch. nation). 11. Anal shields independent or absent. The lack of investigations of feather mites 12. Adanal discs dentate or edentate. during the first half of the twentieth century is 13. Setae a posterolateral, lateral, or anterolat- well demonstrated with the history of this genus. eral to adanal discs.

Oudemans erected Montesauria in 1905, Till 14. Setae a and c3 in trapezoidal arrangement. synonymized the genus with Pterodectes in 1954 15. Setae d 5 setiform to lanceolate. and Vassilev re-established the genus in 1959. 16. Setae pai setiform (spiculiform in amb/y­ However, between 1899 when Trouessart de­ cercus). scribed Pterodectes navicu/a and 1942 when 17. Solenidia

Male 28. Solenidia <11 shorter than w3 on legs I. 1. Epimerites I V-, Y-, or 1£-shaped, with or 29. Setae wa distant from ra and la on legs 1-11. without posterolateral extensions. 30. Setae cG and mG on legs 1-11 setiform or 2. Coxal fields I open or closed, coxal fields spiculiform. II-IV open. 31. Solenidia a1 present on legs 111; setae sR 3. Legs I-IV subequal or legs I enlarged. present or absent on legs Ill. 4. Hysterosomal lobes distinct forming linear, 32. Found on birds of the Passeriformes and V-, or U-shaped cleft. Piciformes (Capitonidae). A GENERIC REVISION OF THE PTERODECTINAE / 59

The following described species are assigned 1877, J. Anat. Physiol., 13: 647; Pterodectes c.: Berlese, to the genus Montesauria; species re-assigned 1886, A.M.S., fasc. 27, no. 9; Pterodectes corvinco/a Oudemans, 1905, Entomol. Ber., 1(23): 225 (new syn­ to the genus are denoted by an asterisk: onymy); Montesauria cy/indricus: Oudemans, 1905, op. cit., 1 (24): 240; M. corvinco/a: Oudemans, 1905, op. cit., Montesauria acotylura (Gaud and Mouchet) * (2)25: 12; P. cy/indricus: Bonnet and Timon-David, 1932, Bull. Soc. Linn. Provences, 5-6: 27; M. cy/indricus: Vas­ Pterodectes acotylurus Gaud and Mouchet, 1957, Ann. silev, 1959, Bulg. Acad. Sci., Proc. Zool. Inst., 8: 49. Parasitol. hum. comp., 32(5-6): 514. Montesauria delicatula (Till)* Montesauria agriocerca (Gaud and Mouchet) * Pterodectes delicatulus Till, 1957, J. Entomol. Soc. S. Pterodectes agriocercus Gaud and Mouchet, 1957, Ann. Africa, 20: 450. Parasitol. hum. comp., 32(5-6): 516. Montesauria dicruri (Gaud and Mouchet) * Montesauria amblycerca (Gaud and Mouchet) * Pterodectes dicruri Gaud and Mouchet, 1957, Ann. Para­ Pterodectes amblycercus Gaud and Mouchet, 1957, Ann. sitol. hum. comp., 32(5-6): 520. Parasitol. hum. comp., 32(5-6): 517-8. Montesauria diplotrema (Gaud and Mouchet) * Montesauria bacillus (Trouessart) * Pterodectes diplotrema Gaud and Mouchet, 1957, Ann. Proctophyl/odes (Pterodectes) bacillus Trouessart, 1885, Parasitol. hum. comp., 32(5-6): 523. Bull. Soc. Etud. sci. Angers, 15: 81; Pterodectes b.: Gaud, 1952, Mem. Inst. sci. Madagascar, Ser. A, 7: 88. Montesauria dispar (Gaud)* Montesauria bilobata (Robin) Pterodectes dispar Gaud, 1953, Ann. Parasitol. hum. comp., 28(3): 202. Pterodectes bilobatus Robin, 1868, Compt. rend., Acad. Sci. Paris, 66(16): 787 (nomen nudum); Proctophyl/odes (Pterodectes) b.: Robin, 1877 in Robin and Megnin, 1877, Montesauria eucyrta (Gaud)* J. Anat. Physiol., 13: 392; Derma/eichus anthi Canestrini, Pterodectes papilla eucyrtus Gaud, 1953, Ann. Parasitol. 1878, Atti R. 1st. Veneto Sci. Let. Arti, Ser. 5, 5: 10; hum. comp., 28(3): 206; P. eucyrtus: Gaud, 1964, Ann. Proctophyllodes alaudae Haller, 1882, Z. wiss. Zool., 36 Mus. roy. Afr. cent., Zool., 80(132): 121. 385, 388 (nomen nudum); Pterodectes b.: Berlese, 1886, A.M.S., fasc. 27, no. 10; Pterocolus b.: Canestrini, 1886, Prosp. Acarof. ital., 2: 296-7; Pterodectes b.: Canestrini, Montesauria eulabis (Buchholz)* 1886, op. cit., 2: 304-5; Proctophyllodes bureschi Vassi­ Dermaleichus eulabis Buchholz, 1869, Bemerk. Gattung lev, 1958, Acad. Sci., Proc. Second Biol. Med. Sci., 4: Dermaleichus, p. 21; Pterocolus e.: Haller, 1878, Zeit. f. 25-30; Pterodectes bureschi: Vassilev, 1959, Compt. Wiss. Zool., 20: 539; Trouessartia e.: Canestrini and rend. Acad. bulg. Sci., 12(3): 244-5. Kramer, 1899, Tierreich, 7: 121. Montesauria brachycaulus (Gaud and Mouchet) * Montesauria eurycalyx (Gaud)* Pterodectes brachycau/us Gaud and Mouchet, 1957, Ann. Parasitol. hum. comp., 32(5-6): 518. Pterodectes eurycalyx Gaud, 1964, Ann. Mus. roy. Afr. cent., Zool., 80(132): 121. Montesauria buphagi (Till)* Montesauria gigas (Gaud and Mouchet) * Pterodectes buphagi Till, 1957, J. Entomol. Soc. S. Africa, 20: 452. Pterodectes gigas Gaud and Mouchet, 1957, Ann. Parasitol. hum. comp., 32(5-6): 523. Montesauria buttikeri (Till)* Montesauria heterocaula (Gaud and Mouchet) * Pterodectes biittikeri Till, 1954, Mogambique doc. trim., (79): 89. Pterodectes heterocaulus Gaud and Mouchet, 1957, Ann. Parasitol. hum. comp., 32(5-6): 526. Montesauria centropa (Gaud and Mouchet) * Montesauria holosticta (Gaud and Mouchet) * Pterodectes centropus Gaud and Mouchet, 1957, Ann. Parasitol. hum. comp., 32(5-6): 519. Pterodectes holostictus Gaud and Mouchet, 1957, Ann. Parasitol. hum. comp., 32(5-6): 527. Montesauria cylindrica (Robin) Pterodectes cylindricus Robin, 1868, Comp. rend. Acad. Montesauria holothyra (Gaud)* Sci. Paris, 66(16): 787 (nomen nudum); Proctophyllodes Pterodectes holothyrus Gaud, 1952, Mem. Inst. sci. Mada­ (Pterode<.tes) cylindricus Robin in Robin and Megnin, gascar, Ser. A, 7(1): 89-90. 60 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

Montesauria hypersticta (Gaud and Mouchet) *, Montesauria reticulifera (Trouessart and new status Neumnan) *, provisional inclusion Pterodectes holostictus hyperstictus Gaud and Mouchet, Pterodectes reticulifer Trouessart and Neumann, 1888, Bull. 1957, Ann. Parasitol. hum. comp., 32(5-6): 527. sci. France Belg., 19: 371.

Montesauria lanceolata (Sugimoto)* Montesauria rosickyi (Cerny)* Pterolichus lanceolatus Sugimoto, 1942, Bull. Sch. Agric. Pterodectes rosickyi Cerny, 1963, Acarologia, 5(4): 649. Forest., Taihoku Imp. Univ., 3:135-6, 147; Pterodectes /.: Gaud and Petitot, 1948, Ann. Parasitol. hum. comp., Montesauria sabiensis (Till)* 23(5-6): 341-2, 343, 346. Pterodectes sabiensis Till, 1954, Moc;:ambique doc. trim., (79): 94. Montesauria leioplax (Gaud and Mouchet) * Pterodectes leioplax Gaud and Mouchet, 1957, Ann. Para­ Montesauria stephanocaula (Gaud)" sitol. hum. comp., 32(5-6): 527. Pterodectes stephanocaulus Gaud, 1953, Ann. Parasitol. hum. comp., 28(5-6): 207. Montesauria listroprocta (Gaud and Mouchet) * Pterodectes listroproctus Gaud and Mouchet, 1957, Ann. Montesauria stictothyra (Gaud)* Parasitol. hum. comp., 32(5-6): 528. Pterodectes papillo var. stictothyrus Gaud, 1953, Ann. Parasitol. hum. comp., 28(5-6): 206; P. s.: Gaud and Montesauria mainati (Trouessart) * Till, 1961, Publ. S. African Inst. Med. Res., 11(L): 256. Proctophyllodes (Pterodectes) mainati Trouessart, 1885, Montesauria synosterna (Gaud and Mouchet) * Bull. Soc. Etud. sci. Angers, 14: 81; Pterodectes m.: Canestrini and Kramer, 1899, Tierreich, 7: 126. Pterodectes synosternus Gaud and Mouchet, 1957, Ann. Parasitol. hum. comp., 32(5-6): 535.

Montesauria merulae (Gaud)* Montesauria trulla (Trouessart) * Pterodectes merulae Gaud, 1957, Bull. Soc. Sci. nat. Proctophyllodes (Pterodectes) trulla Trouessart, 1885, Bull. Maroc, 37: 126, P. (P.) Gaud, 1968, Nat. Hist. Rennell m.: Soc. Etud. sci. Angers, 14: 81; Pterodectes t.: Gaud, Is., Brit. Solomon Isis. 5: 134. 1966, Rev. Zool. Bot. Afr., 73(3-4): 337.

Montesauria navicula (Trouessart) * Montesauria zumpti (Till)* Pterodectes navicula Trouessart, 1899, Bull. Soc. Etud. sci. Pterodectes zumpti Till, 1954, Moc;:ambique doc. trim., Angers, 28: 36. (79): 96.

Montesauria oligosticta (Gaud and Mouchet) * Dolichodectes, new genus Pterodectes oligostictus Gaud and Mauch-et, 1957, Ann. Type species: Proctophyllodes (Pterocolus) ed­ Parasitol. hum. comp., 32(5-6): 531. wardsi Trouessart, 1885. Derivation: Contraction of dolichos, long and Montesauria oxyphylla (Gaud and Petitot) *, Pterodectes. new status The elongated males of this new genus are Pterodectes phyllurus oxyphyllus Gaud and Petitot, 1948, unique in having the anal setae (a) positioned Ann. Parasitol. hum. comp., 23(5-6): 342. posterior to the adanal discs and in having the hysterosomal lobes extended beyond the inser­ Montesauria pachypa (Gaud)* tions of setae d 5 resulting in the dorsal position­ Pterodectes (P.) pachypus Gaud, 1968, Nat. Hist. Rennell ing of these latter setae. The males also have Is., Brit. Solomon Isis. 5: 134-138. considerable portions of the ventral hysterosoma heavily sclerotized, closed coxal fields I-IV, well­ Montesauria papillo (Gaud and Petitot) * developed ventrolateral shields, and the forma­ Pterodectes papillo Gaud and Petitot, 1948, Ann. Parasitol. hum. comp., 23(5-6): 342. tion of an apparent pregenital arch and subgen­ ital shield(s). The latter two structures, the arch Montesauria pardalis (Gaud and Mouchet)* and the subgenital shields, are interpreted as being respectively, thickenings and expansions Pterodectes pardalis Gaud and Mouchet, 1957, Ann. Para­ sitol. hum. comp., 32(5-6): 532. of the surface shields associated with the pos- A GENERIC REVISION OF THE PTERODECTINAE / 61

200µ

cJ' 9

Figs. 29-32. Dolichodectes edwardsi (Trouessart): dorsal and ventral aspects of male (29, 30) and female (31, 32). 62 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM terior epimerites. The coxal fields of the hystero­ 23. Genitocoxal apodemes normal with pregen­ soma are enclosed by a Y-shaped sclerite or ital apodeme Q-shaped. apodeme extending from the genital arch to the 24. Setae /5 lanceolate without terminal fila­ epimerites of coxae Ill. Conversely, the females ment. of this genus are without notable ventral sclero­ 25. Solenidia on legs Ill much longer than l1J tizations; all coxal fields are open and even on legs IV. epimerites I lack posterolateral extensions. The following definition is based on five Male and Female named and two new species: 26. Hysterosomal setae absent: none. 27. Setae '1 inserted on or off humeral shields. Male 28. Solenidia 01 smaller than w3 on legs I. 1. Epimerites I Y-shaped with posterolateral 29. Setae wa distant from ra and la on legs 1-11. extensions. 30. Setae cG and mG on legs 1-11 setiform. 2. Coxal fields I-IV closed. 31. Solenidia 01 and setae sR present on 3. Legs I-IV subequal or legs I and/or IV legs Ill. slightly enlarged. 32. Found on birds of the families Muscicapi­ 4. Hysterosomal lobes elongated forming a dae, Turdidae and Sylviidae (Passeres). deep linear cleft. The following named species are assigned to 5. Supranal concavity usually indistinct. the new genus Dolichodectes; the new combina­ 6. Metapodosomal shields absent. tions are denoted by an asterisk: 7. Ventrolateral shields well developed, often uniting across the venter posterior to ada­ Dolichodectes allocaulus (Gaud and Mouchet) * nal discs, often uniting with surface shields Pterodectes allocau/us Gaud and Mouchet, 1957, Ann. of epimerites IVa. Parasitol. hum. comp., 32(5-6): 517. 8. Pregenital apodeme absent. 9. Genital arch with posterolateral extremities Dolichodectes diplocercus (Gaud and Mouchet)* connected to some portion of surrounding Pterodectes diplocercus Gaud and Mouchet, 1957, Ann. sclerotizations. Parasitol. hum. comp., 32(5-6): 522-3. 10. Genital discs posterior to genital arch and Dolichodectes edwardsi (Trouessart}* often invisible. Proctophyllodes (Pterocolus) edwardsii Trouessart, 1885, 11. Anal shields absent. Bull. Soc. Etud. sci. Angers, 14: 72-3; Pterodectes e.: 12. Adanal discs edentate. Canestrini and Kramer, 1899, Tierreich, 7: 123. 13. Setae a posterior or posterolateral to cen­ ters of adanal discs. Dolichodectes glyphonotus

14. Setae a and c3 in long, almost rectangular (Gaud and Mouchet)* arrangement. Pterodectes g/yphonotus Gaud and Mouchet, 1957, Ann. Parasitol. hum. comp., 32(5-6): 533. 15. Setae d5 lanceolate and inserted on dorsal surfaces of lobes. Dolichodectes platynocercus 16. Setae pai small and setiform. (Gaud and Mouchet) * 17. Solenidia of legs Ill-IV subequal. Pterodectes platynocerus Gaud and Mouchet, 1957, Ann. Parasitol. hum. comp., 32(5-6): 523. Female 18. Epimerites I V-shaped without posterolat- Pedanodectes, new genus eral extensions. Type species: Pterodectes hologaster Gaud, 19. Legs I-IV subequal. 1953. 20. Ambulacra I-IV subequal. Derivation: Contraction of pedanos, short and 21. Hysterosomal terminus articulated with an­ Pterodectes. terior idiosoma and bearing appendages. The new genus shows affinities to the genus 22. Supranal concavity indistinct. Anisodiscus not only in morphological features, A GENERIC REVISION OF THE PTERODECTINAE / 63

200µ

36

Figs. 33-36. Pedanodectes hologaster (Gaud): dorsal and ventral aspects of male (33, 34) and female (35, 36). 64 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

but both groups share a number of hosts. The Male and Female morphological similarities are illustrated by the 26. Hysterosomal setae absent: d1-2 or d1_3. lack of setae d2 in both sexes, indistinct hystero­ 27. Setae Ii inserted on anterolateral angles of somal lobes in the males, and lanceolate setae hysterosomal shield.

/5 and elongated genitocoxal apodemes in the 28. Solenidia a1 smaller than w3 on legs I. females. Species of both genera occur on the 29. Setae wa distant from la and ra on legs 1-11. families Nectariniidae and Sylviidae. 30. Setae cG and mG on legs 1-11 setiform. The following definition is based on three 31. Solenidia <11 and setae sR present on legs named and five new species: 111. 32. Found on birds of the families Nectarini­ Male idae, Pycnonotidae, Dicaeidae, Laniidae 1. Epimerites I Y- or V-shaped with postero- and Sylviidae (Passeres). lateral extensions. The following named species are assigned to 2. Coxal fields I-IV open, rarely I is closed. the new genus Pedanodectes; the new combi­ 3. Legs I-IV subequal. nations are denoted by an asterisk: 4. Hysterosomal lobes weakly developed or absent forming truncated or entire terminus Pedanodectes andrei (Ti II)* without distinct cleft. Pterodectes andrei Till, 1954, Mogambique doc. trim. {79): 5. Supranal concavity indistinct. 86. 6. Metapodosomal shields absent. 7. Ventrolateral shields weakly developed. Pedanodectes hologaster (Gaud)* 8. Pregenital apodeme absent. Pterodectes hologaster Gaud, 1953, Ann. Parasitol. hum. 9. Genital arch with anterior margins fused to comp., 28(3): 204. epimerites IVa; epimerites IVa from each side may or may not connect anterior to Pedanodectes mesocaulus (Gaud and Mouchet) * arch. 10. Genital discs posterior to genital arch and Pterodectes mesocaulus Gaud and Mouchet, 1957, Ann. Parasitol. hum. comp., 32(5-6): 529-31. indistinct. 11. Anal shields absent. Genus Anisodiscus Gaud and Mouchet 12. Adanal discs edentate. Anisodiscus Gaud and Mouchet, 1957, Ann. Parasitol. hum. 13. Setae a posterolateral, lateral, or anterolat- comp., 32(5-6): 502-4; Gaud and Till, 1961, Publ. S. eral to adanal discs. Afr. Inst. Med. Res., 11(L): 246-7. Type species: Pterodectes dolichogaster Gaud, 14. Setae a and c3 in trapezoidal arrangement.

15. Setae d5 setiform or spiculiform. 1953 (by original designation). 16. Setae pai small and setiform. The genus Anisodiscus was erected for a 17. Solenidia

Figs. 37-40. Anisodiscus dolichogaster (Gaud): dorsal and ventral aspects of male (37, 38) and female (39, 40). 66 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

The new definition of the genus is based on Male and Female five named and one new species. 26. Hysterosomal setae absent: pai, d2, and /4• 27. Setae /1 inserted off humeral shields. Male 28. Solenidia a1 smaller than w3 on legs I. 1. Epimerites I V- or Y-shaped without pos­ 29. Setae wa distant from /a and ra on legs 1-11. terolateral extensions. 30. Setae cG and mG on legs 1-11 setiform. 2. Coxal fields 1-11 open, Ill open or closed, IV 31. Solenidia 01 absent and setae sR present closed. on legs Ill. 3. Legs 1-111 subequal, legs IV slightly thick­ 32. Found on birds of the families Nectariniidae ened. and Sylviidae (Passeres). 4. Hysterosomal lobes absent or weakly devel- The following named species are assigned to oped forming shallow U-shaped cleft. the genus Anisodiscus; new combinations are 5. Supranal concavity distinct. denoted by an asterisk: 6. Metapodosomal shields absent. 7. Ventrolateral shields present. Anisodiscus dolichogaster (Gaud) 8. Pregenital apodeme present and connected Pterodectes dolichogaster Gaud, 1953, Ann. Parasitol. hum. to posterior epimerites (absent in mega­ comp., 28(5-6): 202-3; Anisodiscus d.: Gaud and Mou­ chet, 1957, Ann. Parasitol. hum. comp., 32(5-6): 502; discus). A. d.: Gaud and Till, 1961, Publ. S. Afr. Inst. Med. Res., 9. Genital arch well developed and positioned 11(L): 246. between coxae Ill-IV (small in megadiscus and positioned between coxae IV). Anisodiscus eupariphus Gaud and Mouchet 10. Genital discs approximate, visible and posi­ Anisodiscus eupariphus Gaud and Mouchet, 1957, Ann. Parasitol. hum. comp., 32 (5-6): 503. tioned posterior to the genital arch. 11. Anal shields independent and positioned Anisodiscus megacaulus (Trouessart) approximately midway between genital arch Proctophy/lodes (Pterodectes) megacau/us Trouessart, and adanal discs. 1885, Bull. Soc. Etud. sci. Angers, 14: 80; Pterodectes 12. Adanal discs edentate. m.: Canestrini and Kramer, 1899, Tierreich, 7: 125; Anisodiscus m.: Gaud and Mouchet, 1957, Ann. Para­ 13. Setae a distant and anterolateral to adanal sitol. hum. comp., 32(5-6): 504. discs. 14. Setae a and c3 in trapezoidal arrangement. Anisodiscus megadiscus Gaud and Mouchet 15. Setae d5 setiform. Anisodiscus megadiscus Gaud and Mouchet, 1957, Ann. 16. Setae pai absent. Parasitol. hum. comp., 32 (5-6): 504. 17. Solenidia s6 on legs Ill larger than ¢, on Anisodiscus megalurus (Trouessart) * legs IV. Pterodectes megalurus Trouessart, 1899, Bull. Soc. Etud. Female sci. Angers, 28: 37. 18. Epi merites I Y-shaped without posterolat- Genus Proterothrix Gaud, new status eral extensions. Pterodectes (Proterothrix) Gaud, 1968, Nat. Hist. Rennell 19. Legs I-IV subequal. Is., Brit. Solomon lsls., 5: 126. 20. Ambulacra Ill-IV larger than 1-11. Type species: Pterodectes wolffi Gaud, 1962 (by 21. Hysterosomal terminus articulated with an­ original designation). terior idiosoma and bearing appendages. Gaud (1962) first suggested the possibility of 22. Supranal concavity distinct. dividing the genus Pterodectes (s./.) into two 23. Genitocoxal apodemes elongated with pre­ divisions based in part on the positions of setae genital apodeme Q-shaped. a in relation to the adanal discs in the males;

24. Setae /5 lanceolate without terminal fila­ then he said the females would have setae /5 ment. lanceolate with terminal filaments (in /itt.). Later, 25. Solenidia s6 on legs Ill longer than on Gaud (1968) created the taxon Proterothrix as a legs IV. subgenus of Pterodectes and in 'addition to char- A GENERIC REVISION OF THE PTERODECTINAE / 67

200µ

Figs. 41-44. Proterothrix wolffi (Gaud): dorsal and ventral aspects of male (41, 42) and female (43, 44). 68 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM acters mentioned, stated that both sexes had 22. Supranal concavity distinct, round or oval legs I dilated. in shape. Our studies which include the fifteen named 23. Genitocoxal apodemes normal with pregen­ species of Proterothrix and approximately ital apodeme Q-shaped. twenty new species show that legs I may be 24. Setae '5 setiform or lanceolate with terminal dilated and that setae /5 in the females may be filament. setiform with terminal filaments. Additional vari­ 25. Solenidia 0 on legs Ill larger than on IV. ations are seen in the positions of the genital discs in the males; in most species they are Male and Female approximate to each other and to the genital 26. Hysterosomal setae absent: none. arch, but in a few species from Paradisaeidae 27. Setae /1 inserted off or rarely on humeral one pair of discs may be removed a short dis­ shields. tance anterior to the arch (a condition approach­ 28. Solenidia 01 smaller than wa on legs I. ing that of Megalodectes species). 29. Setae wa distant from /a and ra on legs 1-11. 30. Setae cG and mG setiform on legs 1-11. Male 31. Solenidia o1 and setae sR present on legs 1. Epimerites I V-, Y-, or :u-shaped with pos- 111. terolateral extensions. 32. Found on birds of the Coraciiformes, Pici­ 2. Coxal fields I or I and Ill closed. formes, Musophagiformes and Passeri­ 3. Legs I-IV subequal or I dilated. formes. 4. Hysterosomal lobes distinct forming V­ The following named species are assigned to shaped cleft. the genus Proterothrix; new combinations are 5. Supranal concavity distinct, oval or bell- denoted by asterisks: shaped. 6. Metapodosomal shields absent. Proterothrix aculeata (Canes,trini) *, 7. Ventrolateral shields absent. new status, provisional inclusion 8. Pregenital apodeme absent. Proctophyllodes (Pterodectes) mainati var. Trouessart, 9. Genital arch small, between coxae IV and 1885, Bull. Soc. Etud. sci. Angers, 14: 81; Pterodectes independent of epimerites IVa. mainati aculeata Canestrini, in Canestrini and Kramer, 10. Genital discs approximate and anterolateral 1899, Tierreich, 7: 126. to apex of genital arch (rarely one pair of Proterothrix coscinonota Gaud discs positioned slightly anterior). Pterodectes (Proterothrix) coscinonotus Gaud, 1968, Nat. 11. Anal shields independent or fused. Hist. Rennell Is., Brit. Solomon Isis., 5: 126-8. 12. Adanal discs dentate, often similar to Proc- tophyllodes. Proterothrix dicranochaeta Gaud 13. Setae a anteromesal to adanal disc centers. Pterodectes (Proterothrix) dicranochaetus Gaud, 1968, Nat. 14. Setae a and c3 in rectangular arrangement. Hist. Rennell Is., Brit. Solomon Isis., 5: 129-30.

15. Setae d 5 setiform or leaflike. 16. Setae pai setiform and smal I. Proterothrix diminuta {Trouessart) * 17. Solenidia 0 on legs Ill equal to or smaller Pterodectes diminutus Trouessart, 1899, Bull. Soc. Etud. than those of legs IV. sci. Angers, 28: 38; P. phyllurus var. d.: Canestrini and Kramer, 1899, Tierreich, 7: 126. Female Proterothrix emarginata (Trouessart) *, 18. Epimerites I V- or :u--shaped with postero­ new status lateral extensions. 19. Legs I-IV subequal or I dilated. Pterodectes phyllurus emarginatus Trouessart, 1899, Bull. Soc. Etud. sci. Angers, 28: 38. 20. Ambulacra 111-IV subequal or slightly larger than ambulacra 1-11. Proterothrix hymenostoma Gaud 21. Hysterosomal terminus articulated to ante­ Pterodectes (Proterothrix) hymenostomus Gaud, 1968, Nat. rior idiosoma bearing appendages. Hist. Rennell Is., Brit. Solomon Isis., 5: 130-1. A GENERIC REVISION OF THE PTERODECTINAE / 69

Proterothrix modesta (Trouessart) *, Male new status 1. Epimerites I Y- or :u:-shaped with postero­ Pterodectes diminutus modestus Trouessart, 1899, Bull. lateral extensions. Soc. Etud. sci. Angers, 28: 38-9. 2. Coxal fields I and 111 closed or open; coxal fields 11, IV open. Proterothrix paradisiaca (Trouessart) * 3. Legs I-IV subequal. Proctophyllodes (Pterodectes) paradisiacus Trouessart, 4. Hysterosomal lobes distinct forming V­ 1885, Bull. Soc. Etud. sci. Angers, 14: 80; Pterodectes p.: Canestrini and Kramer, 1899, Tierreich, 7: 125. shaped cleft (except manicatus). 5. Supranal concavity distinct, oval or bell- Proterothrix phyllura (Trouessart) * shaped. Pterodectes phyllurus Trouessart, 1899, Bull. Soc. Etud. 6. Metapodosomal shields absent. sci. Angers, 28: 37-8. 7. Ventrolateral shields absent. 8. Pregenital apodeme absent. Proterothrix ranci (Gaud)* 9. Genital arch between coxae IV and inde­ Pterodectes Ranci Gaud, 1952, Mem. Inst. sci. Madagascar, pendent of epimerites IVa. Ser. A, 7(1): 90-1. 10. Genital discs approximate and posterior to Proterothrix sakatai (Sugimoto)* genital arch. 11. Anal shields independent or absent. Proctophyl/odes sakatai Sugimoto, 1940, Bull. Sch. Agric. Forest., Taihoku Imp., Univ., 1: 53; Pterodectes s.: Rad­ 12. Adanal discs dentate. ford, 1953, Parasitol., 43(3,4): 215. 13. Setae a anteromesal to adanal disc centers. 14. Setae a and c3 in rectangular arrangement. Proterothrix schizothyra (Gaud)* 15. Setae d 5 setiform to spiculiform (leaflike in Pterodectes schizothyrus Gaud, 1952, Mem. Inst. sci. securiclatus). Madagascar, Ser. A, 7(1): 91-2. 16. Setae pai small and setiform (spiculiform Proterothrix stenochaeta Gaud in securiclatus). 17. Solenidia 0 on legs Ill-IV subequal. Pterodectes (Proterothrix) stenochaetus Gaud, 1968, Nat. Hist. Rennell Is., Brit. Solomon Isis., 5: 132-4. Female Proterothrix wolffi (Gaud) 18. Epimerites I V- or Y-shaped with or without Pterodectes wolffi Gaud, 1962, Nat. Hist. Rennell Is., Brit. posterolateral extensions. Solomon Isis., 4: 42-4; P. (Proterothrix) w.: Gaud, 1968, op. cit., 5: 134. 19. Legs I-IV subequal. 20. Ambulacra I-IV subequal or Ill-IV slightly Proterothrix xiphiura (Trouessart) * larger than HI. Proctophyllodes (Pterodectes) xiphiurus Trouessart, 1885, 21. Hysterosomal terminus articulated with an­ Bull. Soc. Etud. sci. Angers, 14: 82; Pterodectes x.: terior idiosoma and bearing appendages. Canestrini and Kramer, 1899, Tierreich, 7: 127. 22. Supranal concavity distinct, round or oval in shape. Neodectes, new genus 23. Genitocoxal apodemes elongated with pre­ Type species: Proctophyllodes (Pterodectes) genital apodeme Q-shaped. securiclatus Trouessart and Neumann, 1888. 24. Setae /5 lanceolate with terminal filament. Derivation: Contraction of neos, new and Ptero­ 25. Solenidia 0 on legs Ill longer than 0 on dectes. legs IV. The new genus Neodectes is extremely close to the genus Proterothrix and is distinguished by Male and Female the genital discs posterior to the genital arch 26. Hysterosomal setae absent: none. and by host preference, occurring primarily on 27. Setae Ii inserted on or off humeral shields. the Meliphagidae. 28. Solenidia a1 shorter than w3 on legs I. The definition of the genus is based on two 29. Setae wa distant from ra and /a on legs HI. named and at least five new species. 30. Setae cG and mG setiform on legs 1-1 I. 70 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

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Figs. 45-48. Neodectes securiclatus (Trouessart and Neumann): dorsal and ventral aspects of male (45, 46) and female (47, 48). A GENERIC REVISION OF THE PTERODECTINAE / 71

31. Solenidia o1 and setae sR present on legs 9. Genital arch weakly developed, positioned 111. behind coxae IV; epimerites IVa small. 32. Found on birds of the families Campephagi­ 10. Genital discs distant from each other and dae, Ptilonorhynchidae, Pycnonotidae, Syl­ from genital arch. viidae, and Meliphagidae (Passeres). 11. Anal shields independent. The following species are assigned to the 12. Adanal discs dentate. genus Neodectes; new combinations are de­ 13. Setae a anteromesal to adanal disc centers. noted by asterisks: 14. Setae a and c3 in rectangular arrangement. 15. Setae d 5 setiform. Neodectes manicatus {Trouessart) * 16. Setae pai spiculiform and conspicuous. 17. Solenidia ~ of legs Ill-IV subequal. Proctophyllodes (Pterodectes) manicatus Trouessart, 1885, Bull. Soc. Etud. sci. Angers, 14: 81; Pterodectes m.: Canestrini and Kramer, 1899, Tierreich, 7: 127. Female 18. Epimerites I Y-shaped without posterolat- Neodectes securiclatus eral extensions. (Trouessart and Neumann)* 19. Legs I-IV subequal. Proctophyllodes (Pterodectes) securiclatus Trouessart and 20. Ambulacra I-IV subequal. Neumann, 1888, Bull. Sci. France Belg., 19: 370-1; 21. Hystersomal terminus fused with anterior Pterodectes s.: Vitzthum, 1922, Arch. Naturgesch., A, 88(5): 61-2. idiosoma and bearing appendages. 22. Supranal concavity distinct, round in shape. Megalodectes, new genus 23. Genitocoxal apodemes normal with pregen­ ital apodeme U-shaped. Type species: Proctophyllodes (Pterodectes) 24. Setae /5 lanceolate with terminal filament. major Trouessart, 1885. 25. Solenidia ~ on legs Ill-IV subequal. Derivation: Contraction of megale, large + Pter­ odectes. Male and Female The new monotypic genus contains the largest mites of the Pterodectinae. Close affinities be­ 26. Hysterosomal setae absent: none. tween Mega/odectes and the Proterothrix spe­ 27. Setae /1 not inserted on humeral shields. cies from Paradisaeidae are indicated by setae 28. Solenidia 01 smaller than m3 on legs I. 29. Setae distant from /a and on legs 1-11. pai being spiculiform and setae a anterior to wa ra 30. Setae cG and mG on legs 1-11 spiculiform. the adanal discs in the males and by setae cG on legs I modified as spines in both sexes. 31. Solenidia o1 and setae sR present on legs Unique features include the widely spaced gen­ 111. ital discs and the large posterolateral lamellae 32. Found only on birds of the family Menuridae in the males (Figs. 49, 50). (Passeriformes: Menurae). The definition of the genus is based on the Only the type species is included in this single species. monotypic genus.

Male Megalodectes major {Trouessart) * Proctophyl/odes (Pterodectes) major Trouessart, 1885, 1. Epimerites I Y-shaped without posterolat- Bull. Soc. Etud. sci. Angers, 14: 78-9; Alloptes m.: eral extensions. Canestrini and Kramer, 1899, Tierreich, 7: 115. 2. Coxal fields I-IV open. 3. Legs Ill-IV thickened. The Trochilodectes Group 4. Hysterosomal lobes distinct forming small, This group, characterized by setae wa, ra, and U-shaped cleft. /a approximate on tarsi 1-11 and solenidia o1 5. Supranal concavity distinct, oval in shape. equal to or longer than m3 on legs I, occurs ex­ 6. Metapodosomal shields absent. clusively on the hummingbirds (Trochilidae). 7. Ventrolateral shields well developed. Within the group, which has apparently been 8. Pregenital apodeme absent. associated with these birds for a long period of 72 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

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Figs. 49-52. Megalodectes major (Trouessart): dorsal and ventral aspects of male (49, 50) and female (51, 52). A GENERIC REVISION OF THE PTERODECTINAE / 73 time, little can be said about the intergeneric 10. Genital discs approximate and anterolat- relationships. It is obvious that the genera Tro­ eral to genital arch apex. chilodectes and Xynonodectes are closely re­ 11. Anal shields absent, or present. lated and that the genera Toxerodectes and 12. Adanal discs edentate. Syntomodectes are unique and probably repre­ 13. Setae a lateral or anterolateral to adanal sent separate invasions of feather mites onto discs. the hummingbirds. 14. Setae a and c3 in trapezoidal arrangement. 15. Setae d 5 lanceolate, leaflike, or setiform. Trochilodectes, new genus 16. Setae pai minute and setiform. Type species: Proctophyllodes (Pterodectes) 17. Solenidia ·~ on legs Ill-IV subequal or Ill trochilidarum Trouessart, 1885. slightly shorter than IV. Derivation: Contraction of Trochilus and Ptero­ dectes. Female The males of Trochilodectes are distinguished 18. Epimerites I :u:-shaped with posterolateral by having small metapodosomal shields lateral extensions. to the dorsal hysterosomal shield, epimerites 19. Legs I-IV subequal. IVa connecting anterior to the genital arch, well­ 20. Ambulacra I- IV subequal. developed ventrolateral shields, and coxal fields 21. Hysterosomal terminus articulated with an­ I closed. The males of Xynonodectes lack met­ terior idiosoma and bearing appendages. apodosomal shields, well-developed epimerites 22. Supranal concavity distinct and round in IVa, ventrolateral shields, and have coxal fields I shape. open. Females of the two genera are also easily 23. Genitocoxal apodemes short with pregen­ distinguished by the :u:-shaped epimerites I, Q­ ital apodeme Q-shaped. 24. Setae /5 lanceolate without terminal fila­ shaped pregenital apodeme, and setae /5 without terminal filaments in Trochilodectes and the Y­ ment. shaped epimerites I, angular U-shaped prngeni­ 25. Solenidia ~ on legs Ill-IV subequal. tal apodeme, and setae /5 with terminal filaments in Xynonodectes. Male and Female The definition of the genus is based on two 26. Hysterosomal setae absent: none. named and five new species. 27. Setae '1 inserted on humeral shields. 28. Solenidia 01 subequal to w3 on legs I. Male 29. Setae wa, la, and ra approximate on legs 1. Epimerites I :u:-shaped with posterolateral 1-11. extensions. 30. Setae cG and mG on legs 1-11 setiform. 2. Coxal fields I closed, 11-111 open, and IV 31. Solenidia o1 and setae sR on legs Ill pres­ closed or open. ent. 3. Legs I-IV subequal. 32. Found exclusively on birds of the family 4. Hysterosomal lobes distinct forming U- or Trochilidae (Apodiformes). V-shaped cleft. Two species are assigned to Trochilodectes; 5. Supranal concavity distinct, round or oval the asterisks denote new combinations. in shape. 6. Metapodosomal shields present. Trochilodectes alloptinus {Trouessart) * 7. Ventrolateral shields extending to apex of Proctophyllodes (Pterodectes} alloptinus Trouessart, 1886, terminal cleft. Bull. Soc. Etud. sci. Angers, 16: 149-50; Alloptes a.: 8. Pregenital apodeme absent. Canestrini and Kramer, 1899, Tierreich, 7: 110. 9. Genital arch well developed, between ante­ rior articulations of legs IV or between Trochilodectes trochilidarum {Trouessart)'* coxae Ill-IV; epimerites IVa connecting an­ Proctophyllodes (Pterodectes} trochilidarum Trouessart, 1885, Bull. Soc. Etud. sci. Angers, 14: 82; Pterodectes t.: terior to arch. Canestrini and Kramer, 1899, Tierreich, 7: 127. 74 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

Figs. 53-56. Trochilodectes trochilidarum {Trouessart): dorsal and ventral aspects of male {53, 54) and female {55, 56). A GENERIC REVISION OF THE PTERODECTINAE / 75

Xynonodectes, new genus 23. Genitocoxal apodemes short with pregen­ Type species: Proctophyllodes (Pterodectes) ital apodeme U-shaped with angular cor­ gracilior Trouessart, 1885. ners. Derivation: Contraction of Xynonos, companion 24. Setae /5 narrowly lanceolate with terminal and Pterodectes. filament. The differentiating characters between Xyno­ 25. Solenidia 0 on legs Ill equal to or slightly nodectes and the closely related Trochilodectes longer than those on legs IV. have been stated in the description of the latter Male and Female taxon. The only unique modification of a struc­ 26. Hysterosomal setae absent: /4• ture is the unusual shape of the pregenital apo­ 27. Setae '1 inserted on or near humeral deme of the female. This structure, rather than shields. being Q- or U-shaped, is almost square (Fig. 60). 28. Solenidia a equal to or larger than ms on The definition is based on one named and 1 legs I. one new species. 29. Setae wa, ra, and /a approximate on legs 1-11. Male 30. Setae cG and mG on legs 1-11 setiform. 1. Epimerites I Y-shaped without posterolat- 31. Solenidia a1 and setae sR present on legs eral extensions. 111. 2. Coxal fields I-IV open. 32. Found exclusively on birds of the family 3. Legs I-IV subequal. Trochilidae (Apodiformes). 4. Hysterosomal lobes distinct forming V­ Only one named species is included in this shaped cleft. new genus; the asterisk indicates a new combi­ 5. Supranal concavity distinct, round or oval nation. in shape. 6. Metapodosomal shields absent. Xynonodectes gracilior (Trouessart) * 7. Ventrolateral shields absent. Proctophyllodes (Pterodectes) gracilior Trouessart, 1885, 8. Pregenital apodeme absent. Bull. Soc. Etud. sci. Angers, 14: 80-1; Pterodectes g.: 9. Genital arch weakly developed and posi­ Canestrini and Kramer, 1899, Tiererich, 7: 125. tioned between weakly developed epime­ Toxerodectes, new genus rites IVa. Type species: Pterodectes gladiger hastifolia 10. Genital discs approximate and anterolat- Trouessart, 1899. eral to genital arch. Derivation: Contraction of toxeres, with a bow 11. Anal shields weak, divided. and Pterodectes. 12. Adanal discs edentate. The males of this mite group are unique to 13. Setae a posterolateral to adanal discs. the Pterodectinae-all have the opisthosomata 14. Setae a and c3 in trapezoidal arrangement. broad with sides parallel or divergent behind 15. Setae d 5 setiform. legs IV. The wide terminus of each has a shal­ 16. Setae pai setiform. lowly curved terminal "cleft" into which are 17. Solenidia ~ on legs Ill-IV subequal. directed setae d5, / 5, pai, and pae. The females have two types of termini. One, as illustrated in Female figure 64 has a deep, narrow V-shaped cleft and 18. Epimerites I Y-shaped without posterolat- has the origins of the terminal appendages on eral extensions. the lateral margins of the lobes. The second 19. Legs I-IV subequal. type, in which the cleft is a broad U, has the 20. Ambulacra I-IV subequal. terminal appendages arising from the terminal 21. Hysterosomal terminus articulated with an­ portions of the lobes as is normally observed in terior idiosoma and bearing appendages. the Pterodectinae and Proctophyllodinae. 22. Supranal concavity distinct, round or oval The definition of the new taxon is based on in shape. three named and five new species. 76 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

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Figs. 57-60. Xynonodectes gracilior (Trouessart): dorsal and ventral aspects of male (57, 58) and female (59, 60). A GENERIC REVISION OF THE PTERODECTINAE / 77

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Figs. 61-64. Toxerodectes hastifolia (Trouessart): dorsal and ventral aspects of male (61, 62) and female (63, 64). 78 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

Male 30. Setae cG and mG on legs 1-1 I setiform. 1. Epimerites I 1£- or Y-shaped with postero- 31. Solenidia cr1 and setae sR present on legs lateral extensions. 111. 2. Coxal fields I-IV open, rarely fields I closed. 32. Found exclusively on birds of the family 3. Legs I-IV subequal. Trochilidae (Apodiformes}. 4. Hysterosomal lobes indistinct forming a The following species are assigned to this broad and shallow cleft. new taxon; the asterisks denote the new com­ 5. Supranal concavity distinct, round or oval binations: in shape. 6. Metapodosomal shields absent. Toxerodectes gladiger (Trouessart} * 7. Ventrolateral shields absent. Proctophyllodes (Pterodectes) gladiger Trouessart, 1885, 8. Pregenital apodeme absent. Bull. Soc. Etud. sci. Angers, 14: 82-3; Pterodectes g.: 9. Genital arch moderately developed and po­ Canestrini and Kramer, 1899, Tierreich, 7: 127. sitioned between small epimerites IVa; one species with large pregenital arch situated Toxerodectes gracillimus (Trouessart) * posterior to large anteromesally directed Proctophyllodes (Pterodectes) gracil/imus Trouessart, 1886, Bull. Soc. Etud. sci. Angers, 16: 151; Pterodectes g.: epimerites IVa. Canestrini and Kramer, 1899, Tierreich, 7: 128. 10. Genital discs approximate and positioned anterolateral to genital arch. Toxerodectes hastifolia (Trouessart) *, 11. Anal shields usually absent. new status 12. Adana! discs dentate. Pterodectes gladiger hastifolia Trouessart, 1899, Bull. Soc. 13. Setae a lateral to adanal discs. Etud. sci. Angers, 28: 37.

14. Setae a and c3 in trapezoidal arrangement. Syntomodectes, new genus 15. Setae d5 setiform, spiculiform, or leaflike. 16. Setae pai minute and setiform. Type species: Proctophyllodes (Pterodectes} selenurus Trouessart, 1885. 17. Solenidia ~ of legs Ill-IV subequal. Derivation: Contraction of syntomos, shortened and Pterodectes. Female The last genus is considered to contain one 18. Epimerites I 1£- or Y-shaped with postero- new and one named species. Both contain ex­ lateral extensions. tremely broad mites with reduced lobes in the 19. Legs I-IV subequal. males and females. The males have wide, 20. Ambulacra I-IV subequal. weakly developed terminal lobes, small, often 21. Hysterosomal terminus fused with anterior poorly defined terminal clefts, and anal setae so idiosoma and bearing appendages which distant from the adanal discs that they are sub­ may arise at or lateral to the apices. lateral in position. 22. Supranal concavity distinct, round in shape. The female termini of both species are short­ 23. Genitocoxal apodemes short with pregen­ ened by fusion with the anterior idiosoma and ital apodeme Q-shaped. reduction of the terminal lobes. The female of 24. Setae /5 lanceolate with terminal filament, the type species has uniquely modified hystero­ or setiform. somal lobes and a reduced hysterosomal shield; 25. Solenidia ~ on legs Ill larger than on legs IV. both are considered as species characters. The definition of the genus is based on one Male and Female named and one new species. 26. Hysterosomal setae absent: none. 27. Setae '1 inserted on humeral shields. Male 28. Solenidia cr1 subequal or larger than co3 on 1. Epimerites I 1£-shaped with small postero­ legs I. lateral extensions. 29. Setae wa, la, and ra approximate on legs 2. Coxal fields I-IV open. 1-11. 3. Legs I-IV subequal. A GENERIC REVISION OF THE PTERODECTINAE / 79

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68

Figs. 65-68. Syntomodectes se/enurus (Trouessart): dorsal and ventral aspects of male (65, 66) and female (67, 68). 80 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

4. Hysterosomal lobes indistinct and forming Syntomodectes selenurus {Trouessart} * weakly developed U-shaped cleft. Proctophyllodes (Pterodectes} selenurus Trouessart, 1885, 5. Supranal concavity distinct, round or oval Bull. Soc. Etud. sci. Angers, 14: 84; Pterodectes s.: Canestrini and Kramer, 1899, Tierreich, 7: 127. in shape. 6. Metapodosomal shields absent. SPECIES INCORRECTLY ASSIGNED 7. Ventrolateral shields slightly developed. TO PTERODECTES 8. Pregenital apodeme absent. Proctophyllodes armatus {Banks}, 9. Genital arch well developed and positioned between coxae IV; epimerites IVa well de­ new combination veloped and directed anterior to genital Pterodectes armatus Banks, 1909, Proc. Entomol. Soc. Washington, 11(3): 141; P. a.: Banks, 1915, U.S. Dept. arch. Agric., Office Secy., Rept. (108): 125. 10. Genital discs approximate and anterolat- eral to genital arch apex. Trouessartia rotifer {Trouessart and Neumann}, 11. Anal shields independent. new combination 12. Adanal discs dentate. Proctophyl/odes (Pteroco/us} rotifer Trouessart and Neu­ 13. Setae a lateral to and removed from adanal mann, 1888, Bull. sci. France Belg., 19: 366; Pterodectes r.: Canestrini and Kramer, 1899, Tierreich, 7: 123. discs; positioned sublaterally on venter. Although Oudemans (1905d} and Radford 14. Setae a and c3 in trapezoidal arrangement. (1953} have followed Canestrini and Kramer 15. Setae d 5 setiform. (1899), the species was described in the correct 16. Setae pai small and setiform. taxon, i.e., Trouessartia (=Pterocolus}. 17. Solenidia ~ on legs Ill-IV subequal. UNASSIGNED SPECIES OF PTERODECTES Female 18. Epimerites I u-shaped with small postero- Pterodectes bilaniatus {Trouessart} lateral extensions. Proctophyllodes (Pteroco/us} bilaniatus Trouessart, 1885, 19. Legs I-IV subequal. Bull. Soc. Etud. sci. Angers, 14: 73; Pterodectes b.: Canestrini and Kramer, 1899, Tierreich, 7: 124. 20. Ambulacra I-IV subequal. This species was described from two males 21. Hysterosomal terminus fused with anterior taken from study skins of Protonotaria (=Mnio­ idiosoma; terminal lobes reduced and bear­ tilta} citrea (Boddaert}, 1783 (Parulidae} from ing appendages. the Antilles. The mites are bizarre and without 22. Supranal concavity present, oval or round females, it is impossible to assign the species in shape. to known taxa. The situation is complicated by 23. Genitocoxal apodeme short with pregenital the other mites contained on the type slide, apodeme Q-shaped. namely Allodectes norneri, a species specific to 24. Setae '5 lanceolate with or without a short the Trochilidae. Until we have been able to re­ filament. collect Trouessart's species from either the 25. Solenidia ~ on legs Ill-IV subequal. Parulidae or Trochilidae, it is preferable to main­ Male and Female tain it in an unassigned status. 26. Hysterosomal setae absent: none. Pterodectes intermedius {Trouessart} 27. Setae /1 inserted on or off humeral shields. Proctophyllodes (P.} intermedius Trouessart, 1855, Bull. 28. Solenidia a1 longer than W3 on legs I. Soc. Etud. sci. Angers, 14: 78 (non Proctophyl/odes 29. Setae wa, ra, la approximate on legs 1-11. intermedius Trouessart, 1888 in Poppe, nom. nud.; non 30. Setae cG and mG on legs 1-1 I setiform. Pterodectes intermedius Trouessart and Neumann, 1888}; Pterodectes i. (in part}: Vitzthum, 1922, Arch. 31. Solenidia a1 and setae sR present on legs Naturgesch., A, 88(5): 53-5; P.i.: Atyeo and Braasch, 111. 1966, Bull. Univ. Nebraska St. Mus., 5: 316. 32. Found on birds of the family Trochilidae {Apodiformes}. Pterodectes minor {Berla} At this time, only the type species is assigned Proctophyl/odes minor Berla, 1959, Revta. brasil. Biol., 19(2): 203-4; Pterodectes m.: Atyeo .and Braasch, 1966, to this new genus. Bull. Univ. Nebraska St. Mus., 5: 316. A GENERIC REVISION OF THE PTERODECTINAE / 81

Pterodectes ocelatus Berla ACKNOWLEDGMENTS Pterodectes ocelatus Berla, 1960, An. Acad. Brasil. Cien­ Major research on feather mite systematics cias, 32(1): 102. began at the University of Nebraska in 1960; As explained previously (p. 80), the three spe­ due to personnel transfers, the project shifted cies intermedius, minor, and ocelatus are inter­ to the University of Georgia. All collections ac­ mediate between the two subfamilies. At a future companied us, including the material owned by time, their positions will be critically evaluated. the University of Nebraska. We would like to formally acknowledge this fine institution and Pterodectes phylloproctus Trouessart certain staff members, namely C. Bertrand Proctophyl/odes (Pterodectes) phyl/oproctus Trouessart, 1886, Bull. Soc. Etud. sci. Angers, 16: 150-1; Pterodectes Schultz, Director, and Harvey L. Gunderson, p.: Canestrini and Kramer, 1899, Tierreich, 7: 127-8. Associate Director of the University of Nebraska This species, described from Podargidae, has State Museum, and Roscoe E. Hill, former never been recollected, but from Trouessart's Chairman, Department of Entomology, for the description there is little doubt that the species continuing cooperation given our research pro­ would be assigned to the Pterodectinae. gram for so many years. Our sincere appreciation is expressed to Pterodectes trouessarti Berlese those individuals who have made collections, Pterodectes trouessarti Berlese, 1898, A. M. S., fasc. 88, including type specimens, available for study: no. 8. Edward W. Baker, J. Gaud, L. van der Hammen, The figure of this species has epimerites I free and Max Vachon. Extensive alcoholic collections and divergent, however by adding a transverse were made available by Thomas Aitkin, Trinidad connection between the ends of these struc­ Regional Virus Laboratory; Elliott H. McClure, tures, a typical Trochilodectes species would Migratory Animal Pathological Survey; and result. The drawings and descriptions are based F. K. E. Zumpt, South African Institute for Med­ on slides sent to Berlese by Trouessart and the ical Research. host is listed as Lanius excubitor (Laniidae) We would especially like to thank the curators rather than Trochilidae. However, Trouessart did of the following ornithological collections who considerable collecting from museum study have made facilities available for collecting skins and it is assumed that the host data is ectoparasites from study skins: Academy of incorrect. To date we have been unable to col­ Natural Sciences of Philadelphia, American Mu­ lect this species from either host group. seum of Natural History, British Museum (Nat­ ural History), Chicago Museum of Natural NOMINA NUDA History, Dallas Natural History Museum, Louisi­ Pterodectes rufus Robin ana State University, Michigan State University, Pterodectes rufus Robin, 1877, in Robin and Megnin, 1877, Occidental College, Smithsonian Institution, J. Anat. Physiol., 13: 651. Texas Agricultural and Mechanical University, Pterodectes variolosus Megnin and Trouessart University of Michigan, University of Nebraska State Museum, and the University of Texas Na­ Pterodectes variolosus Megnin and Trouessart, 1884, Bull. Soc. Etud. sci. Angers, 12: 132. tural History Museum. 82 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

REFERENCES FOR AVIAN SYSTEMATICS Mayr, Ernst. 1945. Birds of southwest Pacific. Macmillan Co., New York: 1-316. American Ornithologists' Union. 1961. Check-list of North Mayr, Ernst, and Dean Amadon. 1951. A classification of American birds (5th ed.). Port City Press, Inc., Baltimore: recent birds. Amer. Mus. Nov. (1496): 1-42. 1-691. McClure, H. Elliott. 1963. English vernacular names of the Austin, Oliver L. 1961. Birds of the World. Golden Press, birds of the Malaysian subregion. Malayan Nat. J. 17: New York: 1-316. 75-121. Bannerman, David A. 1953. The birds of West and Equa­ McGregor, Richard C. 1909. A manual of Philippine birds. torial Africa. Oliver and Boyd, London: 1-1526. Bur. Printing, Manila: 1-769. Barrett, Charles. 1947. Australian bird life. Oxford Univ. Munro, George C. 1960. Birds of Hawaii (rev. ed.). Charles Press, Melbourne: 1-239. E. Tuttle Co., Inc., Rutland, Vermont: 1-189. Blake, Emmet Reid. 1953. Birds of Mexico. Univ. Chicago Paynter, Raymond A., Jr. 1955. The ornithogeography of Press: 1-644. the Yucatan Peninsula. Peabody Mus. Nat. Hist. Bull. 9: Bond, James. 1956. Check-list of birds of the West Indies. 1-347. Acad. Nat. Sci., Philadelphia: 1-214. Peters, James Lee. 1931-1969. Check-list of birds of the Cave, Francis 0., and James D. Macdonald. 1955. Birds of world. Vols. 1-7, 9-10, 12, 14-15. Harvard Univ. Press, the Sudan. Oliver and Boyd, London: 1-444. Cambridge. Cayley, Neville W. 1968. What bird is that? A guide to the Peterson, Roger Tory. 1941. A field guide to western birds. birds of Australia (5th ed.). Halstead Press, Sydney: Houghton Mifflin Co., Boston: 1-240. 1-348. ---. 1954. A field guide to the birds of Britain and Cory, C. B., B. Conover, and C. E. Hellmayr. 1918-1949. Europe. Collins, London: 1-318. Catalogue of birds of the Americas. Field Mus. Nat. Hist. ---. 1960. A field guide to the birds of Texas. Hough­ Zool. Ser. 13 (1-11). ton Mifflin Co., Boston: 1-304. Delacour, Jean. 1947. Birds of Malaysia. Macmillan Co., Rand, A. 1936. The distribution and habits of Mada­ New York: 1-382. L. gascar birds. Bull. Amer. Mus. Nat. Hist. 72(5): 143-499. de Schauensee, R. Meyer. 1964. The birds of Colombia Rand, A. and E. Thomas Gilliard. 1968. Handbook of and adjacent areas of South and Central America. Liv­ L., New Guinea birds. Natural History Press, Garden City, ingston Publishing Co., Narberth, Pennsylvania: 1-430. New York: 1-628. Etchecopar, R. D., and Fran~ois Hue, 1964. The birds of Roberts, Austin. 1957. Birds of South Africa (rev. ed.). North Africa from the Canary Islands to the Red Sea. Cape Times Ltd., Cape Town: 1-504. Oliver and Boyd, London: 1-612. Falla, R. A., R. B. Sibson, and E. G. Turbott. 1967. A field Smithe, Frank B. 1966. The birds of Tikal. Natural History guide to the birds of New Zealand and outlying islands. Press, Garden City, New York: 1-350. Houghton Mifflin Co., Boston: 1-254. Smythies, Bertran E. 1953. The birds of Burma. Oliver and Glenister, A. G. 1951. The birds of the Malay Peninsula Boyd, Edinburgh: 1-668. Singapore and Penang. Oxford Univ. Press, London; Taka-Tsukasa, Nobusuke. 1967. The birds of Nippon. Maru­ 1-282. zen Co., Ltd., Tokyo: 1-701. Herklots, G. A. C. 1961. The birds of Trinidad and Tobago. Van Tyne, Josselyn, and Andrew J. Berger. 1961. Funda­ Collins Clear-Type Press, London. 1-287. mentals of ornithology. John Wiley & Sons, Inc., New Johnson, A. W. 1965. The birds of Chile and adjacent York: 1-624. regions of Argentina, Bolivia and Peru. Platt Estableci­ Vaurie, Charles. 1959. The birds of the palearctic fauna. A mientos Graficos, Buenos Aires 1: 1-397; 2: 1-467. systematic reference, order Passeriformes. H. F. & G. Lekagul, Boonsong. 1968. Bird guide of Thailand. Ramin Witherby Ltd., London: 1-762. Press. Bangkok: 1-271. Wetmore, Alexander. 1960. A classification for the birds of Macworth-Praed, C. W., and C. H. B. Grant. 1952. Birds of the World. Smithsonian Misc. Coll. 139(11): 1-37. eastern and northeastern Africa. Longmans, Green and Wildash, Philip. 1968. Birds of South Vietnam. Charles E. Co., London: 1: 1-836; 2: 1-1099. Tuttle Co. Inc., Rutland, Vermont: 1-234. A GENERIC REVISION OF THE PTERODECTINAE / 83

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Les para­ Univ. 1: 40-58. sites et les maladies parasitaires chez l'homme, les ---. 1940b. A list of the Formosan Acarina with refer­ animaux domestiques et les animaux sauvages avec ences. Part I. Families Tyroglyphidae and Analgesidae. lesquels ils peuvent etre en contact. Paris: 136-156. Trans. Nat. Hist. Soc. Formosa 30: 248-258. Megnin, P., and E. L. Trouessart. 1884a. Note sur la mor­ ---. 1942. Studies on the Formosan mites (4th report). phologie et sur la classification des Sarcoptides plumi­ Bull. Sch. Agric. Forest., Taihoku Imperial Univ. 3:124- coles. Bull. Soc. Etud. sci. Angers 12: 131-135. 149. ---,and---. 1884b. Les Sarcoptides plumicoles. J. Till, W. M. 1954. Five new feather mites of the genus Micrograph. 8(2): 92-101. Pterodectes. Moi;:ambique doc. trim. (19): 85-100. Nordberg, Sven. 1936. Biologisch-bkologische untersuch­ ---. 1957. Two new Pterodectes species from passeri­ ungen Ober die Vogelnidicolen. Acta Zool. 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Trouessart, E. L. 1885. Note sur la classification des Anal­ ---. 1958a. Zwei neue Arten Analgesoidea aus Bul­ gesiens et diagnoses d'especes et de genres nouveaux. garien-Proctophy//odes bureschi, Sp. N. und Proctophyl­ Bull. Soc. Etud. sci. Angers 14: 46-91. lodes dontschevi Sp. N. Bulg. Acad. Sci., Proc. Sect. ---. 1886. Diagnoses d'especes nouvelles de Sarcop­ B:01. Med. Sci. 4: 25-30. tides plumicoles (Analgesinae). Bull. Soc. Etud. sci. ---. 1958b. Analgesoidea mites-ectoparasites of birds Angers 16: 85-156. of the Lake Srebrensk near the town of Silestra, Bul­ ---. 1892. Considerations generales sur la classifica­ garia. Akad. nauk U.S.S.R., Zool. Zh. 37(9): 1325-1328. tion des Acariens suivies d'un essai de classification ---. 1959a. Analgesoidea im Gefieder von Rabenvo­ nouvelle. Rev. Sci. nat. Quest, Ser. 2: 20-54. geln der Familia Corvidae in Bulgarian. Bulg. Acad. ---. 1896. Sur la classification des Sarcoptides plumi­ Sci., Proc. Zool. Inst. 8: 45-51. co!es. Bull. Soc. entomol. France 20: 419-422. ---. 1959b. Fur die Wissenschaft neue Analgesoidea ---. 1899. Diagnoses preliminaires d'especes nouvelles aus Bulgarian. Compt. rend. Acad. bulg. Sci. 12(3): d'Acariens plumicoles. Additions et corrections a la 243-245. sousfamille des Analgesines. Bull. Soc. Etud. sci. Angers ---. 1960. Feather mites (Analgesoidea) on some birds 28: 1-62. in Bulgaria. Bulg. Acad. Sci., Proc. Zool. Inst. 9: 431-437. ---. 1916. Revision des genres de la sousfamille des ---. 1962a. Study of the species composition, biology Analgesinae, ou Sarcoptides plumicoles. Bull. Soc. zool. and ecology of feather acari (Analgesoidea) on birds France 40: 207-223. from the environments of Petrich and Gotse Delchev. Trouessart, E. L., and P. Megnin. 1883a. Sur le polymor­ Bulg. Acad. Sci., Nat. Centers Disease in Petrich and phisme sexuel et larvaire des Sarcoptides plumicoles. Gotse Delchev Areas: 141-166. Compt. rend. Acad. Sci. Paris 79: 1319-1321. ---. 1962b. New feather mite (Analgesoidea) for Bul­ ---, and ---. 1883b. Sur la morphologie des Sar­ garian fauna. lzv. Zool. Inst. Sofiya 12: 237-238. coptides plumicoles. Compt. rend. Acad. Sci. Paris 97: 1500-1502. Ventura, Arnold K. 1968. Ectoparasites of Jamacian birds. Caribbean J. Sci. ---, and ---. 1884. Sur la classification des Sar­ 8(3-4): 165-72. coptides plumicoles. Compt. rend. Acad. Sci. Paris 98: Vitzthum, H. Graf. 1921. Acarologische Beobachtungen (5. 155-157. Reihe.). Arch. Naturgesch., A, 87(4): 1-77. Trouessart, E. L., and G. Neumann. 1888. Diagnoses ---. 1922a. Acarologische Beobachtungen. Zoo!. Jahrb. d'especes nouvelles de Sarcoptides plumicoles (Anal­ 44(6): 517-564. gesinae). Bull. sci. France Belg. 19: 325-380. ---. 1922b. Acarologische Beobachtungen. 6. Reihe. Turk, F. A. 1953. A synonymic catalogue of British Acari: Die Gattung Proctophyl/odes Robin 1868. Arch. Natur­ Part II. Ann. Mag. Nat. Hist., Ser. 12; 6(62): 81-5. gesch., A, 88(5): 1-86. Valle, Antonio. 1955. Revisione dell' Acaroteca Canestrini. ---. 1923. Vogel-Acaridae. Rep. Sci. Results Norwe­ 1st. Zool. Anat. Comp. Univ. Padova, 1955: 3-37 (also in: gian Exped. Novaya Zemlya 1921(8): 2-10. Atti Accad. Padova. Mem. Cl. Sci. Mat. e Nat. 67: ---. 1929. Ordnung Milben, Acari. In Tierwelt Mittel­ 67-101). europas, Leipzig 3(7): 1-112. Vassilev, I. D. 1957. Acariens (Analgesoidea) sur les ---. 1942. Acarina. In Bronn, H. G. Klassen und Ord­ plumes des oiseaux en Bulgaria. Compt. rend. Acad. nungen des Tierreichs. Bd. 5, Abt. 4, Buch 5, Lief. 6: bulg. Sci. 10(4): 337-339. 891-898. A GENERIC REVISION OF THE PTERODECTINAE / 87

INDEX acotylura, Montesauria, 59 diminutus, Pterodectes, 68 intermedius, Proctophyllodes, 80 acotylurus, Pterodectes, 59 diminutus modestus, Pterodectes, 69 intermedius, Pterodectes, 52, 80 aculeata, Proterothrix, 68 diplocercus, Dolichodectes, 62 KEY TO GENERA, 52 agriocerca, Montesauria, 59 diplocercus, Pterodectes, 62 lanceolata, Montesauria, 60 agriocercus, Pterodectes, 59 diplotrema, Montesauria, 59 lanceolatus, Pterodectes, 60 alaudae, Proctophyllodes, 59 diplotrema, Pterodectes, 59 lanceolatus, Pterolichus, 60 allocaulus, Dolichodectes, 62 Diproctophyllodes, 51 leioplax, Montesauria, 60 allocaulus, Pterodectes, 62 dispar, Montesauria, 59 leioplax, Pterodectes, 60 Allodectes, 51 dispar, Pterodectes, 59 listroprocta, Montesauria, 60 Alloptes, 49, 52 Dolichodectes, 43, 48, 51, 53, 54, 60 listroproctus, Pterodectes, 60 ALLOPTINAE, 51, 52 dolichogaster, Anisodiscus, 65, 66 mainati, Montesauria, 60 alloptinus, Alloptes, 73 dolichogaster, Pterodectes, 64, 66 mainati, Proctophyllodes, 60 alloptinus, Proctophyllodes, 73 edwardsi, Dolichodectes, 61, 62 mainati, Pterodectes, 48, 60 alloptinus, Pterodectes, 73 edwardsi, Pterodectes, 62 mainati aculeata, Proctophyllodes, 68 alloptinus, Trochilodectes, 73 edwardsii, Proctophyllodes, 60, 62 mainati aculeata, Pterodectes, 68 amblycerca, Montesauria, 59 edwardsii, Pterocolus, 60, 62 major, Alloptes, 71 amblycercus, Pterodectes, 59 emarginata, Proterothrix, 68 major, Megalodectes, 46, 48, 54, 71, andrei, Pedanodectes, 64 emarginatus, Pterodectes, 68 72 andrei, Pterodectes, 64 eucyrta, Montesauria, 59 major, Proctophyllodes, 71 Anisodiscus, 39, 43, 44, 48, 49, 51, eucyrtus, Pterodectes, 59 major, Pterodectes, 71 53, 54, 64 eulabis, Dermaleichus, 59 manicatus, Neodectes, 71 Anisophyllodes, 51 eulabis, Montesauria, 59 manicatus, Proctophyllodes, 71 anthi, Dermaleichus, 59 eulabis, Pterocolus, 59 manicatus, Pterodectes, 71 armatus, Proctophyllodes, 80 eulabis, Trouessartia, 59 megacaulus, Anisodiscus, 45, 66 armatus, Pterodectes, 80 eupariphus, Anisodiscus, 66 megacaulus, Proctophyllodes, 66 bacillus, Montesauria, 59 eurycalyx, Montesauria, 59 megacaulus, Pterodectes, 66 bacillus, Proctophyllodes, 59 eurycalyx, Pterodectes, 59 megadiscus, Anisodiscus, 43, 44, 64, bacillus, Pterodectes, 59 Favettea, 51 66 bilaniatus, Proctophyllodes, 80 gigas, Montesauria, 59 Megalodectes, 43, 51, 43, 54, 68, 71 bilaniatus, Pterocolus, 80 gigas, Pterodectes, 59 megalurus, Anisodiscus, 66 bilaniatus, Pterodectes, 80 gladiger, Proctophyllodes, 78 megalurus, Pterodectes, 66 bilineatus, Pterodectes, 56 gladiger, Pterodectes, 78 merulae, Montesauria, 60 bilobata, Montesauria, 59 gladiger, Toxerodectes, 78 merulae, Pterodectes, 60 bilobatus, Proctophyllodes, 59 gladiger hastifolia, Pterodectes, 75, mesocaulus, Pedanodectes, 64 bilobatus, Pterocolus, 59 78 mesocaulus, Pterodectes, 64 bilobatus, Pterodectes, 59 glyphonotus, Dolichodectes, 62 minor, Proctophyllodes, 80 brachycaula, Montesauria, 59 glyphonotus, Pterodectes, 62 minor, Pterodectes, 52, 80 brachycaulus, Pterodectes, 59 gracilior, Proctophyllodes, 75 modesta, Proterothrix, 69 Bradyphyllodes, 51 gracilior, Pterodectes, 75 Monojoubertia, 51, 52 buphagi, Montesauria, 59 gracilior, Xynonodectes, 75, 76 Montesauria, 39, 43, 44, 49, 51, 53, buphagi, Pterodectes, 59 gracilis, Proctophyllodes, 56 54, 58 bureschi, Proctophyllodes, 59 gracilis, Pterodectes, 56 MORPHOLOGY, 40 bureschi, Pterodectes, 59 gracillimus, Proctophyllodes, 78 Dorsal ldiosoma, 41 butti keri, Montesauria, 59 gracillimus, Pterodectes, 78 Ventral ldiosoma, 41 biittikeri, Pterodectes, 59 gracillimus, Toxerodectes, 78 Male Genital Region, 43 centropa, Montesauria, 59 hastifolia, Toxerodectes, 77, 78 Female Genital Region, 44 centropus, Pterodectes, 59 Hemipterodectes, 51 Legs, 44 corvincola, Montesauria, 59 heterocaula, Montesauria, 59 Chaetotaxy, 44 corvincola, Pterodectes, 59 heterocaulus, Pterodectes, 59 muticus, Pterodectes, 56 coscinonota, Proterothrix, 68 hirundinis, Dermaleichus, 56 navicula, Montesauria, 60 coscinonotus, Pterodectes, 68 hologaster, Pedanodectes, 63, 64 navicula, Pterodectes, 60 crassus, Proctophyllodes, 56 hologaster, Pterodectes, 62, 64 Neodectes, 43, 44, 51, 53, 54, 69 crassus, Pterodectes, 56 holosticta, Montesauria, 59 nordestensis, Pterodectes, 56 cylindrica, Montesauria, 57, 59 holostictus, Pterodectes, 59 norneri, Allodectes, 80 cylindricus, Proctophyllodes, 58,59 holostictus hyperstictus, Pterodectes, norneri, Alloptes, 51 cylindricus, Pterodectes, 59 60 norneri, Proctophyllodes, 51 delicatula, Montesauria, 59 holothyra, Montesauria, 59 Nycteridocaulus, 51 delicatulus, Pterodectes, 59 holothyrus, Pterodectes, 59 ocelatus, Pterodectes, 52, 81 Dermaleichus, 49 HOST-PARASITE RELATIONSHIPS, oligosticta, Montesauria, 60 dicranochaeta, Proterothrix, 68 52 oligostictus, Pterodectes, 60 dicranochaetus, Pterodectes, 68 hymenostoma, Proterothrix, 68 oxyphylla, Montesauria, 60 dicruri, Montesauria, 59 hymenostomus, Pterodectes, 68 pachypa, Montesauria, 60 dicruri, Pterodectes, 59 hypersticta, Montesauria, 60 pachypus, Pterodectes, 60 diminuta, Proterothrix, 68 interifolia, Pterodectes, 56 papilla, Montesauria, 60 88 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

INDEX

papillo, Pterodectes, 60 ranci, Proterothrix, 69 stictothyra, Montesauria, 60 papillo eucyrtus, Pterodectes, 59 ranci, Pterodectes, 69 synosterna, Montesauria, 60 papillo stictothyrus, Pterodectes, 60 reticulifera, Montesauria, 60 synosternus, Pterodectes, 60 paradlsiaca, Proterothrlx, 69 reticulifer, Pterodectes, 60 Syntomodectes, 51, 53, 73, 78 paradisiacus, Proctophyllodes, 69 rhodesiensis, Pterodectes, 56 Tanyphyllodes, 51 paradisiacus, Pterodectes, 69 rosickyi, Montesauria, 60 TAXONOMY, 49 pardalis, Montesauria, 60 rosickyi, Pterodectes, 60 Historical Account, 49 pardalis, Pterodectes, 60 rotifer, Proctophyllodes, 80 Synonymies, 49 Pedanodectes, 43, 48, 51, 53, 54, 62 rotifer, Pterocolus, 80 Deposition of Type Material, 50 Philepittalges, 51 rotifer, Pterodectes, 80 Descriptive Terminology, 50 phylloproctus, Proctophyllodes, 81 rotifer, Trouessartia, 80 Key to Genera, 52 phylloproctus, Pterodectes, 81 rufus, Pterodectes, 81 Toxerodectes, 43, 51, 53, 73, 75 phyllura, Proterothrix, 45, 69 rutilus, Proctophyllodes, 54, 56 trochilidarum, Proctophyllodes, 73 phyllurus, Pterodectes, 69 rutilus, Pterodectes, 54, 55, 56 trochilidarum, Pterodectes, 73 phyllurus diminutus, Pterodectes, 68 sabiensis, Montesauria, 60 trochilidarum, Trochilodectes, 73, 74 phyllurus emarginatus, Pterodectes, sabiensis, Pterodectes, 60 Trochilodectes, 43, 44, 48, 50, 51, 68 sakatai, Proctophyllodes, 69 52, 53, 71, 73, 75, 81 phyllurus oxyphyllus, Pterodectes, 60 sakatai, Proterothrix, 69 trouessarti, Pterodectes, 81 platynocerus, Dolichodectes, 62 sakatai, Pterodectes, 69 Trouessartia, 49 platynocercus, Pterodectes, 62 schizothyra, Proterothrix, 69 TROUESSARTIINAE, 51 Proctophyllodes, 40, 44, 47, 49, 51, schizothyrus, Pterodectes, 69 trulla, Montesauria, 60 54, 68 securiclatus, Neodectes, 70, 71 trulla, Proctophyllodes, 60 PROCTOPHYLLODIDAE, 44, 51 securiclatus, Proctophyllodes, 69, 71 trulla, Pterodectes, 48, 60 PROCTOPHYLLODINAE, 51, 52 securiclatus, Pterodectes, 69, 71 turdinus, Pterodectes, 58 Proterothrix, 39, 43, 44, 48, 49, 51, selenurus, Proctophyllodes, 78, BO variolosus, Pterodectes, 81 52, 53, 54, 66, 68, 69, 71 selenurus, Pterodectes, 78, BO wolffi, Proterothrix, 66, 67, 69 Pterocolus, 49 selenurus, Syntomodectes, 79, BO wolffi, Pterodectes, 66, 69 Pterodectes, 39, 43, 44, 48, 49, 51, sialiarum, Proctophyllodes, 56 xiphiura, Proterothrix, 48, 69 52, 53, 54, 58 sialiarum, Pterodectes, 56 xiphiurus, Proctophyllodes, 69 PTERODECTINAE, 49, 51, 52 stenochaeta, Proterothrix, 69 xiphiurus, Pterodectes,, 69 Pterolichus, 49 stenochaetus, Pterodectes, 69 Xynonodectes, 43, 46, 51, 53, 73, 75 Pteronyssus, 49 stephanocaula, Montesauria, 60 zumpti, Montesauria, 60 Pterophagus, 49 stephanocaulus, Pterodectes, 60 zumpti, Pterodectes, 60 Ptyctophyllodes, 51 THE BOARD OF REGENTS J. G. Elliott, Scottsbluff Kermit R. Hansen, Omaha Robert R. Koefoot, M.D., Grand Island James H. Moylan, Omaha Robert J. Prokop, M.D., Wilber Robert L. Raun, Minden Edward Schwartzkopf, Lincoln Kermit Wagner, Schuyler G. Robert Ross, Lincoln, Corporation Secretary

THE CHANCELLOR D. B. Varner

THE PRESIDENT, UNIVERSITY OF NEBRASKA-LINCOLN Joseph Soshnik