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5-1971

A Revision of the Feather Mite Genus Brephosceles (Proctophyllodidae: Alloptinae)

Paul C. Peterson

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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 4 The University of Nebraska State Museum MAY, 1971

Paul C. Peterson

A Revision of the Feather Mite Genus Brephosceles (Proctophyllodidae: Alloptinae) Paul C. Peterson

A Revi,sion of the Feather Mite Genus Brephosceles (Proctophyllodidae: Alloptinae)

BULLETIN OF The University of Nebraska State Museum VOLUME 9 NUMBER 4 MAY, 1971 BULLETIN OF VOLUME 9, NUMBER 4 THE UNIVERSITY OF NEBRASKA STATE MUSEUM MAY, 1971

Pp. 89-172, Figs. 1-162

ABSTRACT

A Revision of the Feather Mite Genus Brepbosceles (Proctophyllodidae: Alloptinae)

Paul C. Peterson

A systematic revision is presented for the genus 8rephosceles. Data on external morph­ ology, zoogeography, and host-parasite relationships are included. Thirteen named and thirty new species are recognized and described. The new species and the type hosts are: 8rephosceles afribycis, from Afribyx lateralis (Charadriidae); 8. anhimae, from Anhima cor­ nuta (Anhimidae); 8. belonopteri, from 8elonopterus chilensis cayennensis (Charadriidae); 8. bilobatus, from Puffinus puffinus (Procellariidae); 8. chilensis, from 8elonopterus chilensis lampronotus (Charadriidae); 8. collaracus, from Charadrius collaris (Charadriidae); 8. con­ strictus, from Charadrius bicinctus (Charadriidae); 8. diomedei, from Diomedea chrysostoma (Diomedeidae);8. discidicus, from Cygnus bewickii (Anatidae); 8. disjunctus, from Ptero­ droma /eucoptera hypo/euca (Procellariidae); 8. haematopi from Haematopus ostralegus longirostris (Haematopodidae); 8. hoplopteri, from Hoplopterus duvancelii (Charadriidae); 8. inornatus, from Charadrius leschenaultii (Charadriidae); 8. lanceolatus, from Oceano­ droma castro bangsi (Hydrobatidae); 8. /obivanelli, from Lobivanel/us indicus atronuchalis (Charadriidae); 8. longipenis, from Oceanodroma markhami tristrami (Hydrobatidae); 8. /ongistriatus, from Charadrius vociferus ternominatus (Charadriidae); 8. lunatus, from Ptero­ droma externa externa (Procellariidae); 8. pachyptilae, from Pachyptila desolata (Procel­ lariidae); 8. parvatus, from Puffinus pacificus chlororhynchos (Procellariidae); 8. prolatus, from Pachyptila desolata (Procellariidae); 8. pterodromae, from Pterdroma leucoptera hypo­ /euca (Procellariidae); 8. puffini, fromPuffinus gravis (Procellariidae); 8. selenopeltatus, from Puffin us pacificus cuneatus (Procellariidae); 8. stephanibycis, from Stephanibyx melano­ pterus (Charadriidae); 8. marlae, from Puffinus tenuirostris (Procellariidae); 8. triangularis, from Pterodroma cooki defilippiana (Procellariidae); 8. tuberi, from Charadrius alexandrinus seebohmi (Charadriidae); 8. turgidus, from Haematopus palliatus palliatus (Haematopodi­ dae); 8. virginiensis, from Charadrius wilsonia wilsonia (Charadriidae). Peterson1

A Revision of the Feather Mite Genus Brephosceles (Proctophyllodidae: Alloptinae)

INTRODUCTION representing 59 species of birds have contained species of Brephosceles. This study is directed toward the taxonomy, morphology, zoogeogra­ The· genus Brephosce/es Hull with forty-five phy, and host-parasite relationships of species species is one of the largest taxa of the feather of the genus Brephosceles. mite subfamily Alloptinae (Proctophyllodidae). These mites form symbiotic associations with birds in eight non-passeriform families repre­ senting five avian orders. Each of these orders COLLECTION AND PREPARATION may support one or more of six morphologically OF MATERIAL distinct mite species groups; however, each Specimens for this study were acquired by mite group is restricted to a single order of birds field collections, loans, and examination of mu­ (one exception). In the majority, one host-one seum study skins. Field collected birds were parasite relationships are evidenced for the examined with a dissecting microscope to gain Brephosceles species. experience for judging the validity of infesta­ Prior to the present study and that of Peterson tions found on museum study skins. If consider­ and Atyeo (1968), only eleven species of Breph­ able numbers of mites were present (50+), the osceles were known and information concerning birds were washed in water with a mild deter­ host-parasite relationships and zoogeography gent and the decanted washings were preserved was limited. During this and related studies, over in 70 per cent ethyl alcohol for future examina­ 18 000 birds or bird skins representing every tion. bi;d family have been examined for feather The most efficient and profitable method of mites. Additionally, thousands of vials and slides collecting is the examination of bird study skins of feather mites have been received from col­ in museum collections. In contradistinction to lectors throughout the world. Against this back­ blood-feeding mites, feather mites are seden­ ground of material, 370 individual collections tary even after death of the bird. As a result, the ~I C. Peterson: Assistant Curator of Entomology, mites can be seen in position along the barbs University of Nebraska State Museum, and lnstruct~r, De­ partment of Entomology, University of Nebraska, Lincoln, of the feathers; in this way accurate host rec­ Nebraska. ords are obtained and contamination is minimal. Present Position: Assistant Professor of Biology, Depart­ Through field experience, infestations of these ment of Biology, Youngstown State University, Youn~sto~n, mites can accurately be assessed by the use of Ohio, and Research Associate in Entomology, Umvers1ty of Nebraska State Museum, Lincoln, Nebraska. a dissecting microscope. The mites can be 89 90 I BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM scraped from the feather and transferred to vials sexes and consequently is of little diagnostic containing 70 per cent ethyl alcohol. Collecting value. Typically hysterosomal shields are illus­ data are recorded and cross referenced with a trated in Figures 1 and 2; in a few species the host card file. males may have the shields reduced in size Mite specimens are rehydrated and cleared and/or divided by a mid-dorsal transverse mem­ in lactophenol at 93°C. for ten minutes, then branous area or suture. The most meaningful mounted in Hoyer's mounting medium. Freshly morphological differences among the various mounted specimens are placed in a drying oven groups and between sexes are found in the at 50°C. for three days after which the cover terminal structures of the hysterosoma. slips are sealed with a commercial ringing com­ The bifurcate male terminus or lobar region pound. can be considered as two posterior prolonga­ tions of the hysterosoma or conversely, the terminal portions of the idiosoma separated by a deep cleft. The anterodorsal surface of each MORPHOLOGY lobe may be continuous with the dorsal surface In contrast to the relatively simple body con­ of the anterior hysterosoma (Fig. 9) or the sur­ formation in many proctophyllodid genera, the faces may be partially (Fig. 13) or completely males of the genus Brephosceles are distinct. (Fig. 33) separated by a membranous area or The hysterosomal terminus is divided into two suture. The length of the lobes is subject to con­ well-developed lobes with a repositioning of siderable variation either between or within the specific setae. The female configuration is com­ species groups. Each lobe bears a series of paratively simple; the hysterosoma is elongated, marginal lamellae. The inner margin has one either bilobate or entire, and the terminal lobes (rarely two) inter/obar Jame/Jae (IL) which may may bear well-developed lamellae. or may not connect anteriorly to the lamella of It is not the intent in this section to present a the opposite side. The outer margin may have an detailed morphological analysis of each species extralobar Jamel/a (EL) extending for the entire group, but rather to identify those features which length of the lobe, restricted to the antero- or are most important in species differentiation. posterolateral surface or absent. The apex of Drawings of a hypothetical male and female each lobe has one or two small postlobar Jame/­ Brephosceles species are presented for illustra­ Jae (PL); when two are present, the mesal mem­ tive purposes (Figs. 1-4). ber is the largest and the lateral lamella is ventral to the largest terminal seta. The shapes Gnathosoma of the postlobar lamellae vary and the margins The adult gnathosoma of species in Breph­ are classified as either entire or dentate. osceles and related alloptine genera provides The hysterosomae of the females usually ter­ important diagonstic characters. In contrast to minate in two distinct lobes which may or may the Proctophyllodinae, the gnathosoma of the not be extended by terminal lamellae. The hys­ Alloptinae is much longer than wide and the terosomal terminus is classified as entire, weakly movable digits extend beyond the apices of the or distinctly bilobate; if bilobed the interlobar pedipalpi. In the Brephosceles marginiventris cleft (JC) separating them is subject to consider­ group the movable digits are greatly lengthened, able variation in size and may support a hyaline spoon-shaped, and have marginal serrations. interlobar Jamel/a (IL). Continuous with the in­ Other than the cited variations, the mouth parts terlobar cleft, a supranal cleft (SaCI) usually are similar to those in the genus Proctophy/­ extends anteriorly along the midline connecting lodes as described by D. E. Johnston (Atyeo a shallow depression, the supranal concavity and Braasch, 1966). (SaCo); the latter supports the copulatory pore. ldiosoma Often the supranal cleft and supranal concavity Dorsal Jdiosoma. The propodosoma is gener­ are difficult to delimit and either may be present ally uniform within a species group and between or absent according to the species group. The A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 91 external opening of the primary spermathecal nizable patterns in both sexes. Each epimerite duct is positioned in the supranal concavity or has an anterior and a posterior branch and terminates on the interlobar lamella. these delimit the coxal fields. Additionally, the Ventral /diosoma. The structures of the epim­ epimerites may support small integumental erites, the shape of the pregenital apodeme, sclerotizations (surface fields) of varying shapes the position of setae and the development of which are characterized as well, moderately, or the genital organ, form the major diagnostic fea­ poorly developed. In Brephosceles, epimerites I tures in the genus Brephosce/es. Collectively, are without exception fused medially in a Y and these characters define six morphologically dis­ coxal fields I and II are always open. The termi­ tinct species groups. nology of these structures follows Atyeo and Structurally, the epimerites form easily recog- Braasch (1966).

d'

h

---·······-PO re ✓-='"--H+---HCI

2 Figs. 1 and 2. A hypothetical Brephosceles sp., male (1), female (2), dorsal aspects of idiosomae. Setal designations: d,-d, dorsal; h, humeral; /,-/,, lateral; pae, postanal external; pai, postanal internal; see, external scapular; sh, subhu­ meral;sci, internal scapular; ve, external vertical; vi, internal vertical; HCI, hysterosomal cleft; IC, interlobar cleft; Sa­ C/, supranal cleft; Saco, supranal concavity. 92 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

cf

3 A Figs. 3 and 4. A hypothetical Brephosceles sp., male (3), female (4), ventral aspects of idiosomae. Seta! designations: a, adanal; Ci-Ca, central; cxa, coxal; s, sternal. A, anus; AD, adanal discs; EL, extralobar lamellae; Ep1-Ep,, epimerites; GD, genital discs; GO, genital organ; IL, interlobar lamellae; PaA, preanal apodeme; PgA, pregenital apodeme; PL, postlobar lamellae; Sgs, subgenital shield.

In the male, epimerites associated with legs tions are simple and similar to those of Procto­ Ill (EPa) and IV (Ep4) are variable in shape and phyl/odes species described by Atyeo and position. In some species epimerites Ill are co­ Braasch (1966). alesced with the humeral shields, in others inde­ Posteriorly there are two specialized cup pendent. Epimerites IV (Ep4) and the posterior shaped structures, the adanal discs and their branch of epimerites Ill (Ep3a) coalese and an­ associated adanal shields. In Brephosce/es the teriorly may fuse or nearly fuse with the anterior disc cylinder is very short, the longitudinal axis branch of epimerites Ill forming closed coxal being shorter than the transverse axis. The cor­ field Ill. Epimerites IVa and the pregenital apo­ ona is involuted and attached securely by folds deme (PgA) are often fused with epimerites IV, of the integument. According to Popp (1967) forming closed coxal field IV. In the majority of these accessory genital organs operate as a species, however, the coxosternal configura- piston and cylinder in a vacuum pump and hold A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 93 the opisthodorsal region of the female trito­ small sclerites supports the exterior integumen­ nymph during copulation. Additionally, an ad­ tal folds. Two pairs of atrophied genital discs hesive substance, produced by a pair of special are positioned near the posterolateral extremi­ glands near the discs and secreted through ties of the pregenital apodeme, either anterior them, smooths the corrugated dorsal integument or posterior to setae C2 depending on the spe­ of the tritonymphal instar. cies group. Many species are characterized in part by The spermatheca per se is a sac-like structure the presence or absence or a preanal apodeme located at the anterior end of a long primary (PaA). This apodeme is usually small and spermathecal duct which opens posteriorly to rounded in the males, and large and crescentric the outside of the body. The primary duct can in the females (Fig. 4). In the males the preanal be expanded apically (Fig. 144), tapering distally apodeme may be fused with the subgenital (Fig. 142), or simple and tubular (Fig. 156). shield. Closely associated with the spermathecal head Male Genital Region. The male genitalia and is a pair of secondary ducts which lead to the their relationships with other components of the ovaries. The secondary ducts vary considerably ventral hysterosoma supply important criteria in in their origins and lengths and in some cases the differentiation of both species and species from the uniform tubular configuration. groups. Although differing considerably between species groups, the genitalia are usually located ldiosomal Chaetotaxy at the level of the articulations of legs Ill or IV. The nomenclatorial system for idiosomal The genital organ varies considerably in chaetotaxy was developed by Atyeo and Gaud length and width. It is supported anteriorly and (1966) and has been used successfully for many laterally by a prominent genital arch (Fig. 3) genera of sarcoptiform feather mites. The sys­ which may or may not be connected to the inner tem is applied here to Brephosceles. margin of the pregenital apodeme (PgA). Di­ Dorsal ldiosoma (Figs. 1 and 2). On the propo­ rectly beneath the genital organ is a small sub­ dosomal shield, the external scapular setae (se) genital shield (SgS) of variable shape and size are long and conspicuous and approximate to which may or may not bear setae C2, the small internal scapular setae (si). Near the Species groups are based primarily on the anterior margin of the shield is a small unpaired development of the pregenital apodeme (PgA) microseta, the internal vertical seta (vi). A pair in the male. This structure may have the follow­ of external vertical setae (ve) may be present ing shapes: inverted V (group II, Fig. 15), open V or absent; if present, they are small and often (group 111, Fig. 75), open but fused with epim­ difficult to observe. erites IV forming closed coxal field Ill and/or Anterolaterally, each humeral shield supports IV (groups IV and V, Figs. 95, 107), trapezoid three setae: the long humeral seta (h}, the and open (group VI, Fig. 120), and trapezoidal shorter, variously shaped subhumeral seta (sh), and fused (group VI, Fig. 115). Only in groups and the first lateral seta (/1). In the genus Breph­ IV, V, and VI are the posterior limits of the gen­ osceles the seta sh is positioned either at the ital arch fused to the inner margin of the pre­ same level or posterior to seta h. Seta '1 is usu­ genital apodeme. Located on the pregenital ally positioned on the extreme anteromedial apodeme, anterior to the genital organ are two margin of the humeral shield. Structurally seta pairs of atrophied genital discs. 11 and sh may be setiform, spiculiform, lanceo­ Female Genital Region. The female genital late or bluntly rounded. region is circumscribed by a well-developed Theoretically, the hysterosoma bears five pregenital apodeme (PgA, Fig. 4). The apodeme horizontal (transverse) rows of setae, each row may be crescentric or fused posteriorly with consisting of a dorsal pair (d) and a lateral pair epimerites IV in the shape of a Moresque arch. (/) designated as d1-5 and 11_5 respectively. Setae Posterior to the pregenital apodeme, the genital d1 are absent in the genus Brephosceles. In opening appears as an inverted V. A pair of those genera or life stages with a simple body 94 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM conformation these rows are easily delimited. at the level of the articulations of legs Ill and IV However, due to the elongation of the hystero­ respectively. On the opisthosoma anterior to the somal lobes in Brephosce/es males, there has terminal lobes, setae d4 and /3 are usually posi­ been a corresponding shift in positions of vari­ tioned in a relatively straight line; setae d3 are ous setae. In addition to dorsal and lateral setae, anterior to d 4 at variable distances. Setae d5 on two pairs of setae, the internal postanals (pai) the terminus of the lobe and setae h on the and external postanals (pae), are apparent. lateral margin are the largest and most conspic­ These postanal setae are differently positioned uous setae on the hysterosoma. The small, often in the two sexes. In the male, the internal post­ inconspicuous setae /4 are located anterior to anal setae are inserted on the mesal margin of setae /5• the hysterosomal lobes, immediately posterior The internal postanal setae (pai) are minute to setae d3• The external postanal setae are and inserted on the dorsal hysterosomal lobes, always positioned on the postlamellae, approxi­ anteromesal to setae d5• The postanal external mate to setae h. In the female, the internal post­ setae (pae) are ventral in position and due to anal setae are positioned on the internal margins the striations on the venter, extremely difficult of the terminal cleft formed by the lobes. The to observe. external postanal setae are inserted ventral to Ventral ldiosoma (Figs. 3 and 4). There are setae a. six pairs of ventral setae; the sternal setae (s), Considering the portion of the male hystero­ the coxal setae (CX3), three pairs of central setae soma anterior to the level of the lobes, there (c1_3) and one pair of adanal setae (a). The post­ are only three pairs of setae (d2, Ii, /2) which anal setae which have secondarily shifted (ex­ represent the first and second transverse rows cept pae), the sternal, coxal, and adanal setae of dorsohysterosomal setae. Setae d2 are always are constant in position and are respectively microsetae, while setae /2 may be elongate. The located between epimerites I and II, between only remaining structure on the anterior portion epimerites Ill and IV, and lateral to the anal of the hysterosomal shield is a small pore lo­ opening. cated anterior to hysterosomal cleft (between In the males, setae c1 are located anterior to setae 12 and /3). the genital apparatus at approximately the level The remaining setae (d3_5 and /3_5) are vari­ of the subhumeral setae (sh). Setae C2 are pos­ ously positioned on the hysterosomal terminus terior to the genital apparatus and may or may according to species or species group. In some not be associated with the subgenital shield. species, setae d3 are positioned on the anterior Setae c3 are found in the region of coxal field IV, one-third of the lobe and setae d4 approximate lateral to the genital apparatus. In the females, to the internal postanal setae (pai). Setae d3 setae c1 and C2 are in close proximity to the may also be positioned at the midlength of the pregenital apodeme whereas setae Cs are pos­ lobe; if this occurs setae d4 are anterior to setae terior, at approximately the level of trochan­ pai. Setae d5 and the external postanal setae ters IV. (pae) are variable in length and always associ­ ated with the postlobar and extralobar lamellae Leg Morphology Including Chaetotaxy respectively. The external postanal setae are Typically each leg of an analgoid mite is positioned immediately ventral to setae /4. The comprised of seven distinct segments: coxa, tro­ largest and most conspicuous setae (/5) are ap­ chanter, femur, genu, tibia, tarsus, and pretar­ proximate to setae /4 which are variable in shape sus. The coxae are incorporated into the ventral and length. Setae /3 are usually positioned on and ventrolateral surfaces of the idiosoma; rem­ the outer margin, at midlength of lobe. nants form partially sclerotized epimera which In the females, the setal rows are easily dis­ deliniate the coxal fields. tinguished. As setae d1 are absent, the hystero­ Leg morphology and leg chaetotaxy have soma bears four pair of dorsal setae and five played an integral part in defining genera and pair of lateral setae. Setae d2 and /2 are located species within the Alloptinae and in differentiat- A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 95

5

II 6

vF mG

111 7

IV 8

Figs. 5-8. Brephosce/es charadrii Dubinin, male, antiaxial (postaxial) aspects• of leg I (5); leg II (6); leg Ill (7); leg IV (8). 96 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM ing the subfamilies of the Proctophyllodidae. Leg Ill The Alloptinae are separated from the related trochanter ...... 0 ...... 0 ...... 1 ...... 1 Proctophyllodinae and Trouessartinae by the femur ...... O ...... 0 ...... 0 ...... 0 genu ...... (1) ...... (1) ...... (1) ...... (1) reduced seta a2 on genu II, the shape of the tibia ...... 1 ...... (1) ...... 1(1) ...... 1(1) or (1) apotele (an inverted T or transverse bar), and tarsus ...... 4 ...... 4 ...... 5 ...... 5 the fusion of the femur and genu. This partial fusion of the femur and genu can be observed Leg IV from the postaxial aspect of legs I-IV (see Figs. trochanter - ...... 0 ...... 0 ...... 0 5-8). The tibia and tarsus in adult Brephosce/es femur ...... - ...... 0 ...... 0 ...... O species are equal both in length and width. The genu ...... - ...... 0 ...... O ...... O pretarsus is expanded to form a bell-like or tibia ...... - ...... 0 ...... (1) ...... (1) ovoid ambulacrum which may or may not have tarsus ...... - ...... 3 ...... 5 ...... 5 () = solenidia an apical point. The leg chaetotaxy of Brephosceles species nearly parallels Proctophyllodes (Atyeo and Braasch, 1966). Figures 5-8 position and name the setae in the alloptine genus Brephosceles HOST-PARASITE RELATIONSHIPS using the nomenclature of Atyeo and Gaud Alloptine mites are commonly found in tan­ (1966). The adult and tritonymph bear the same dem between the barbs of the primary and sec­ compliment of setae and solenidia with the ex­ ondary remiges of the wings and the rectrices ception of tibia Ill in which setae kT may be of the tail. Growing evidence suggests that present or absent. The corresponding larva and Brephosceles species may occupy a specific protonymph lack setae sR on the trochanters ecological niche. Dubinin (1949, 1951) reported and omega3 on tarsus I. On genu II a solenidian that the secondary feathers of the wings are the appears in the protonymphal stage with a corre­ primary loci during moderate and light infes­ sponding increase of one tactile seta on tarsus tations. In heavy infestations the mites are Ill. Leg IV of the protonymph has only three distributed throughout the wings and tail. Spe­ setae: d, r, and w. The adult bears the normal cific biological and ecological data-ecological full compliment of solenidia and tactile setae. niches, food preferences, temperature and hu­ The following table summarizes the number of midity tolerances-are requisite. Hopefully with tactile setae and solenidia applicable to a par­ renewed interest in feather mites, investigations ticular life stage. Numbers not enclosed within of these phenomena will be conducted. parenthesis indicate numbers of tactile setae; The present study cites thirty-four species of numbers enclosed by parenthesis indicate the Brephosceles (75 per cent) which are known number of solenidia. The femur and genu are from only a single bird species, ten (22 per cent) treated as distinct segments. which occur on more than one species in the TABLE I same genus, and two species which inhabit two Developmental Leg Chaetotaxy or more families of birds. The details of these Leg I associations using the arbitrary categories of larva protonymph tritonymph adult Atyeo and Braasch (1966) for host-parasite rela­ trochanter ...... O ...... 0 ...... 1 ...... 1 femur ...... 1 ...... 1 ...... 1 ...... 1 tionships are discussed below. genu ...... 2(1) ...... 2(1) ...... 2(1) ...... 2(1) In Brephosceles, there are six morphologically tibia ...... 1 (1) ...... 1 (1) ...... 1 (1) ...... 1 (1) distinct mite species groups; each mite group is tarsus ...... 7(1 ) ...... 7(1 ) ...... 7(2) ...... 7(2) restricted to a single avian order (except the two Leg II species currently assigned to group V). Types of trochanter ...... 0 ...... O ...... 1 ...... 1 host specificities within an avian order vary but femur ...... 1 ...... 1 ...... 1 ...... 1 generally they are one parasite-one host associ­ genu ...... 2 ...... 2(1) ...... 2(1) ...... 2(1) tibia ...... •...... 1(1) ...... 1(1) ...... 1(1) ...... 1(1) ations. The six species groups of the genus tarsus ...... 7(1) ...... 7(1) ...... 7(1) ...... 7(1) Brephosceles and their hosts are: A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 97

Group I (Brephosceles forficiger} ger, suggests support of Storer's tentative Gaviiformes conclusions. Gaviidae Group II. Members of this group are restricted to two familes of the Procellariiformes. With Group II (the decapus group} three exceptions, these mites are host specific Procellariiformes at the species level as indicated below. The Hydrobatidae exceptions are host specific at the genus level. Procellariidae Hydrobatidae Group Ill (the marginiventris group} Hydrobates pelagicus Brephosce/es pe/agicus Procellariiformes Vitzthum, 1921 Diomedeidae Oceanodroma castro Brephosceles decapus Hydrobatidae (Gaud) 1953 Procellariidae Oceanodroma castro /eucorhoa Group IV (the anatina group} Oceanodroma markhami Brephosce/es /ongipenis, Anseriformes tristrami new species Anatidae Procellariidae Anhimidae Pachyptila desolata Brephosceles pachyptilae, Group V (the geranoxenus group} new species Charadriiformes Brephosce/es prolatus, new species Burhinidae Pterodroma leucoptera Brephosce/es pterodromae, Gruiformes hypoceuca new species Gruidae Pterodroma externa Brephosceles lunatus, cervicalis new species Group VI (the charadrii group} Pterodroma externa Charadriiformes externa Charadriidae Pterodroma cookii Brephosceles triangularis, defilippiana new species Group I currently contains one species, Puffinus tenuirostris Brephosceles superbus Brephosceles forficiger (Megnin and Trouessart} Dubinin, 1952 from the loons Gavia immer and G. arctica. The Brephosce/es marlae new species species is known from one male and two fe­ males. With the exception of Dubinin's record Group Ill. In contradistinction to strict species of a mite resembling B. forficiger from two arctic specificity in group II, the species included in loons, this species has never been recollected group Ill are primarily host specific at the genus from the type host. Thus it must be assumed level and occur on three families, Procellariidae, the description is based on a small and re­ Hydrobatidae, and Diomedeidae. The broadest stricted population or an incorrect host identi­ host range in this group is exemplified by fication. Brephosce/es disjunctus known from four of Assuming the host determination is correct, twenty-three species of Pterodroma and B. par­ Brephosceles forficiger exemplifies a situation vatus from four of eighteen species of Puffinus. in which acarine taxonomy may complement re­ Extensive collecting from related bird species cent avian systematics. Storer (1956), utilizing will undoubtedly extend the host ranges of these comparative morphology and behavior, tenta­ and other included species within the group. If tively places the loons in Charadriiformes. Gavia two or more species of Brephosceles occur on is not closely related to Hesperornis as previ­ the same host species or the same individual, ously thought, but remains in primitive posi­ then it is probable that discrete ecological tion by its bionomics. Based on comparative niches are involved. As a case in point, Breph­ morphological characters which are shared by osceles bi/obatus from Puffinus puffinus is com­ feather mites of the Gaviiformes and Charadri­ monly found in association with B. parvatus from iformes (group VI}, the mite species, B. forfici- the same bird species. 98 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

The only example of interfamilial host speci­ Nettapus auritus Brephosce/es lambda ficity in Brephosceles is evidenced by B. mar­ (Trouessart) 1885 giniventris. This species occurs on the bird Anhimidae genera Diomedea (Diomedeidae) and Macron­ Anhima cornuta Brephosce/es anhimae, ectes (Procellariidae) .The host-parasite rela­ new species tionship is not surprising since the ranges of the Group V. This seemingly artificial group is two bird species overlap considerably and represented by two species, each known from breeding areas are restricted: this may increase separate orders. Brephosceles chaubaudi has the chances for repopulation on another host. a particularly broad host preference and can be The avian hosts and their respective parasites considered as host specific to the family Bur­ are listed below. hinidae (Charadriiformes). The mite species is Diomedeidae recorded from representatives of the three in­ Diomedea chrysostoma Brephosceles diomedei, cluded bird genera, Burhinus, Orthorhamphus, new species and Esacus. In contrast, Brephosceles geranox­ Diomedea exulans enus is known from one host species, Balearica Diomedea melanophris pavonina regu/orum (Gruiformes). To date, this melanophris Diomedea chlororhyn­ Brephosceles marginiventris is the only example of a species group occurring chos (Trouessart) 1899 in more than one order. Group VI. Specific to the Charadriiformes, Procellari idae this group is the largest in total number of mite Macronectes giganteus species. Extensive collecting from all charad­ Oceanodroma castro Brephosce/es /anceo/atus, bangsi new species riiform families has produced Brephosceles Pterodroma /eucoptera Brephosceles disjunctus, species from only Charadriidae and Haematopo­ hypo/euca new species didae. Additional differentiation in this group Pterodroma leucoptera can be observed within the two avian families. masafuerae Males of new species from Haematopodidae Pterodroma cookii defilippiana are unique in that the pregenital apodeme is Pterodroma externa free whereas males specific to the family Cha­ cervicalis radriidae have the pregenital apodeme fused. Puffinus pacificus Brephosce/es parvatus, Females from both avian families are not dis­ chlorohynchus new species tinctive. For further discussion see the taxonomy Puffinus gravis Puffinus puffinus section, p. 99. Puffinus tenuirostris Host specificity at the species level is prev­ Puffinus puffinus Brephosceles bilobatus, alent throughout group VI. Known associations new species occur among the following pairs of hosts and Group IV. The members of group IV form a parasites. morphologically unique group based on closed Charadriiformes or nearly closed coxal fields Ill and IV and the Haematopodidae absence of extralobar lamellae. Host parasite relationships are limited to the Anseriformes and Haematopus ostral- Brephosce/es haematopi, gus longirostris new species all known species are host specific at the spe­ Haematopus palliatus Brephosceles turgidus, cies level. The mites with their respective hosts palliatus new species are listed as follows: Charadriidae Anseriformes Belonopterus chi/en- Brephosce/es belonopteri, Anatidae sis cayennensis new species Anas platyrhynchos Brephosce/es anatina Belonopterus chi/en- Brephosceles chilensis, Dubinin, 1951 sis chilensis new species Cygnus bewickii Brephosceles discidicus, Belonopterus chi/en- Brephosceles chi/ensis, new species sis lampronatus new species A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 99

Charadrius bicinctus Brephosce/es constrictus, Dubinin (1951) essentially redefined the genus new species Brephosceles. Although his generic diagnosis Charadrius col/aris Brephosceles collaracus, was general, his concept of the genus was cor­ new species Charadrius mongolus Brephosce/es inornatus, rect; four named and three new species were new species included and keys for the identification of males Charadrius tricol/aris Brephosceles histionotus and females were provided. Subsequently, nu­ bifrontatus Gaud and Mouchet, 1957 merous related genera have been erected from Charadrius tricollaris Brephosce/es histionotus new and described species of Brephosceles forbesi Gaud and Mouchet, 1957 Charadrius tricollaris Brephosce/es histionotus (s.1.) and Alloptes (s.1.). tricollaris Gaud and Mouchet, 1957 In 1968, Peterson and Atyeo further restricted Charadrius wilsonia Brephosce/es virginiensis, the definition of the genus Brephosce/es on the new species basis of the relative development, positions, and Hoplopterus duvan- Brephosceles hoplopteri, absence of specific setae, the structures of the celii (=H. ventralis) new species Lobivanellus indicus Brephosceles lobivanelli, male termini and genital regions, and the condi­ atronuchalis new species tions of epimerites I. Only fourteen species were Rhinoptilus africanus Brephosce/es furcifer retained in the genus Brephosce/es {s.s.); these africanus (=Cur- (Trouessart) 1885 species were listed with their pertinent synono­ sorius bicinctus) mies. The remaining species are host specific at the Deposition of Type Material genus level. In retrospect, definitive statements about In the descriptive sections, the major reposi­ host-parasite relationships cannot be made with tories for the primary and secondary types are certainty, particularly with respect to monoxenic denoted by the following abbreviations: associations. However, in the genus Brephosce­ AMNH: American Museum of Natural History, les natural groups are recognized in association New York, New York 10024. with specific avian orders. Within these groups, host specificity varies but in general one host­ BM: Bernice P. Bishop Museum, Honolulu, one parasite is the most prevalent. Hawaii 96819. BMNH: British Museum (Natural History), Cromwell Road, London S.W. 7, Eng­ TAXONOMY land. Historical Account Gaud: Dr. J. Gaud, Direction de la Santa, 3 In 1934, Hull erected the genus Brephosceles rue de Fougeres, Rennes, Ille et Vilaine, for two species of feather mites characterized France. primarily by males having well-developed termi­ LAS: Leningrad Academy of Science, Lenin­ nal lobes bearing lamellae on their free margins. grad, Russia. Prior to this description, Trouessart {1885, 1886) had placed all species with the above charac­ NU: Division of Entomology, University of ters in one of five subgenera of Proctophy/lodes Nebraska State Museum, Lincoln, Ne­ (s.l.). The subgenera included A/loptes, Troues­ braska 68503. sartia (=Pteroco/us), Proctophy/lodes, Ptero­ RNH: Rikjmuseum van Naturlikje Historie, dectes, and Pterophagus. Diagnostic characters Leiden, Netherlands. were given for each species group. Alloptes and SAIMR: South African Institute for Medical Re­ Pterocolus were further subdivided into four search, Hospital Street, Post Office Box subgroups based on the conformation of the 1038, Johannesburg, Republic of South male hysterosomal terminus and the length of Africa. the fourth pair of legs, and in the female, the presence or absence of lobes and/or gladiform SEA: Stazione Entomologica Agraria, via Ro­ appendages. mana 15-17, Florence, Italy. 100 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

SIU: Department of Zoology, Southern Illi­ Distance between external scapular setae. nois University, Carbondale, Illinois Measured center to center. 62901. Seta/ types. Setiform: long and hairlike; lance­ olate: dagger or spear-shaped, tapering to a TC: The Trouessart Collection, c/o Dr. Max point; spiculiform: slender and needle-like. Vachon, 61 rue de Bufton, 75-Paris, Length, hysterosomal shield. Measured along France. the midline from the anterior margin to setae h. UGA: Department of Entomology, University Width, hysterosomal shield. The widest por­ of Georgia, Athens, Georgia 30601. tion, usually at the level of setae /i. Length, hysterosomal lobes. Measured from USNM: United States National Museum, Wash­ the anterior margin of the hysterosomal cleft to ington, D. C. 20560. setae /5. ZSBS: Zoologische Sammlung des Bayer­ Margin of postlamel/ae. Entire: smooth, not ischen Staates, Menzingstrasse 67, incised and without projections; dentate: having Munchen 19, Germany. many small indentations. Coxa/ fields. Closed: completely surrounded ZSZM: Zoologisches Staatsinstitute und Zoolo­ by epimerites; open: epimerites end freely. gisches Museum, von-Melle-Park 10, Within each species group of Brephosce/es Hamburg 13, Germany. the females are remarkably uniform in their characteristics. The formal description of the Characters and Descriptive Methods female has been shortened; duplication of infor­ General morphological characters useful in mation is thereby avoided. species differentiation have been discussed in the morphology section. Many of the character­ Female (Figs. 2 and 4) istics utilized in the formal descriptions are Length. Distance between the apices of the subject to individual preference. The characters pedipalps and setae d5• and descriptive methods parallel the sequence Primary spermatheca/ duct. Expanded prox­ found in the formal descriptions. imally: an enlarged, sclerotized portion of the Phase-contrast microscopes were used in duct near the spermathecal head, ending ab­ this study. Measurements (in microns} were ruptly. Tapering distally: enlarged, not heavily made with an ocular micrometer and drawings sclerotized and tapering gradually to the were made with a Wild microscope equipped supranal concavity. with a drawing attachment. Measurement scales included with illustrations are marked as WM, Family PROCTOPHYLLODIDAE GO, HS, or SP, indicating they are used for the Megnin and Trouessart whole mount, genital organ, hysterosomal shield or spermatheca respectively. Genus BREPHOSCELES Hull Ptero/ichus (in part), Megnin and Trouessart, Male (Figs. 1 and 3) 1884, J. Microgr., 8(7): 380-381. Length, excluding postlamel/ae. Distance be­ Proctophyllodes (Pterocolus} (in part}, Troues­ tween apex of pedipalp and bases of terminal sart, 1885, Bull. Soc. Etud. Sci. Angers, 14: 72, setae /5• 73, 74. Width. The widest part of the idiosoma, usu­ Proctophyllodes (Al/optes} (in part}, Trouessart, ally at level of setae sh. 1886, Bull. Soc. Etud. Sci. Angers, 16: 145-146. Length, propodosomal shield. Anterior margin Alloptes (Al/optes} (in part}, Trouessart, 1899, to the greatest length of posterior margin, in­ Bull. Soc. Etud. Sci. Angers, 28: 58-59. cluding mesal projections. Brephosceles Hull, 1934, Trans. North. Nat. Width, proposomal shield. Distance across the Union, 1: 205. widest part of shield, excluding the projections Brephosceles, Dubinin, 1949, Akad. Nauk. around setae se. S.S.S.R., Zool. Inst., Parasitol. Sborn., 11: 221. A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 101

Brephosceles, Dubinin, 1951, Akad. Nauk. fused anteriorly; pregenital apodeme S.S.S.R., Zool. Inst., Parasitol. Sborn, 13: 220- of female free (Procellariiformes) ...... 227...... group Ill, p. 124 Brephosceles, Dubinin, 1952, Akad. Nauk. 3. Male with coxal field IV closed; pregen- S.S.S.R., Zool. Inst., Trudy, 12: 262-264. ital apodeme free...... 4 Brephosce/es, Gaud, 1959, Bull. Soc. Sci. Nat. Male with coxal field IV open; pregenital Phys. Maroc, 37(2): 114-115. apodeme fused ...... 6 Brephosceles, Gaud and Mouchet, 1957, Ann. 4. Male with genital organ positioned at Parasitol. Hum. Comp., 32(5-6): 239-241. level of trochanter IV. Setae c2 at same Brephosceles, Gaud and Till, 1961, Publ. So. Afr. level or slightly posterior to setae c3• Inst. Med. Res., 11 (L): 239-241. From Anseriformes ...... group IV, p. 133 Brephosceles, Peterson and Atyeo, 1968, Bull. Male with genital organ anterior to level Univ. Nebraska St. Mus., 8(4): 217-236. of trochanter IV. Setae C2 anterior to Mites ectoparasitic on Gaviidae (Gaviiformes), setae Ca ...... 5 Diomedidae, Procellariidae, Hydrobatidae (Pro­ 5. Female with hysterosomal terminus dis­ cellariiformes), Fregatidae (Pelecaniformes), tinctly bilobate. From Charadriiformes Anhimidae, Anatidae (Anseriformes), Gruidae and Gruiformes ...... group V, p. 140 (Gruiformes), Haematopodidae, Charadriidae, Female with hysterosomal terminus en­ Burhinidae, Glareolidae, Laridae (Charadrii­ tire. From Gaviiformes (Brephosc/es formes). Both sexes with epimerites I Y-shaped, forficiger) ...... group I, p. 101 femora and genua of all legs partially fused, 6. Male with pregenital apodeme fused an­ setae vi present, setae ve present or absent. teriorly; coxal field IV open (Charadrii- Males with legs IV equal to or larger than legs formes) ...... group VI, p. 144 Ill, with terminus bilobate and bearing large Male with pregenital apodeme free; marginal lamellae and 1 pair of long terminal coxal field IV open (Charadriiformes) setae, coxal fields Ill-IV open or closed, genital ...... group VI, p. 144 organ minute with simple reflexion. Female similar to A//optes species, with terminal cleft, Brephosce/es forficiger with pregenital apodeme independent or con­ (Megnin and Trouessart) nected to epimerites IV to form a Moresque Ptero/ichus forficiger Megnin and Trouessart, arch as in Pterodectes. 1884, J. Microgr., 8(7): 380-381, Fig. 48(a,b). Type species: Pterolichus forficiger Megnin Type host: Gavia immer (=Colymbus g/acialis) and Trouessart, 1884 (by original designation). (Gaviidae). Pteroco/us forficiger, Berlese, 1898, A. M. S., Key to the species groups fasc. 88, no. 5. 1. Genital arch of male not fused with inner Pterolichus (Eupterolichus) forficiger, Canes­ margin of pregenital apodeme...... 2 trini and Kramer, 1899, Tierreich, 7: 50. Genital arch of male fused with inner Brephosce/es forficiger, Dubinin, 1951, Akad. margin of pregenital apodeme...... 3 Nauk. S.S.S.R., Zool. Inst., Parazitol. Sborn., 2. Both sexes with setae '1 short, bluntly 13: 222. rounded; surface fields well devel­ Brephosceles forficiger, Radford, 1953, Para­ oped. Pregenital apodeme of male in sitol., 43(3,4): 202. shape of inverted V; pregenital apo­ Brephosce/es forficiger, Radford, 1958, Rev. deme of female fused to epimerites Brasil. Entomol., 8: 109. lVa forming a Moresque arch. From Brephosceles forficiger, Vassilev, 1960, Bulgar­ Procellariiformes ...... group II, p. 103 ian Acad. Sci., Proc. Zool. Inst., 9: 436. Both sexes with setae '1 setiform; sur­ Brephosceles forficiger, Peterson and Atyeo, face fields moderate to weakly devel­ 1968, Bull. Univ. Nebraska St. Mus., 8(4): oped. Pregenital apodeme of male not 219-220. 102 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

Figs. 9-12. Brephosce/es torficiger (Megnin and Trouessart), male (lectotype): dorsal aspect (9); ventral aspect (10); female (syntype): dorsal aspect (11); ventral aspect (12). A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 103

The males of Brephosceles forficiger are dis­ indicates that Brephosceles forficiger probably tinguished from related species by the free represents a small unique population in North­ condition of the pregenital apodeme, the fusion ern Europe. Vassilev (1960) reported only one of the genital arch with the pregenital apodeme male and one female from Gavia arctica and and the closed coxal fields Ill and IV. Females Dubin in (1951) reports this species from a loon. are characterized by the hysterosomal terminus The author was unable to recollect this species being entire and the free condition of the pre­ from numerous bird study skins in North Amer­ genital apodeme. ica. The redescription and drawings are from MALE (lectotype). Length, excluding post­ the lectotype male and syntype female. lamellae, 407u; width, 172u. Dorsal idiosoma: Propodosomal shield 112u in length, 112u in HOSTS width; without lacunae; without external vertical Gaviidae setae; distance between external scapular setae, Gavia immer Megnin and 106u. Scapular shields well developed. Setae (Brunnich) 1764 Trouessart, 1884 11 setiform, 10u in length, positioned on well (=Colymbus glacialis) developed humeral shield; setae sh spiculiform Gavia arctica Present study 18u in length, approximate to seta h. Hystero­ (Linne) 1758 Dubinin, 1951 somal shield 246u in length, 88u in width, ante­ Vassilev, 1960 rior margin straight. Hysterosomal lobes 70u in

length; setae d3 positioned at apex of lobe, an­ Group II-the decapus group terior to level of 13 ; setae d4 at terminus of lobe, approximate to pai. lnterlobar and postlobar The grouping of the thirteen species included lamellae well developed. All legs subequal. Ven­ within this complex is based primarily on the tral idiosoma: Epimerites I fused in a Y; surface shape of the pregenital apodeme in both the fields weakly developed. Pregenital apodeme males and the females. In the males this struc­ free, coxal fields Ill and IV closed. Genital organ ture is shaped as an inverted V, with genital 16u in length, not extending beyond setae c2; discs on the anteromedial margin. The female subgenital shield present. Adanal shields not pregenital apodeme is horse-shoe shaped and fused medially; not joined to pregenital apo­ the distal ends are fused with epimerites IV deme. forming a Moresque arch (except Brephosceles FEMALE (syntype). Length, 438u; width, 156u. prolatus which lacks this connection). A number Dorsal idiosoma: Propodosomal shield 104u in of additional characters pertinent to this species length, 93u in width; without lacunae; without group are listed below. external vertical setae; distance between scap­ Pertinent characters for group differentiation, ular setae, 96u. Setae 11 setiform, 8u in length, males: positioned on well developed humeral shield; 1. Surface fields of epimerites I-IV well devel­ setae sh spiculiform 13u in length, slightly pos­ oped. terior to seta h. Hysterosomal shield 264u in 2. Genital apparatus free (i.e., not fused with length, 83u in width; anterior margin straight; inner margin of the pregenital apodeme.) setae d 3 and d 4 in rectangular arrangement, 3. External vertical setae (ve) present. separated by 52u. Hysterosomal terminus entire, 4. lnterlobar, postlobar, and extralobar lam­ without terminal lamellae. Ventral idiosoma: ellae present. Surface fields weakly developed. Preanal apo­ 5. Setae h and sh at approximately the same deme absent. level. Type material. From Gavia immer (Gaviidae): 6. Setae 11 bluntly rounded, positioned on an­ lectotype male (TC), 4 female syntypes (TC), teromedial margin of humeral shield. Northern Europe. The date and collector are 7. Setae d4 at midlength of hysterosomal unknown. lobes, not approximate to setae pae.

Remarks. The rarity of recollected material 8. Setae d 5, 14 and pae long and setiform. 104 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

Pertinent characters for group differentiation, 8. Genital organ extending to or slightly females: beyond the origin of setae c2 •••••••••••••• 9 1. Setae C2 anterior to genital discs. Genital organ never extending to or be- 2. Ventral margins of hysterosoma heavily yond the level of setae C2 ...... 10 sclerotized. 9. Genital organ extremely broad at length, 3. Hysterosomal terminus bilobate with well­ terminating abruptly to a fine point developed terminal lamellae. apically; position of genital apparatus Key to the species of group II subject to mounting procedure ...... (decapus group) ...... pachyptilae, n. sp., p. 110 1. Male with surface fields of epimerites I Genital organ not broad at length, ex- and II fused to form a ventral propo- tending to level of setae C2 ...... dosomal shield ...... 2 ...... puffini, n. sp., p. 118 Male with surface fields of epimerites I 10. Posteriomedial margins of genital ap­ and II separate...... 4 paratus rounded; genital organ not 2. Genital organ extending beyond level of extending beyond these margins; fe­ setae C2••······ ... .longipenis, n. sp., p. 108 male with crescent-shaped preanal Genital organ not extending beyond apodeme .. .. se/enopeltatus, n. sp., p. 123 level of setae C2...... 3 Posteriomedial margins of genital ap­ 3. Genital apparatus triangular in shape paratus not fused; female with preanal with two small pores at apices; female apodeme absent....mar/ae, n. sp., p. 120 with setae /3 slightly posterior to setae d4 ...... decapus (Gaud), p. 104 Brephosce/es decapus (Gaud) Genital apparatus not triangular in Al/optes decapus (Gaud), 1953, Ann. Parasitol. shape, without pores; female with Hum. Comp., 28(3): 196-197, Fig. 2. Type host: setae /3 anterior to setae d4 ...... Oceanodroma castro (Hydrobatidae) ...... superbus Dubinin, p. 106 Alloptes decapus, Radford, 1958, Rev. Brasil. 4. Hysterosomal lobes of male elongate, at Entomol., 8: 111. least one-half the length of the idio­ Brephosceles decapus, Gaud and Till, 1961, soma; setae d4 posterior to the lateral Publ. So. Afr. Inst. Med. Res., 1HL): 241. setae 13 ...... prolatus, n. sp., p. ii Hysterosomal lobes less than one-half Brephosce/es decapus, Peterson and Atyeo, the length of the idiosoma; setae d4 1968, Bull. Univ. Nebraska St. Mus., 8(4): 218-219. anterior to setae /3•••••••••••••••••••••••••••••••••• 5 5. Male with setae C2 positioned on well- Brephosce/es decapus, B. superbus and B. developed subgenital shield...... 6 longipenis are unique to group I. The surface Male with setae C2 not positioned on fields associated with epimerites I and II of each subgenital shield ...... 8 species are fused to form a distinct ventral 6. Subgenital shield crescent shaped; a shield. Males of B. decapus may be separated preanal apodeme may or may not be from the related B. prolatus by the length of the fused with the subgenital shield ...... hysterosomal lobes (228u), from B. superbus by ...... lunatus, n. sp., p. 113 the irregular shape of the ventral shield . Subgenital shield not crescent shaped.... 7 MALE (paratype). Length, excluding postlam­ 7. Male with subgenital shield triangular, ellae, 466u; width, 176u. Dorsal idiosoma: Pro­ setae C2 positioned at apices; genital podosomal shield 117u in length, 108u in width; organ indistinct triangularis, n. sp., p. 115 setae vi, ve present, scapular setae (si, se) in­ Male with subgenital shield variously cluded within shield. Scapular shields well de­ shaped; genital organ extending to veloped. Setae /1 short, spiculiform, Bu in length, level of setae C2 ...... positioned on well developed humeral shield; ...... pterodromae, n. sp., p. 118 setae sh spiculiform, 18u in length, approximate A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 105

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Figs. 13-17. Brephosceles decapus (Gaud), male (paratype): dorsal aspect (13); genital organ (14); ventral aspect (15); female (paratype): dorsal aspect (16); ventral aspect (17). 106 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM to seta h. Hysterosomal shield 284u in length, HOSTS 91 u in width; anterior margin shallowly concave. Hydrobatidae Mesal margin of interlobar lamellae straight; Oceanodroma castro Gaud, 1953 postlobar and extralobar lamellae well devel­ (Harcourt) 1851 Present study oped. Hysterosomal lobes 168u in length; setae Oceanodroma castro /eucor­ Present study d3 positioned at extreme anterior margin of hoa (Vieillot) 1817 complete supranal cleft; setae d 4 at posterior Brephosceles superbus Dubinin one-third of lobe, separated from setae d3 by Brephosce/es superbus Dubinin, 1949, Akad. 91 u, from pai by 48u. Setae d5, pae and '4 long, setiform. Ventral idiosoma: Epimerites I fused Nauk S.S.S.R., Zool. Inst., Trudy, 11: 221. in a Y; surface fields of epimerites I and II fused (Nomen nudum). to form a ventral shield. Pregenital apodeme Brephosceles superbus Dubinin, 1952, Akad. fused to epimerites IVa in shape of inverted V. Nauk S.S.S.R., Zool. Inst., Trudy, 12: 262-264, Genital organ 13u in length, not extending to Fig. 7. Type host: Puffinus tenuirostris (Pro­ cellariidae). level of setae c2• Setae C2 not associated with subgenital shield. Adanal shield well developed Brephosceles superbus, Peterson and Atyeo, with setae a. Preanal apodeme present but 1968, Bull. Univ. Nebraska St. Mus., 8(4): 220. reduced. Brephosceles superbus is distinguished from related species by the setiform condition of FEMALE (paratype). Length, excluding termi­ setae sh and the irregular shape of the ventral nal lamellae, 53u; width, 179u. Dorsal idiosoma: shield (surface fields of epimerites I and II). Propodosomal shield 130u in length, 114u in Additionally, the small dimensions of length and width; setae vi, ve present, scapular setae (si, width of the male are unique to the group. se) included on shield. Humeral shield well de­ MALE. Length, excluding postlamellae, 350u; veloped bearing setae 11 at extreme anterior width, 119u. Dorsal idiosoma: Propodosomal margin; subhumeral setae spiculiform, 16u in shield 91 u in length, 88u in width; setae vi, ve length. Hysterosomal shield 329u in length, 80u present, scapular setae (si, se) included on in width; setae d 3 , d 4 in rectangular arrange­ shield. Scapular shields well developed. Setae ment, separated by 47u. Hysterosomal lobes /1 bluntly rounded, 5u in length, positioned on with well developed, pointed terminal lamellae. well developed humeral shield; setae sh seti­ Ventral idiosoma: Surfase fields of epimerites I form, 10u in length, approximate to seta h. Hys­ and II fused to form ventral shield. Narrow con­ terosomal shield 241u in length, 88u in width; nection between surface fields associated with anterior margin straight. Mesal margin of inter­ epimerites Ill and IV absent. Preanal apodeme lobar lamellae slightly concave; postlobar and small, rounded. extralobar lamellae well developed. Hystero­ Type Material. From Oceanodroma castro somal lobes 137u in length; setae d 3 positioned (Hydrobatidae); holotype male (Gaud}, allotype on anterior one-third of complete supranal cleft; female (Gaud), 3 male, 2 female paratypes, setae d4 at midlength of lobe, seperated from Dakar, on the coast of Senegal, date and col­ setae d 3 by 36u, from pai by 44u, setae d 5, pae lector unknown. Paratypes deposited: (Gaud) and l4 long and setiform. Ventral idiosoma: (French Institute of North Africa). Epimerites I fused in a Y; surface fields fused Material examined. Hydrobatidae: 1 male, 2 to form a ventral shield. Pregenital apodeme females (paratypes) from Oceanodroma castro, fused to epimerites IVa in shape of an inverted Senegal; 1O males, 9 females from Oceano­ V. Genital organ 10u in length, not extending droma castro leucorhoa, Newfoundland. beyond setae C2. Setae C2 not positioned on sub­ Remarks. The drawings and redescriptions of genital shield. Adanal shields well developed the male and female are based on the para­ bearing setae a. Preanal apodeme present. types. The illustration of the genital organ is FEMALE. Length, excluding terminal lamellae, based on a specimen from Oceanodroma castro 428u; width, 119u. Dorsal idiosoma: Propodo­ leucorhoa. somal shield 91u in length, 98u in width; setae A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 107

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Figs. 18-22. Brephosceles superbus Dubinin, male: dorsal aspect (18); genital organ (19); ventral aspect (20); female: dorsal aspect (21); ventral aspect (22). 108 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM vi, ve present, scapular setae (si, se) included on setae d4 at posterior one-third of lobe, sepa­ shield. Humeral shield well developed bearing rated from setae d3 by 65u, from pai by 44u. setae /1 at extreme anteromedial angle; sub­ Setae d 5 and pae relatively short, setiform. Ven­ humeral setae spiculiform, 10u in length. Hyster­ tral idiosoma: Epimerites I fused in a Y; surface osomal shield 334u in length, 88u in width; setae fields well developed. Pregenital apodeme fused d3, d4 in trapezoidal arrangement. Hysterosomal to epimerites IVa in an inverted V. Genital organ lobes with pointed lamellae. Ventral idiosoma: 13u in length not extending beyond insertions Surface fields well developed. Narrow connec­ of setae C2. Subgenital shield absent. Adanal tion between surface fields associated with shields poorly developed not bearing setae a. epimerites Ill and IV absent. Preanal apodeme Preanal apodeme present. moderately developed. FEMALE (allotype). Length, excluding termi­ Type material. From Puffinus tenuirostris (Pro­ nal lamellae, 466u; width, 93u. Dorsal idiosoma: cellariidae): holotype male (LAS) allotype female Propodosomal shield 109u in length, 83u in (LAS), 3 male, 12 female paratypes, July 10, width; setae vi, ve present, scapular setae si 1939, Rogers Bay. Collector unknown. Paratypes included on shield, se on lateral margin. Hu­ are deposited at the Leningrad Academy of meral shield well developed bearing setae /1 at Science. extreme anteriomedial margin; subhumeral Material examined. Procellariidae: 1 male, 1 setae spiculiform, 21 u in length. Hysterosomal female (paratypes) from Puffinus tenuirostris. shield 306u in length, 96u in width; setae d3 , d4 in trapezoidal arrangement, separated by 57u. HOSTS Hysterosomal lobes without pointed terminal Procellariidae lamellae. Ventral idiosoma: Surface fields mod­ Puffinus tenuirostris Dubinin, 1952 erately developed. Narrow connection between (Temminck) 1835 Present study surface fields associated with epimerites Ill and Brephosceles prolatus, new species IV absent. Preanal apodeme present. Type material. From Pachyptila desolata (Pro­ The length of the hysterosomal lobes, the cellariidae): holotype male (BMNH), allotype position of setae d3 and the condition of setae female (BMNH), December 31, 1957, Falkland sh distinguish this species from the related Islands (Dependencies Survey); 1 male, 2 fe­ Brephosceles pachyptilae. In B. prolatus the male paratypes, December 18, 1956, 3 female hysterosomal lobes are 228u in length, setae d3 paratypes, December 20, 1956, 2 male, 1 female is at midlength of the lobes and setae sh is paratypes, November 29, 1956, South Orkney Is­ setiform. Comparatively, in B. pachyptilae the lands, W. L. N. Tickell; 1 male, 1 female para­ lobes do not exceed 135u in length, setae d3 is types, February 1, 1965, Heard Island, P. Temple. positioned on the anterior one-third of lobe and Paratypes deposited: Gaud, UGA, USNM. setae sh is spiculiform. Remarks. The species is named for the length­ MALE (holotype). Length, excluding postlam­ ened hysterosomal lobes. The drawings of the ellae, 513u; width, 179u. Dorsal idiosoma: Pro­ male and female are from the holotype and allo­ podosomal shield 104u in length, 80u in width; type respectively. Illustrations of the genital setae vi, ve present, scapular setae (si, se) in­ organ are from the paratypes. cluded on shield. Scapular shields well devel­ oped. Setae /1 setiform, 10u in length, positioned HOSTS on well developed humeral shield; setae sh Procellariidae spiculiform, 18u in length, approximate to setae Pachyptila desolata (Gmelin) Present study h. Hysterosomal shield 352u in length, 104u in 1789 width; anterior margin straight. Mesa! margin of interlobar lamellae undulate; postlobar and Brephosceles /ongipenis, new species extralobar lamellae well developed. Hystero­ This new species, although related to Breph­ somal lobes 228u in length; setae d 3 positioned osce/es decapus and B. superbus, can easily be at midlength of lobe; supranal cleft incomplete; distinguished by the length of the genital organ. A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 109

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Figs. 23-27. Brephosceles prolatus, new species, male (holotype): dorsal aspect (23); genital organ (24); ventral aspect (25); female (allotype): dorsal aspect (26); ventral aspect (27). 110 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

In B. longipenis the genital organ greatly sur­ one male and one female. Additional collecting passes the origin of setae C2 whereas in the from phylogentically related avian species will related species the genital organ is small and undoubtedly extend the range of this species. inconspicuous. The species is named for the length of the MALE (holotype). Length, excluding postlam­ genital organ. ellae, 461 u; width, 161 u. Dorsal idiosoma: Pro­ HOSTS podosomal shield 135u in length, 109u in width; Hydrobatidae setae vi, ve present, scapular setae (si, se) in­ Oceanodroma markhami Present study cluded on shield. Scapular shields well devel­ tristrami Salvi n, 1896 oped. Setae 11 bluntly rounded, 8u in length, positioned on well developed humeral shield; Brephosceles pachyptilae, new species setae sh spiculiform, 21 u in length, approximate This new species, although closely related to to setae h. Hysterosomal shield 272u in length, Brephosceles prolatus, is unique in respect to 93u in width; anterior margin shallowly concave the genital organ. In B. pachyptilae this struc­ with heavy sclerite. Mesal margin of interlobar ture is extremely broad basally and tapers to a lamellae straight, postlobar and extralobar lam­ fine point distally. The species is additionally ellae well developed. Hysterosomal lobes 166u characterized by the length of the hysterosomal in length; setae d3 at anterior one-third of com­ lobes (135u), setae d3 on anterior one-third of plete supranal cleft; setae d4 at midlength of lobe and the spiculiform setae sh. lobe, separated from setae d3 by 67u, from pai MALE (holotype). Length, excluding lamellae, by 21 u. Setae d 5 , pae and l4 long and setiform. 420u; width, 171 u. Dorsal idiosoma: Propodoso­ Ventral idiosoma: Epimerites I fused in a Y; mal shield 119u in length, 108u in width; setae surface fields of epimerites I and II fused to vi, ve present, scapular setae (si, se) included form a ventral shield. Pregenital apodeme fused on shield. Scapular shields well developed. to epimerites IVa in shape of inverted V. Genital Setae l1 bluntly rounded, Bu in length, positioned organ 57u in length, extending beyond setae C2. on well developed humeral shield; setae sh Setae c2 not associated with subgenital shield. spiculiform, 18u in length, approximate to setae Adanal shields well developed, bearing seta a. h. Hysterosomal shield 259u in length, 93u in Preanal apodeme present. width; anterior margin straight. Mesal margin of FEMALE (allotype). Length, excluding terminal interlobar lamellae straight; postlobar and extra­ lamellae, 482u; width, 169u. Dorsal idiosoma: lobar lamellae well developed. Hysterosomal

Propodosomal shield 11 ?u in length, 106u in lobes 135u in length; setae d3 positioned on width; setae vi, ve present, scapular setae (si, anterior one-third of complete supranal cleft; se) included in shield. Humeral shield well de­ setae d4 at midlength of lobe, separated from veloped, bearing setae Ii at extreme anterome­ setae d3 by 26u, from pai by 47u. Setae d5, pae dial angle; subhumeral setae spiculiform, 16u in and 14 long and setiform. Ventral idiosoma: length. Hysterosomal shield 308u in length, 101 u Epimerites I fused in a Y; surface fields well in width; setae d3 , d 4 in trapezoidal arrangement, developed. Pregenital apodeme fused to epimer­ separated by 52u. Hysterosomal lobes with long ite IVa in an inverted V. Genital organ 31 u in tapering terminal lamellae. Ventral idiosoma: length, extremely thick at base and tapers to a Epimerites I and II with surface fields well devel­ fine point, extending beyond insertion of setae oped. Narrow connection between surface fields c2 • Subgenital shield absent. Adanal shields well associated with epimerites Ill and IV absent. developed bearing setae a. Preanal apodeme Preanal apodeme small, rounded. present. Type material. From Oceanodroma markhami FEMALE (allotype). Length, excluding terminal tristrami (Hydrobatidae): holotype male (USNM), lamellae, 472u; width, 168u. Dorsal idiosoma: allotype female (USNM), January 6, 1913, Laysan Propodosomal shield 122u in length, 111 u in Island, G. Willett. width; setae vi, ve present, scapular setae (si, Remarks. This species is known from only se) included on shield. Humeral shield well de- A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 111

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Figs. 28-32. Brephosce/es /ongipenis, new species, male (holotype): dorsal aspect (28); genital organ (29); ventral aspect (30); female (allotype): dorsal aspect (31); ventral aspect (32). 112 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

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Figs. 33-38. Brephosceles pachptilae, new species, male {holotype): dorsal aspect {33); illustrations of genital organ {34, 35); ventral aspect {36); female {allotype): dorsal aspect {37); ventral aspect {38). A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 113 veloped bearing setae 11 at extreme anterome­ effectively separate this female from other spe­ dial angle; sub humeral setae sh spiculiform, 21 u cies included in group I. Therefore, until the in length. Hysterosomal shield 290u in length, acquisition of a male specimen, the species is

93u in width; setae d3 , d4 in trapezoidal arrange­ retained without further differentiation. ment separated by 42u. Hysterosomal lobes with MALE. Unknown. pointed terminal lamellae. Ventral idiosoma: FEMALE. Length, excluding terminal lamellae, Surface fields separate, well developed. Narrow 420u; width, 132u. Dorsal idiosoma: Propodoso­ connection between surface fields associated mal shield well developed; setae vi, ve present, with epimerites Ill and IV absent. Preanal apo­ scapular setae (se) as long as one-half the body deme reduced. width. Humeral shield well developed, bearing Type material. From Pachypti/a desolata (Pro­ setae '1 at extreme anteromedial angle; sub­ cellariidae): holotype male (BMNH), allotype humeral setae spiculiform, approximately two­ female (BMNH), 7 male, 7 female paratypes, thirds the body width. Hysterosomal shield January 7, 1958 and 1 male, 2 female paratypes, heavily sclerotized; with supranal concavity. December 31, 1957, Falkland Islands (Depend­ Hysterosomal lobes with terminal lamellae. Ven­ encies Survey); 3 male, 8 female paratypes, tral idiosoma: Surface fields well developed; October 13, 1955, 1 male paratype, November 9, without narrow connection between surface 1955, 2 male, 4 female paratypes, December 18, fields associated with epimerites Ill and IV. Pre­ 1956, 1 male, 3 female paratypes, December 18, anal apodeme apparently absent. 1956, 3 male paratypes, December 26, 1956, 1 Type material. From Hydrobates pelagicus male, 2 female paratypes, December 20, 1956, (Hydrobatidae): November 16, 1920, North Sea, South Orkney Islands, W. L. N. Tickell; 1 male, collector unknown. Primary types are deposited 1 female paratypes, February 1, 1965, Heard in the personal collection of Dr. H. G. Vitzthum. Island, P. Temple. Paratypes deposited: BMNH, HOSTS Gaud, LAS, NU, RNH,SEA, UGA,ZSBS,ZSZM. Hydrobatidae The formal descriptions are based on the holo­ Hydrobates pelagicus (L.) Vitzthum, 1921 type and allotype. Drawings are from the para­ 1758 types. Brephosceles lunatus, new species Remarks. The species is named for the type host. Whole mount drawings and the illustrations The length of the genital organ and the shape are from the holotype and allotype. of the subgenital shield distinguish this species from the related Brephosceles triangularis. In B. HOSTS lunatus the genital organ is distinct and the sub­ Procellariidae genital shield semicircular in contrast to the Pachypti/a desolata (Gmelin) Present study indistinct genital organ and the triangular sub­ 1789 genital shield in B. triangularis. MALE (holotype). Length, excluding post­ Brephosceles pelagicus (Vitzthum) 1921 lamellae, 472u; width, 192u. Dorsal idiosoma: Pterodectes pelagicus Vitzthum, 1921, Arch. f. Propodosomal shield 140u in length, 124u in Naurgesch., A, 87(4): 66-69, Figs. 48 and 49. width; setae vi, ve present, scapular setae (si, Type host: Hydrobates pelagicus (L.). se) included on shield. Scapular shields well Brephosceles pelagicus, Peterson and Atyeo, developed. Setae '1 bluntly rounded, Bu in 1968, Bull. Univ. Nebraska St. Mus., 8(4): 220. length, positioned on well developed humeral Due to the absence of type material or speci­ shield; setae sh spiculiform, 18u in length, ap­ mens collected from the type host, the following proximate to setae h. Hysterosomal shield 280u description is an interpretation and paraphras­ in length, 101u in width; anterior margin straight. ing of information taken from the original de­ Mesal margin of interlobar lamellae undulate; scription and drawings of the female (Vitzthum, postlobar and extralobar lamellae well devel­ 1921, pages 66-69). It will be noted that none of oped. Hysterosomal lobes 148u in length; setae the characters presented in this description d3 positioned on anterior one-third of complete 114 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

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Figs. 39-44. Brephosceles lunatus, new species, male (holotype): dorsal aspect (39); illustrations of genital organ (40, 41); ventral aspect (42); female (allotype): dorsal aspect (43); ventral aspect (44). A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 115

supranal cleft; setae d4 at midlength of lobe, Brephosceles triangularis, new species separated from setae d 3 by 21 u, from pai by 65u. The triangular shape of the subgenital genital Setae d 5, pae and /4 long and setiform. Ventral shield and the indistinct genital organ form a idiosoma: Epimerites I fused in a Y; surface unique combination of characters. All observed fields separate, well developed. Pregenital apo­ male specimens had setae C2 positioned on the deme fused to epimerites IVa in shape of in­ apex of the subgenital shield. verted V. Genital organ 21 u in length, not MALE (holotype). Length, excluding postlam­ extending beyond setae c2. Setae C2 positioned ellae, 435u; width, 171u. Dorsal idiosoma: Pro­ on crescentric subgenital shield. Adanal shields podosomal shield 127u in length, 106u in width; well developed bearing setae a. Preanal apo­ setae vi, ve present, scapular setae (si, se) deme fused to subgenital shield. included on shield. Scapular shields well devel­ FEMALE (allotype). Length, excluding terminal oped. Setae /1 bluntly rounded, Su in length, lamellae, 513u; width, 171u. Dorsal idiosoma: positioned on well developed humeral shield, Propodosomal shield 137u in length, 117u in seta sh spiculiform, 10u in length, approximate width; setae vi, ve present, scapular setae (si, to setae h. Hysterosomal shield 259u in length, se) included on shield. Humeral shield well de­ 98u in width; anterior margin slightly concave. veloped bearing setae /1 at anteromedial angle; Mesal margin of interlobar lamellae relatively subhumeral setae spiculiform, 23u in length. straight; postlobar and extralobar lamellae well Hysterosomal shield 306u in length, 99u in width; developed. Hysterosomal lobes 137u in length; setae d 3, d4 in rectangular arrangement, sepa­ setae d 3 positioned on anterior one-third of com­ rated by 31 u. Hysterosomal lobes with terminal plete supranal cleft; setae d4 at midlength of lamellae bluntly rounded. Ventral idiosoma: Sur­ lobe, separated from setae d3 by 29u, from pai face fields separate, well developed. Narrow by 49u. Setae d5, pae and /4 long and setiform. connection between surface fields associated Ventral idiosoma: Epimerites I fused in a Y; sur­ with epimerites Ill and IV present. Preanal apo­ face fields well developed. Pregenital apodeme deme well developed crescentric shaped. fused to epimerites IVa in shape of inverted V. Type material. From Pterodroma externa ex­ Genital organ indistinct. Seta C2 positioned on terna (Procellariidae): holotype male (USNM), anterior margin of triangular shaped subgenital allotype female (USNM), 10 male, 13 female shield. Adanal shields well developed bearing paratypes, near Masatierra Island, Chile, Janu­ seta a. Preanal apodeme fused to posterior mar­ ary 21, 1914, and paratypes, December 30, 1913, gin of subgenital shield. C. Beck. Paratypes deposited: AMNH, BMNH, FEMALE (allotype). Length, excluding terminal Gaud, LAS, RNH, SAIMR, USNM, ZSBS. lamellae, 484u; width, 156u. Dorsal idiosoma: Additional material. Procellariidae: 3 males Propodosomal shield 125u in length, 106u in from Pterodroma externa cervica/is, Kermadec width; setae vi, ve present, scapular setae (si, Islands. se) included on shield. Humeral shield well de­ Remarks. The drawings of the male and fe­ veloped bearing setae '1 at anteromedial angle; male are from the holotype and allotype re­ subhumeral setae spiculiform, 18u in length. spectively. Illustrations of genital organ and Hysterosomal shield 298u in length, 88u in width; variations in the shape of the subgenital shield setae d 3, d 4 in rectangular arrangement sepa­ are from the paratypes. The species is named rated by 36u. Hysterosomal lobes with bluntly for the crescentric shape of the subgenital rounded terminal lamellae. Ventral idiosoma: shield. Surface fields separate, well developed. Narrow HOSTS connection between surface fields associated Procellariidae with epimerites Ill and IV absent. Preanal apo­ Pterodroma externa externa Present study deme present. (Salvin) 1876 Type material. From Pterodroma cookii defilip­ Pterodroma externa cervica/is Present study piana (Procellariidae): holotype male (USNM), (Salvin) 1891 allotype female (USNM), 1 male, 1 female para- 116 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

46

250u 50u

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Figs. 45-49. Brephosce/es triangularis, new species, male (holotype): dorsal aspect (45); genital organ (46); ventral aspect (47); female (allotype): dorsal aspect (48); ventral aspect (49). A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 117

. . .

. t . 51

250u 50u

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Figs. 50-54. Brephosceles pterodromae, new species, male (holotype): dorsal aspect (50); genital organ (51); ventral aspect (52); female (allotype): dorsal aspect (53); ventral aspect (54). 118 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM types, 155 miles S.E. Talara, Peru, September 1, se) included in shield. Humeral shield well de­ 1924, C. L. Fagen. Paratypes deposited: UGA. veloped, bearing setae /1 at extreme anterome­ Remarks. The drawings of the male, female dial angle; subhumeral setae spiculiform, 23u in and the illustration of the genital organ are from length. Hysterosomal shield 307u in length, 104u the holotype and allotype. The species is named in width; setae d 3, d4 separated by 13u, in rec­ for the triangular shape of the subgenital shield. tangular arrangement. Hysterosomal lobes with pointed terminal lamellae. Ventral idiosoma: HOSTS Epimerites I and II with surface fields well devel­ Procellari idae oped. Surface fields of epimerites Ill and IV Pterodroma cookii defilippiana Present study joined by narrow connection. Preanal apodeme (Giglioli and Salvadori) 1869 well developed, semicircular. Brephosce/es pterodromae, new species Type material. From Pterodroma /eucoptera hypoleuca (Procellariidae): holotype male (SIU), This new species is more closely related to allotype female (SIU), 4 male, 6 female para­ B. triangularis than to any other species in types, Midway Island, December 7, 1959, J. C. group I. The males are distinguished primarily Downey; Paratypes: 4 males, 7 females, Green by (1) the triangular subgenital shield, (2) the Island, Kure Atoll, Hawaiian Islands, March 7, genital organ extending to level of setae c2. The 1965, 4 females, March 6, 1965, 3 males, 2 fe­ females are characterized by the semicircular males, Lisianski Island, February 14, 1963; 2 preanal apodeme and the short distance be­ males, 2 females, Bonin Island, February, 1910; tween setae ds and d4. 1 male, 4 females, Pacific Ocean, July 9, 1964. MALE {holotype). Length, excluding postlam­ Paratypes deposited: BM, Gaud, LAS, UGA, ellae, 474u; width, 189u. Dorsal idiosoma: Pro­ USNM. podosomal shield 130 in length, 125u in width; Remarks. The drawings of the male and fe­ setae vi, ve present, scapular setae (si, se) in­ male are from the holotype and allotype respec­ cluded in shield. Scapular shield well developed. tively. The illustration of the genital organ is Setae '1 bluntly rounded, 1Ou in length, posi­ from a paratype. The species is named for the tioned on well developed humeral shield; setae type host. sh spiculiform, 16u in length, approximate to HOSTS setae h. Hysterosomal shield 303u in length, Procellariidae 111 u in width; anterior margin straight. Masai Pterodroma leucoptera hypo- Present study margin of interlobar lamellae undulate, post­ Jeuca (Salvin) 1888 lobar, and extralobar lamellae well developed. Hysterosomal lobes 145u in length; setae d3 at Brephosceles puffini, new species apex of complete supranal cleft; setae d4 at mid­ In Brephosceles puffini the genital organ ex­ length of lobe, separated from setae d3 by 39u, tends to the level of setae C2, the subgenital from pai by 57u. Setae d5, pae, and /4 long and shield is absent and in the female the preanal setiform. Ventral idiosoma: Epimerites I fused apodeme is small and circular. This combina­ in a Y; surface fields separate, well developed. tion of characters sufficiently distinguishes this Pregenital apodeme fused to epimerites IVa in new species from the following B. marlae in shape of inverted V. Genital organ 26u in length, which the genital organ extends nearly to the extending to level of setae C2. Setae C2 posi­ origins of setae C2 and the preanal apodeme of tioned on anterior margin of small triangular the female is absent. shield. Adanal shield triangular in shape, bear­ MALE (holotype). Length, excluding postlam­ ing seta a on medial apex. Preanal apodeme ellae, 420u; width, 158u. Dorsal idiosoma: Pro­ present. podosomal shield 119u in length, 93u in width; FEMALE (allotype). Length, excluding terminal setae vi, ve present, scapular setae (si, se) in­ lamellae, 512u; width, 176u. Dorsal idiosoma: cluded on shield. Scapular shields well devel­ Propodosomal shield 143u in length, 127u in oped. Setae /1 bluntly rounded, Su in length, width; setae vi, ve present, scapular setae (si, positioned on well developed humeral shield; A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 119

56

200u 50u

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Figs. 55-59. Brephosce/es puffini, new species, male (holotype): dorsal aspect (55); genital organ (56); ventral aspect (57); female (allotype): dorsal aspect (58); ventral aspect (59). 120 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM setae sh spiculiform, 13u in length, approximate HOSTS to setae h. Hysterosomal shield 246u in length, Procellariidae 80u in width; anterior margin straight. Mesal Puffinus gravis (O'Reilly) Present study margin of interlobar lamellae relatively straight; 1818 postlobar and extralobar lamellae well devel­ Puffinus griseus (Gmelin) Present study oped. Hysterosomal lobes 130u in length; setae 1789 d3 positioned on anterior one-third of complete Brephosceles marlae, new species supranal cleft; setae d4 at midlength of lobe, The males of this new species may be sepa­ separated from setae ds by 31 u, from pai by 47u. rated from the former related two species by Setae d5, pae and l4 long and setiform. Ventral the length of the genital organ and the absence idiosoma: Epimerites I fused in a Y; surface of the subgenital shield. In Brephosceles mar­ fields well developed. Pregenital apodeme fused lae the genital organ extends nearly to the to epimerites IVa in shape of inverted V. Genital level of setae C2 whereas in B. selenopeltatus organ 16u in length, extending to level of setae this structure does not surpass the posterior c2 • Setae c2 not associated with subgenital margin of the well developed subgenital shield. shield. Adanal shields well developed bearing In the females, the preanal apodeme is absent. setae a. Preanal apodeme present. MALE (holotype). Length, excluding postlam­ FEMALE (allotype). Length, excluding termi­ ellae, 415u; width, 171u. Dorsal idiosoma: Pro­ nal lamellae, 456u; width, 148u. Dorsal idiosoma: podosomal shield 117u in length, 99u in width; Propodosomal shield 119u in length, 93u in setae vi, ve present, scapular setae (si, se) in­ width; setae vi, ve present, scapular setae (si, cluded on shield. Scapular shield well devel­ se) included on shield. Humeral shield well de­ oped. Setae 11 bluntly rounded, 8u in length, veloped bearing setae '1 at anteromedial angle; positioned on well developed humeral shield; subhumeral setae spiculiform, 16u in length. setae sh spiculiform, 1Ou in length, approximate Hysterosomal shield 272u in length, 83u in to setae h. Hysterosomal shield 239u in length, width; setae d 3 , d4 in trapezoidal arrangement 91 u in width; anterior margin straight. Mesal separated by 44u. Hysterosomal lobes with margin of interlobar lamellae shallowly concave; pointed terminal lamellae. Ventral idiosoma: postlobar and extralobar lamellae well devel­ Surface fields separate, well developed. Narrow oped. Hysterosomal lobes 104u in length; setae connection between surface fields associated d 3 positioned at anterior one-third of complete with epimerites Ill and IV absent. Preanal apo­ supranal cleft; setae d4 at posterior one-third of deme present, but reduced. lobe, separated from setae d3 by 34u, from pai

Type material. From Puffinus gravis (Procel­ by 41 u. Setae d 5, pae and l4 long and setiform. lari idae): holotype male (NU), allotype female Ventral idiosoma: Epimerites I fused in a Y; sur­ (NU), 6 male, 14 female paratypes, July 27, face fields separate well developed. Pregenital 1962, Kent Island, Brunswick, Canada, K. Hy­ apodeme fused to epimerite IVa in the shape of land, G. West, A. Moorhouse; 1 male, 3 female an inverted V. Genital organ 13u in length, ex­ paratypes, July 8, 1945, Cape Freels, Newfound­ tending to level of setae C2. Setae C2 not asso­ land, H. S. Peters, T. D. Burleigh; 3 male para­ ciated with subgenital shield. Adanal shield types, June 1, 1896, Sable Island, Nova Scotia, variously shaped, bearing seta a. Preanal apo­ J. Dwight, Jr.; 1 male, 1 female paratypes, Au­ deme present, but reduced. gust 30, 1894, Labrador Coast. Paratypes depos­ FEMALE (allotype). Length, excluding terminal ited: AMNH, Gaud, LAS, UGA, USNM. lamellae, 467u; width, 153u. Dorsal idiosoma: Additional material. Procellariidae: 3 males, 9 Propodosomal shield 114u in length, 104u in females from Puffinus griseus, Canada; 3 fe­ width; setae vi, ve present, scapular setae (si, males from P. griseus, Newfoundland. se) included on shield. Humeral shield well de­ Remarks. The drawings of the male and veloped bearing setae '1 at anteromedial angle; female are from the holotype and allotype re­ subhumeral setae spiculiform, 10u in length. spectively. The species is named for the host. Hysterosomal shield 282u in length, 88u in width; A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 121

250u SOu

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Figs. 60-64. Brephosceles marlae, new species, male (holotype): dorsal aspect (60); genital organ (61); ventral aspect (62); female (allotype): dorsal aspect (63); ventral aspect (64). 122 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

~ !, 66

2SOu 50u

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Figs. 65-69. Brephosceles selenopeltatus, new species, male (holotype): dorsal aspect (65); genital organ (66); ventral as­ pect (67); female (allotype): dorsal aspect (68); ventral aspect (69). A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 123

setae d3 , d4 in trapezoidal arrangement, sepa­ and l4 long and setiform. Ventral idiosoma: rated by 39u. Hysterosomal lobes with pointed Epimerites I fused in a Y; surface fields well de­ terminal lamellae. Ventral idiosoma: Surface veloped. Pregenital apodeme fused to epimer­ fields separate, well developed. Narrow connec­ ites IVa in shape of an inverted V. Genital organ tion between surface fields associated with 11 u in length not extending beyond setae c2 • epimerites Ill and IV absent. Preanal apodeme Setae C2 not positioned on divided subgenital absent. shield. Adanal shields well developed bearing Type material. From Puffinus tenuirostris (Pro­ setae a. Preanal apodeme present. cellariidae): holotype male (USNM), allotype FEMALE (allotype). Length, excluding terminal female (USNM), 2 male, 4 female paratypes, lamellae, 466u; width, 53u. Dorsal idiosoma: Little Sitkin Island, Alaska, June 26, 1936, 0. J. Propodosomal shield 119u in length, 99u in Murie; Paratypes: 1 male, 1 female, S. of Unimak width; setae vi, ve present, scapular setae (si, Pass, Aleutian Islands, July 29, 1888; 7 females, se) included on shield. Humeral shield well de­ Cooper Island, Commander Islands, June 18, veloped bearing setae 11 at extreme anterome­ 1912. Paratypes deposited: AMNH, BM, NU, dial angle; subhumeral setae spiculiform, 13u in UGA. length. Hysterosomal shield 280u in length, 86u Remarks. The drawings of the male and fe­ in width; setae d 3, d4 in trapezoidal arrangement, male are from the holotype and allotype respec­ separated by 36u. Hysterosomal lobes with apex tively. The illustration of the genital organ is of terminal lamellae rounded. Ventral idiosoma: from a paratype. Surface fields separate, well developed. Narrow connection between surface fields associated HOSTS with epimerites Ill and IV absent. Preanal apo­ Procellariidae deme well developed, crescentric in shape. Puffinus tenuirostris Present study Type material. From Puffinus pacificus cune­ (Temminck) 1835 atus (Procellariidae): holotype male (USNM), Brephosceles selenopeltatus, new species allotype female (USNM), 2 male, 3 female paratypes, San Benedicto Island, Revillagigedo This new species can be separated from the Islands, Mexico, May 12, 1925. Paratypes: 1 related Brephosceles puffini and B. marlae male, 6 females, Nihoa Island, Hawaiian Islands, by the presence of a well defined subgenital June 2, 1902, W. K. Fisher. shield and the genital organ which does not Additional material. Procellariidae: 2 males, 6 extend beyond the origin of setae C2. The fe­ females from Puffinus pacificus chlorohynchus, males can be distinguished by the crescentric Eastern Island, Midway Atoll, Central Pacific; 1 shape of the preanal apodeme. male, 7 females from Puffinus pacificus, Pacific MALE (holotype). Length, excluding postlam­ Ocean; 2 males, 2 females from Puffinus bulleri, ellae, 407u; width, 158u. Dorsal idiosoma: Pro­ Chile. podosomal shield 112u in length, 93u in width; Remarks. The three recorded host species are setae vi, ve present, scapular setae (si, se) currently assigned to the avian subgenus Thello­ included on shield. Scapular shields well devel­ droma. It is probable that Brephosceles seleno­ oped. Setae '1 bluntly rounded, au in length, peltatus is host specific to this taxa. The draw­ positioned on well developed humeral shield; ings of the male and female are from the halo­ setae sh spiculiform, 10u in length, approximate type and allotype respectively. The illustration to setae h. Hysterosomal shield 239u in length, of the genital organ is from one of the para­ 86u in width; anterior margin straight. Mesal types. The species is named for the crescentric margin of interlobar lamellae straight; postlobar preanal apodeme of the female. and extralobar lamellae well developed. Hystero­

somal lobes 127u in length; setae d 3 positioned HOSTS on anterior one-third of complete supranal cleft; Procellariidae

setae d4 at midlength of lobe, separated from Puffinus pacificus cuneatus Present study setae d 3 by 23u, from pai by 57u. Setae d 5, pae Salvin, 1888 124 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

Puffinus pacificus ch/oro­ Present study Epimerites IVa fused to pregenital apo­ hynchus Lesson, 1831 deme; female with hysterosomal ter­ Puffinus bulleri Salvin Present study minus weakly bilobate or entire...... 3 1888 3. Male with pregenital apodeme diver­ gent; surface fields associated with Group Ill-the marginiventris group epimerites IV and Illa weakly devel­ In their typical form, the major characters oped, not fused with pregenital apo- employed to differentiate group Ill from group deme ...... parvatus, n. sp., p. 133 II are easily observable, that is, the male pre­ Male with pregenital apodeme not diver­ genital apodeme is not fused anteromedially in gent; surface fields well developed.... 4 group Ill versus the fusion of the pregenital 4. Male with surface fields of epimerites apodeme in group II. In both sexes the surface Illa and IV encompassing entire coxal fields are variously conformed; a ventral propo­ field; female with setae d4 setiform ...... dosomal shield may be present or absent de­ ...... bilobatus, n. sp., p. 131 pending upon the species. The females are Male with surface fields of epimerites recognized by the elongate chelicery and the Illa and IV not encompassing entire hysterosomal terminus which is typically entire coxal field; female with setae si and or weakly bilobate. A preanal apodeme may be d4 lanceolate .... /anceo/atus, n. sp., p. 129 present or absent. 5. Male with two marginally serrate postlo­ bar lamellae; female terminus weakly Pertinent characters for group recognition, bilobate, preanal apodeme absent...... males: ...... marginiventris (Trouessart) p. 124 1. Pregenital apodeme fused anteromedially. Male with one postlobar lamellae, with­ 2. Overall dimensions smaller than other out serrations; female terminus entire, groups. preanal apodeme present...... 3. External vertical setae (ve) absent. .... diomedei (Atyeo and Peterson) p. 127 4. Setae '1 and sh setiform. Brephosceles marginiventris (Trouessart) 5. Setae d4 approximate to setae pai. Proctophyl/odes (Alloptes) marginiventris (Trou­ Pertinent characters for group recognition, essart, 1899, Bull. Soc. Etud. Sci. Angers, 28: females: 58-59. Type host: Diomedea chlorohynchos 1. Setae C2 anterior to genital discs. (Diomedeidae). 2. Ventrolateral margins of hysterosoma not Brephosceles marginiventris, Peterson and At­ heavily sclerotized. yeo, 1968, Bull. Univ. Nebraska St. Mus., 8(4): 3. Hysterosomal terminus entire or weakly 220. bilobate. Brephosce/es marginiventris is related to the 4. Chelicery elongate, movable digit often following species B. diomedei. The males are spoon-shaped with marginal serrations. characterized in part by the absence of a ven­ tral shield, surface fields of epimerites Ill well Key to the species of group Ill developed and legs IV which surpass the hys­ (marginiventris group) terosomal terminus by the length of the tarsus. 1. Male with surface fields of epimerites I The females are distinguished by the absence and 11 fused to form a ventral propo- of the preanal apodeme and by the hysteroso­ dosomal shield ...... 2 mal terminus which is weakly bilobate. Male with surface fields of epimerites I MALE (lectotype). Length, excluding post­ and II separate...... 5 lamellae, 415u; width, 169u. Dorsal idiosoma: 2. Epimerites IV of male fused to pregenital Propodosomal shield 109u in length, 83u in apodeme; female with hysterosomal width; without lacunae; without external vertical terminus bilobate; preanal apodeme setae; distance between external scapular setae, crescentric ...... disjunctus, n. sp., p. 129 70u. Scapular shields well developed. Setae '1 A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 125

2SOu

Figs. 70-73. Brephosceles marginiventris (Trouessart), male {lectotype): dorsal aspect (70); ventral aspect (71); female {syntype): dorsal aspect (72); ventral aspect (73). 126 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

200u

74

Figs. 74-77. Brephosceles diomedei, new species, male (holotype): dorsal aspect (74); ventral aspect (75); female (allo­ type): dorsal aspect (76); ventral aspect (77). A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 127 setiform, Su in length, positioned on well devel­ distinguished by legs IV not surpassing the hys­ oped humeral shield; setae sh Su in length, terosomal lobes and by the smoothly rounded approximate to setae h. Hysterosomal shield interlobar lamellae. In contradistinction to B. 244u in length, 93u in width; anterior margin marginiventris the females of B. diomedei pos­ straight. Hysterosomal lobes 104u in length; sess a well developed preanal apodeme. setae d3 positioned at midlength of lobe, the MALE (holotoype). Length, excluding post­ same level as setae 13 ; setae d4 and pai approxi­ lamellae, 363u; width, 116u. Dorsal idiosoma: mate. Postlobar lamellae with margin serrate; Propodosomal shield 88u in length, 62u in width; interlobar and extralobar lamellae well devel­ without lacunae; without external vertical setae; oped. Legs IV hypertrophied, surpassing termi­ distance between external scapular setae, 57u. nus by the length of the tarsi. Ventral idiosoma: Scapular shields well developed. Setae 11 seti­ Epimerites I fused in a Y; surface fields moder­ form, Su in length, positioned on well developed ately developed. Pregenital apodeme free, fused humeral shield; setae sh 10u in length, approxi­ to epimerites IVa. Genital organ Su in length, mate to setae h. Hysterosomal shield 220u in not extending beyond setae c2 ; subgenital shield length, 62u in width; anterior margin straight. present. Adanal shield moderately developed; Hysterosomal lobes 11 Ou in length; setae d3 without setae a. positioned at posterior one-third of lobe, pos­ FEMALE (syntype). Length, 433u; width, 145u. terior to setae 13 ; setae d4 and pai approximate. Dorsal idiosoma: Propodosomal shield 109u in lnterlobar, extralobar and postlobar lamellae length, 88u in width; without lacunae; without well developed. All legs subequal. Ventral idio­ external vertical setae; distance between scapu­ soma: Epimerites I fused in a Y; surface fields of lar setae, 80u. Setae '1 setiform, 10u in length, epimerites I and II fused to form a ventral shield. positioned on well developed humeral shield; Coxal field IV heavily sclerotized. Pregenital setae sh Su in length, approximate to setae h. apodeme free, fused to epimerites IVa. Genital Hysterosomal shield 259u in length, 93u in width; organ Su in length, not extending beyond setae anterior margin straight. Hysterosomal terminus c2 ; subgenital shield apparently absent. Adanal weakly bilobate; without terminal lamellae. Ven­ shield weakly developed; without setae a. tral idiosoma: Surface fields weakly developed. FEMALE (allotype). Length, 384u; width, 119u. Preanal apodeme absent. Dorsal idiosoma: Propodosomal shield 99u in Type material. From Diomedea chlorohynchos length, 67u in width; without lacunae; without (Diomedeidae): lectotype male (TC), syntype fe­ external vertical setae; distance between scap­ male (TC), lie St. Paul. ular setae, 62u. Setae '1 setiform, Su in length, Additional material. Procellariidae: 14 males, positioned on well developed humeral shield; 4 females from Macronectes giganteus, South setae sh 1Ou in length, approximate to setae h. Georgia Island. Hysterosomal shield 220u in length, 67u in Remarks. To date, repeated attempts to rec­ width; anterior margin straight. Hysterosomal ollect this species from Diomedea have proved terminus entire, without terminal lamellae. Ven­ futile. tral idiosoma: Surface fields separate, well de­ HOSTS veloped. Preanal apodeme crescentric. Diomedeidae Type material. From Diomedea chrysostoma Diomedea chlorohynchos Trouessart, 1899 (Diomedeidae); holotype male (BM), allotype Present study Gmelin, 1789 female (BM), 7 male, 7 female paratypes. Sep­ Procellariidae tember 17, 1963, South Georgia Island, H. B. Macronectes giganteus Present study Clagg. Paratypes deposited: AMNH, BM, LAS, (Gmelin) 1789 SEA, UGA, ZSBS. Brephosceles diomedei Atyeo and Peterson, Additional material. Diomedeidae: 7 males, 8 1970, Pacific Ins. Monog., 23:121-151. females from Diomedea melanophris, South Closely related to Brephosceles margmt­ Georgia Island; 1 male, 1 female from Diomedea ventris, the males of this species can be exulans, South Georgia Island. 128 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

200u

Figs. 78-81. Brephosceles disjunctus, new species, male (holotype): dorsal aspect (78); ventral aspect (79); female (allo­ type): dorsal aspect (80); ventral aspect (81). A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 129

Remarks. The drawings of the male and fe­ FEMALE (allotype). Length, 329u; width, 104u. male are from the holotype and allotype respec­ Dorsal idiosoma: Propodosomal shield 86u in tively. The species is named for the type host. length, ?Ou in width; without lacunae; without external vertical setae; distance between scap­ HOSTS ular setae, 57u. Setae 11 setiform, 8u in length, Diomedeidae positioned on well developed humeral shield; Diomedea chrysostoma Present study setae sh 10u in length, at same level as setae h. Atyeo and Peterson, 1970 Hysterosomal shield 202u in length, ?Ou in width; Forster, 1785 anterior margin straight. Hysterosomal terminus Diomedea exulans L. Present study weakly bilobate, without terminal lamellae. Ven­ 1758 tral idiosoma: Surface fields well developed. Diomedea melanophris Present study Preanal apodeme crescentric. Temminck, 1828 Type material. From Pterodroma leucoptera hypoleuca (Procellariidae): holotype male (SIU), Brephosceles disjunctus, new species allotype female (SIU), 3 male, 4 female para­ types, December 17, 1959, Midway Island, J. C. The males of this new species can be sepa­ Downey. Paratypes deposited: AMNH, SEA, rated from the related Brephosceles lanceolatus UGA. by the condition of the pregenital apodeme, the Additional material (Procellariidae): 1 male, 1 length of the genital organ and the presence of female from Pterodroma cookii de fileppiana, a well developed surface field IV. In B. disjunc­ 155 miles Southeast Talara, Peru; 1 male, 1 tus the genital organ extends to the level of female from Pterodroma leucoptera masafuerae, setae C2, the pregenital apodeme is not fused Chile; 1 male, 1 female from Pterodroma externa to epimerite IVa forming an inverted V, and sur­ cervica/is, Sunday Island, Kermadec Islands. face field IV is well developed. Males of B. Remarks. The drawings of the male and fe­ lanceolatus have a minute genital organ, the male are from the holotype and allotype respec­ surface field IV moderately developed, and epi­ tively. The species is named for the pregenital merites IVa fused with the pregenital apodeme. apodeme which is not fused to epimerites IVa. MALE (holotype). Length, excluding postlam­ ellae, 306u; width, 112u. Dorsal idiosoma: Pro­ HOSTS podosomal shield 91 u in length, 62u in width; Procellariidae without lacunae; without external vertical setae; Pterodroma leucoptera Present study distance between external scapular setae, 52u. hypoleuca (Salvin) 1888 Scapular shields well developed. Setae 11 seti­ Pterodroma leucoptera masa- Present study form, 8u in length, positioned on well developed fuerae Lonnberg, 1921 humeral shield; setae sh 18u in length, approxi­ Pterodroma cookii de Present study mate to seta h. Hysterosomal shield 182u in fileppiana (Giglioli and length, 62u in width; anterior margin straight. Salvadori) 1869 Hysterosomal lobes 62u in length; setae d3 posi­ Pterodroma externa cervicalis Present study tioned at posterior one-third of lobe, at same (Salvin) 1875 level as setae 13 ; setae d4 and pai approximate. lnterlobar and postlobar lamellae well devel­ Brephosceles lanceolatus, new species oped. All legs subequal. Ventral idiosoma: In addition to the combination of characters Epimerites I fused in a Y; surface fields of epi­ mentioned in the former species, Brephosceles merites I and II fused to form a ventral shield. lanceolatus can easily be distinguished by the Pregenital apodeme free, not fused to epimer­ hysterosomal chaetotaxy of the female. In this ites IVa. Genital organ 8u in length, extending species, setae da is absent and setae d4 is lance­ to level of setae c2 ; subgenital shield weakly olate in contrast to the related B. disjunctus in

developed. Adanal shield weakly developed; which setae d 3 is present and setae d4 is seti­ without seta a. form. 130 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

ISOu

Figs. 82-85. Brephosceles lanceolatus, new species, male (holotype): dorsal aspect (82); ventral aspect (83); female (allo­ type): dorsal aspect (84); ventral aspect (85). A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 131

MALE (holotype). Length, excluding post­ Brephosceles bilobatus, new species lamellae, 277u; width, 109u. Dorsal idiosoma: Of the male species which have the surface Propodosomal shield 70u in length, 62u in width; fields of epimerites I and II fused to form a ven­ without lacunae; without external vertical setae; tral shield, this new species can be separated distance between external scapular setae, 57u. by the fusion of the pregenital apodeme with Scapular shields well developed. Setae '1 seti­ epimerites IVa and by the total length (304u). form, 5 u in length, positioned on well developed In addition, the females of this new species humeral shield; setae sh 13u in length, slightly have a small but distinctly bilobate hysterosomal posterior to setae h. Hysterosomal shield 176u terminus. in length, 62u in width; anterior margin straight. MALE (holotype). Length, excluding post­ Hysterosomal lobes 91 u in length; setae d3 posi­ lamellae, 304u; width, 91 u. Dorsal idiosoma: tioned on posterior one-fourth of lobe, separated Propodosomal shield 78u in length, 60u in width; from pai by 10u. Setae d4 extremely small, pos­ without lacunae; without external vertical setae; terior to setae pai. lnterlobar, postlobar and distance between external scapular setae, 47u. extralobar lamellae well developed. All legs sub­ Scapular shields well developed. Setae '1 seti­ equal. Ventral idiosoma: Epimerites I fused in a form, 8u in length, positioned on well developed Y; surface fields of epimerites I and II fused to humeral shield; setae sh 13u in length, approxi­ form a ventral shield. Pregenital apodeme free, mate to setae h. Hysterosomal shield 182u in fused to epimerites IVa. Genital organ au in length, 57u in width; anterior margin straight. length extending beyond setae c2; subgenital Hysterosomal lobes 99u in length; setae d3 posi­ shield present. Adanal shield weakly developed; tioned on posterior one-third of lobe, separated without seta a. from pai by 13u. Setae d4 long and setiform. FEMALE (allotype). Length, 314u; width, 106u. lnterlobar and postlobar lamellae well de­ Dorsal idiosoma: Propodosomal shield 91 u in veloped. All legs subequal. Ventral idiosoma: length, 70u in width; without lacunae; without Epimerites I fused in a Y; surface fields of epi­ external vertical setae; distance between scap­ merites I and II fused to form a ventral shield. ular setae, 67u. Setae '1 spiculiform, 16u in Pregenital adopeme free. Genital organ 8u in length, positioned on well developed humeral length, extending beyond setae c2; subgenital shield; setae sh 21 u in length, approximate to shield absent. Adanal shield weakly developed; seta h. Hysterosomal shield 163u in length, without seta a. 65u in width; anterior margin straight; setae d4 FEMALE (allotype). Length, 332u; width, 93u. lanceolate. Hysterosomal terminus weakly bilo­ Dorsal idiosoma: Propodosomal shield 88u in bate; without terminal lamellae. Ventral idio­ length, 65u in width; without lacunae; without soma: Surface fields moderately developed. external vertical setae; distance between scap­ Preanal apodeme reduced in size. ular setae, 49u. Setae '1 setiform, 8u in length, Type material. From Oceanodroma castro positioned on well developed humeral shield; bangsi (Hydrobatidae): holotype male (USNM), setae sh 10u in length, approximate to seta h. allotype female (USNM), 1 male, 4 female para­ Hysterosomal shield 202u in length, 60u in width; types, April 4, 1891, Galapagos Islands, C. H. anterior margin straight. Hysterosomal terminus Townsend. Paratypes deposited: UGA, USNM. with lobes, without terminal lamellae. Ventral Remarks. The drawings of of the male and idiosoma: Surface fields moderately developed. female are from the holotype and allotype re­ Preanal apodeme absent. spectively. The species is named for the lance­ Type material. From Puffinus puffinus (Procel­ olate setae d4. lari idae): holotype male (USNM), allotype female (USNM), 2 male, 3 female paratypes, 1963, HOSTS Johnston Island, North Pacific, A. B. Amerson. Hydrobatidae Paratypes deposited: USNM, UGA. Oceanodroma castro bangsi Present study Remarks. The drawings of the male and fe­ Nichols, 1914 male are from the holotype and allotype respec- 132 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

ZOOu

Figs. 86-89. Brephosceles bilobatus, new species, male (holotype): dorsal aspect (86); ventral aspect (87); female (allo­ type): dorsal aspect (88); ventral aspect (89). A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 133 tively. The spceies is named for the bilobate Type material. From Puffinus pacificus chloro­ hysterosomal terminus of the female. hynchos (Procellariidae); holotype male (USNM), allotype female (USNM), 6 male paratypes, July HOSTS 15, 1962, Eastern Island, Midway Atoll, Central Procellari idae Pacific, H. I. Fisher. Paratypes deposited: AMNH, Puffinus puffinus (Brunnich) Present study Gaud, LAS, SEA, SIU, UGA. 1764 Additional material. Procellariidae: 1 male, 2 females from Puffinus gravis, Canada; 1 male Brephosceles parvatus, new species from Puffinus puffinus, Central Pacific; 1 male from Puffinus tenuirostris, Alaska. Unique to group Ill, Brephosceles parvatus is Remarks. The drawings of the male and fe­ the smallest species in total dimensions. How­ male are from the holotype and allotype respec­ ever, the conformation of the epimerites, the tively. free pregenital apodeme and the setiform con­ HOSTS dition of setae '1 clearly establish a close Procellari idae relationship with other species of the group. Puffinus pacificus chloro­ Present study MALE (holotype). Length, excluding post­ hynchos Lesson, 1831 lamellae, 261 u; width, 75u. Dorsal idiosoma: Puffinus puffinus (Brunnich) Present study Propodosomal shield 73u in length, 52u in width; 1764 without lacunae; without external vertical setae; Puffinus gravis (O'Reilly) Present study distance between external scapular setae, 34u. 1818 Scapular shields well developed. Setae '1 seti­ Puffinus tenuirostris Present study form, Su in length, positioned on well developed (Temminck) 1835 humeral shield; setae sh Su in length, slightly posterior to seta h. Hysterosomal shield 156u in Group IV-the lambda group length, 49u in width; anterior margin straight. Ectoparasitic on the Anseriformes, this group Hysterosomal lobes 65u in length; setae d3 posi­ is recognized primarily by the greatly shortened tioned at posterior one-third of lobe; setae d4 hysterosomal lobes and by the truncate appear­ and pai approximate. lnterlobar and postlobar ance of these lobes distally. All species have lamellae well developed; extralobar lamellae ab­ weakly developed surface fields and coxal fields sent. All legs subequal .Ventral idiosoma: Epi­ IV closed. Coxal field Ill is open or closed de­ merites I fused in a Y; surface fields fused to pending upon the species. form a ventral shield. Pregenital apodeme ap­ Based on characters of the coxosternal parently fused anteriorly, genital discs present skeleton, it is believed that group IV and the on anterior projections. Genital organ Su in following group V are evolutionary intermediates length, extending beyond setae c2 ; subgenital between group VI (Charadriiformes) and groups shield present. Adanal shield weakly developed; II and 111 (Procellariiformes). without seta a. Pertinent characters for group recognition, FEMALE (allotype). Length, 298u; width, 86u. males: Dorsal idiosoma: Propodosomal shield 78u in 1. Coxal field IV closed; coxal field Ill open length, 55u in width; without lacunae; without or closed. external vertical setae; distance between scap­ 2. Surface fields weakly developed. ular setae, 42u. Setae 11 setiform, Su in length, 3. Hysterosomal lobes truncate in appear- positioned on well developed humeral shield; ance and reduced in length. setae sh Bu in length, approximate to seta h. 4. Extralobar lamellae absent. Hysterosomal shield 182u in length, 52u in width; 5. Setae '1 setiform.

anterior margin straight. Hysterosomal terminus 6. Setae d3 approximate to setae pai. entire, without terminal lamellae. Ventral idio­ Pertinent characters for group recognition, soma: Surface fields moderately developed. females: Preanal apodeme present. 1. Pregenital apodeme free. 134 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

l50u

Figs. 90-93. Brephosceles parvatus, new spceies, male (holotype): dorsal aspect (90); ventral aspect (91); female (allo­ type): dorsal aspect (92); ventral aspect (93). A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 135

2. Hysterosomal terminus bilobate or entire. teromedial margin of well developed humeral 3. Setae C2 posterior to genital discs. shield; setae sh lanceolate, 13u in length, pos­ 4. Hysterosomal pore at level of setae da. terior to setae h. Hysterosomal shield 231 u in Key to the species of group IV length, 78u in width; anterior margin straight. (lambda group) Hysterosomal lobes 78u in length; setae d3 posi­ 1. Male with adanal shields fused medially; tioned at midlength of lobe, at same level as hysterosomal cleft absent...... setae la, setae d4 at terminus of lobe, approxi­ ...... lambda (Trouessart) p. 135 mate to setae pai. lnterlobar and postlobar lam­ Male with adanal shields separate...... 2 ellae well developed; extralobar lamellae absent. 2. Genital organ extending to or beyond All legs subequal. Ventral idiosoma: Epimerites level of setae c2; coxal fields Ill and I fused in a Y; surface fields weakly developed. IV closed; female terminus distinctly Pregenital apodeme free; coxal field Ill nearly bilobate ...... anhimae, n. sp., p. 138 closed, IV closed. Genital organ 16u in length, Genital organ not extending beyond not extending beyond setae c2; subgenital shield setae c2; coxal field Ill closed, IV open 3 present. Adanal shields fused, notjoined to pre­ genital apodeme. 3. Male with setae 11 spiculiform ...... anatina Dubinin p. 138 FEMALE (syntype) .Length, 461u; width, 145u . Male with setae '1 setiform ...... Dorsal idiosoma: Propodosomal shield 101 u in ...... discidicus, n. sp., p. 135 length, 91u in width; without lacunae; without external vertical setae; setae se included on Brephosceles lambda (Trouessart) shield, separated from each other by 83u. Setae Pterolichus lambda Trouessart, 1885, Bull. Soc. '1 spiculiform, Su in length, positioned on well Etud. Sci. Angers, 14: 72. Type host: Nettapus developed humeral shield; setae sh lanceolate, auritus (Anatidae). 13u in length, posterior to seta h. Hysterosomal Alloptes lambda, Canestrini and Kramer, 1899, shield 301u in length, 75u in width; anterior mar­ Tierreich, 7: 113-114. gin straight; setae d3 and d4 in trapezoidal ar­ Alloptes lambda, Gaud, 1952, Mem. Inst. Sci. rangement, separated by 39u. Hysterosomal Madagascar, Ser. A, 7(1): 87. terminus distinctly bilobate, with terminal lam­ Alloptes lambda, Radford, 1953, Parasitol., 43 ellae. Ventral idiosoma: Surface fields weakly (3,4): 213. developed. Preanal apodeme absent. Alloptes lambda, Radford, 1958, Rev. Brasil. Ent., Type material. From Nettapus auritus (Anati­ 8: 118. dae): lectotype male {TC), 1 male, 2 female Brephosceles lambda, Peterson and Atyeo, Bull. syntypes (TC), Malagasy Republic. Univ. Nebraska St. Mus., 8(4): 220. Additional material. (Anatidae): 9 males, 6 fe­ The male of this long known species is dis­ males from Nettapus auritus, Malagasy Republic. tinguished from the related Brephosceles dis­ Remarks. The drawings of the male and fe­ cidicus by the shape of the adanal shield and male are from the lectotype and syntype respec­ the truncate appearance of the hysterosomal tively. lobes. The adanal shields are fused medially in HOSTS B. lambda whereas in the following species the Anatidae adanal shields are separate. The females are Nettapus auritus (Boddaert) Trouessart, 1885 characterized as distinctly bilobate in B. lambda, 1783 Present study and weakly bilobate in B. discidicus. MALE (lectotype). Length, excluding postlam­ Brephosceles discidicus, new species ellae, 397u ;width, 147u. Dorsal idiosoma: Pro­ Related to Brephosceles lambda, the males of podosomal shield 117u in length, 83u in width; this new species are characterized by separate without lacunae; without external vertical setae; adanal shields and by a complete supranal cleft. distance between external scapular setae, 75u. The hysterosomal termini of the females are Scapular shields well developed. Setae '1 seti­ weakly bilobate, separating this species from B. form 21 u in length, positioned on extreme an- lambda and the following species, B. anatina. 136 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

200u

I . I . . . . ~-- ' . : : . ~-

Figs. 94-97. Brephosceles lambda (Trouessart), male (lectotype): dorsal aspect (94); ventral aspect (95); female (syntype): dorsal aspect (96); ventral aspect (97). A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 137

200u 99

Figs. 98-101. Brephosceles discidicus, new species, male (holotype): dorsal aspect (98); ventral aspect (99); female (allotype): dorsal aspect (100); ventral aspect (101). 138 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

MALE (holotype). Length, excluding post­ Brephosceles anatina Dubinin lamellae, 306u; width, 109u. Dorsal idiosoma: Brephosceles anatina Dubinin, 1951, Akad. Nauk Propodosomal shield 91 u in length, 75u in width; S.S.S.R., Zool. Inst., Parazitol. Sborn., 13: 223- without lacunae; without external vertical setae; 224, Fig. 52. Type host: Anas platyrhynchos distance between external scapular setae, 67u. (Anatidae). Scapular shields well developed. Setae 11 seti­ Brephosceles anatina, Peterson and Atyeo, 1968, form, 5u in length, positioned on well developed Bull. Univ. Nebraska St. Mus., 8(4): 218. humeral shield; setae sh setiform, au in length, Due to the absence of type material or speci­ slightly posterior to seta h. Hysterosomal shield mens collected from the type host, the following 171 u in length, 62u in width; anterior margin descriptions have been extrapolated from the straight. Hysterosomal lobes 44u in length; setae original description and illustrations (Dubinin, d3 positioned at midlength of lobe, anterior to 1951). The lanceolate setae 11 and sh distinguish level of setae 13; setae d4 at terminus of lobe, this species from the former two species in approximate to setae pai. lnterlobar and post­ group IV. lobar lamellae well developed, extralobar lamel­ lae absent. All legs subequal. Ventral idiosoma: MALE. Length, 290-300u. Dorsal idiosoma: Epimerites I fused in a Y; surface fields weakly Propodosomal and scapular shields well devel­ developed. Pregenital apodeme free; coxal field oped. Setae 11 lanceolate, positioned on anterior 111 nearly closed, IV closed. Genital organ 13u in margin of humeral shield; setae sh lanceolate posterior to setae h. Hysterosomal lobes shorti length, not extending beyond setae c2 ; subgen­ ital shield present. Adana! shield not fused setae d3 positioned on anterior one-third of lobe· medially, not fused with pregenital apodeme. anterior to level of setae 13 ; setae d 4 at terminu~ of lobe. lnterlobar and postlobar lamellae well FEMALE (allotype). Length, 415u; width, 153u. developed, extralobar lamellae absent. All legs Dorsal idiosoma: Propodosomal shield 96u in subequal. Ventral idiosoma: Epimerites I fused length, fused laterally with scapular shields; !n a Y; surface fields weakly developed. Pregen- without lacunae; without external vertical setae; 1tal apodeme free; coxal field 111 nearly closed, distance between scapular setae, 83u. Setae '1 IV closed. Genital organ extending to level of setiform, 5u in length, positioned on well devel­ setae c2; subgenital shield apparently absent. oped humeral shield; setae sh spiculiform, 10u Adana! shields not fused medially; not joined to in length, slightly posterior to setae h. Hystero­ pregenital apodeme. somal shield 244u in length, 75u in width; an­ FEMALE. Length, 370-380u. Dorsal idiosoma: terior margin shallowly concave; setae d 3 and d 4 in rectangular arrangement, separated by 52u. Hysterosomal terminus weakly bilobate; without Hysterosomal terminus weakly bilobate without terminal lamellae. Ventral idiosoma: Surface terminal lamellae. Ventral idiosoma: Surface fields weakly developed. Preanal apodeme fields weakly developed. Preanal apodeme absent. absent. Type material. From Anas platyrhynchos (Ana­ Type material. From Cygnus bewicki (Anati­ tidae): The type is deposited at the Leningrad dae): holotype male (Gaud), allotype female Academy of Science, Leningrad, Russia. (Gaud), 2 male, 8 female paratypes, October, HOSTS 1953, Zoo Amsterdam, J. Gaud. Paratypes de­ Anatidae posited: Gaud, UGA. Anas platyrhynchos L. Dubinin, 1951 Remarks. The drawings of the male and 1758 female are from the holotype and allotype respectively. Brephosceles anhimae, new species HOSTS The closed coxal fields Ill and IV, the length Anatidae of the genital organ, and the dentate postlobar Cygnus bewicki Yarrel Present study lamellae characterize this species. Well devel­ 1830 oped hysterosomal lobes distinguish the female. A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 139

200u

\ 104 \ Figs. 102-105. Brephosceles anhimae, new species, male (ho lotype): dorsal aspect (102); ventral aspect (103); female (al­ lotype): dorsal aspect (104); ventral aspect (105). 140 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

MALE (holotype}. Length, excluding pest­ HOSTS lamellae, 314u; width, 109u. Dorsal idiosoma: Anhimidae Propodosomal shield 83u in length, 73u in width; Anhima cornuta (L.} Present study without lacunae; without external vertical setae; 1766 distance between external scapular setae, 60u. Group V-the geranoxenus group Scapular shields weakly developed. Setae / 1 Consisting of only two species, this group is setiform, 4u in length, not positioned on humeral distinguished from the proceeding group IV by shield; setae sh setiform, Bu in length, slightly the prolongation of the hysterosomal lobes, the posterior to setae h. Hysterosomal shield 184u closed coxal field IV, and setae d3 which is in length, 67u in width; anterior margin straight. positioned near the supranal cleft. Setae d4 is Hysterosomal lobes 62u in length; setae d3 posi­ approximate to setae pai. tioned at midlength of lobe, anterior to level of Ectoparasitic on two orders of birds, Grui­ la; setae d4 at terminus of lobe, approximate to formes and Charadriiformes, it is probable that setae pai. lnterlobar lamellae well developed, this group is highly artificial and that in fact extralobar lamellae absent. Postlobar lamellae there are actually two species groups involved. with 4 dentations. All legs subequal. Ventral However, after extensive collecting throughout idiosoma: Epimerites I fused in a Y; surface both avian orders, no additional species were fields moderately developed. Pregenital apo­ found applicable to this group. deme free; coxal field Ill and IV closed. Genital Pertinent characters for group recognition, organ 1Bu in length, not extending beyond setae males: c2; subgenital shield absent. Adana! shields not 1. Hysterosomal lobes elongate with supranal fused medially, not fused with pregenital cleft. apodeme. 2. Coxal field IV closed, coxal field Ill open FEMALE (allotype}. Length, 394u; width, 109u. or closed. Dorsal idiosoma: Propodosomal shield 88u in 3. Extralobar lamellae present. length, fused laterally with scapular shield, with­ 4. Setae '1 setiform. out lacunae; without external vertical setae; dis­ 5. Setae d4 approximate to setae pai. tance between scapular setae, 67u. Setae /1 Pertinent characters for group recognition, setiform, 3u in length, not positioned on humeral females: shield; setae sh spiculiform, Bu in length, pos­ 1. Pregenital apodeme free. terior to setae h. Hysterosomal shield 241 u in 2. Setae c3 posterior to genital discs. length, 73u in width; anterior margin shallowly 3. Hysterosomal terminus bilobate. concave; setae d3 and d4 in rectangular arrange­ ment, separated by 67u. Hysterosomal terminus Key to the species of group V distinctly bilobate, without terminal lamellae. (geranoxenus group} Ventral idiosoma: Surface fields moderately de­ 1. Postlobar lamella of male with six to veloped. Preanal apodeme present. seven dentations; setae sh setiform Type material. From Anhima cornuta (Anhimi­ ...... geranoxenus Peterson p. 140 dae}: holotype male (MU}, allotype female (MU}, Postlobar lamella of male without denta- 5 male, 11 female paratypes, December 15, 1940, tions; setae sh lanceolate ...... Andes, Ecuador, collector unknown. Paratypes ...... chaubaudi Gaud p. 142 deposited: AMNH, Gaud, MU, SAIMR, UGA. Brephosceles geranoxenus Peterson Remarks. To date, this is the only species of Brephosce/es geranoxenus Peterson, 1968, Ann. Brephosce/es known from the Anhimidae. Col­ Roy. Mus. Cent. Africa, Zool., 8(164): 80-82. lections from phylogenetically related birds will The dentations on the postlobar lamellae, the undoubtedly produce new species applicable to setiform seta sh and closed coxal fields IV, suffi­ this group. The drawings of the male and female ciently distinguish this species from the related are from the holotype and allotype respectively. Brephosceles chaubaudi. The females are vir­ The species is named for the type host. tually indistinguishable. A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 141

JOOu

108 I Figs. 106-109. Brephosce/es geranoxenus Peterson, male: dorsal aspect (106); ventral aspect (107); female: dorsal aspect (108); ventral aspect (109). 142 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

MALE (holotype). Length, excluding post­ HOSTS lamellae, 485u; width, 158u. Dorsal idiosoma: Gruidae Propodosomal shield 114u in length, 106u in Balearica regulorum Gaud, 1968 width; without lacunae; without external vertical (Bennet) Present study setae; distance between external scapular setae, 96u. Scapular shields moderately developed. Brephosceles chaubaudi Gaud Setae /1 setiform, 16u in length, positioned on Brephosceles chaubaudi Gaud, 1968, Ann. Roy. well developed humeral shield; setae sh seti­ Mus. Cent. Africa, Zool., 8(164): 78-80. form 18u in length, slightly posterior to setae h. This species is separated from the related Hysterosomal shield 316u in length, 104u in Brephosceles geranoxenus by the lanceolate width; anterior margin straight. Hysterosomal setae sh, the open coxal field IV and the margin lobes 112u in length; setae d 3 positioned on of the postlobar lamellae which is entire. Addi­ extreme anterior margin of lobe, anterior to tionally, a small pore is located on the pregen­ setae /3; setae d4 at terminus of lobe, approxi­ ital apodeme of the male, at the level of setae C3. mate to setae pai. lnterlobar and extralobar MALE. Length, excluding postlamellae, 474u; lamellae well developed. Postlobar lamellae with width, 169u. Dorsal idiosoma: Propodosomal 6-7 dentations. All legs subequal. Ventral idio­ shield 112u in length, 93u in width; without soma: Epimerites I fused in a Y; surface fields lacunae; without external vertical setae; dis­ poorly developed. Pregenital apodeme free; tance between external scapular setae, 88u. coxal field 111 and IV closed. Genital organ 18u Scapular shields well developed. Setae 11 seti­ in length, not extending beyond setae c2; sub­ form, 1Ou in length, positioned on well devel­ genital shield present. Adanal shields not fused; oped humeral shield; setae sh lanceolate, 18u in not joined to pregenital apodeme. length, posterior to setae h. Hysterosomal shield FEMALE (allotype). Length, 482u; width, 150u. 293u in length, 73u in width; anterior margin Dorsal idiosoma: Propodosomal shield 117u in straight. Hysterosomal lobes 137u in length; length, 104u in width; without lacunae; without setae d3 positioned on anterior one-third of lobe, external vertical setae; distance between scap­ anterior to level of setae 13 ; setae d4 at terminus ular setae, 96u. Setae /1 setiform, 13u in length, of lobe, approximate to setae pai. lnterlobar and positioned on well developed humeral shield; postlobar lamellae well developed. Extralobar setae sh spiculiform 13u in length, slightly pos­ lamellae apparently absent. All legs subequal. terior to setae h. Hysterosomal shield 298u in Ventral idiosoma: Epimerites I fused in a Y; sur­ length, 81 u in width; anterior margin shallowly face fields weakly developed. Pregenital apo­ concave; setae d 3 and d4 in trapezoidal arrange­ deme free; coxal field Ill nearly closed, IV ment; separated by 80u. Hysterosomal terminus closed. Genital organ 8u in length, not extend­ distinctly bilobate; without terminal lamellae. ing beyond setae c3 ; subgenital shield absent. Ventral idiosoma: Surface fields weakly devel­ Adanal shields not fused medially, not con­ oped. Preanal apodeme absent. nected to pregenital apodeme. Type material. From Balearica regulorum FEMALE. Length, 482u; width, 163u. Dorsal (Gruidae): holotype male (AMNH), allotype fe­ idiosoma: Propodosomal shield 119u in length, male (AMNH), 3 male, 4 female paratypes; June 109u in width; without lacunae; without external 3, 1929, Nyassaland, R. Boulton and L. Boulton: vertical setae; distance between scapular setae,

Paratypes: 2 male, 1 female, October, 1962, 91 u. Setae 11 setiform, 8u in length, positioned Lochinvar, North Rhodesia, F. Zumpt. Paratypes on anterior margin of well developed humeral deposited: Gaud, UGA, USNM. shield; setae sh lanceolate, 18u in length, pos­ Remarks. The drawings of the male and terior to seta h. Hysterosomal shield 285u in female are from the holotype and allotype length, 65u in width; anterior margin slightly respectively. concave; setae d3 and d4 in rectangular arrange- A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 143

Figs. 110-113. Brephosceles chaubaudi Gaud, male: dorsal aspect (110); ventral aspect (111); female: dorsal aspect (112); ventral aspect (113). 144 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM ment, separated by 44u. Hysterosomal terminus Pertinent characters for group recognition, distinctly bilobate, with short terminal lamellae. males: Ventral idiosoma: Surface fields weakly devel­ 1. Pregenital apodeme trapezoidal and free oped. Preanal apodeme absent. (Haematopodidae); trapezoidal and fused Type material. From Burhinus vermiculatus (Charadriidae). (Burhinidae): holotype male (Gaud), allotype 2. All surface fields weakly developed. female (Gaud), 3 male, 3 female paratypes 3. Coxal fields I-IV open. (Gaud), February, 1958, Yaounde, Cameroun, J. 4. Setae d4 approximate to setae pai. Mouchet. 5. Setae sh posterior to setae h. Material examined. Burhinidae: 8 males, 2 Pertinent characters for group recognition, females from Orthorhamphus magnirostris, Aus­ females: tralia; 6 males, 5 females, Philippines; 2 males, 1. Pregenital apodeme free. 4 females from Esacus recurivirostris, Thailand. 2. Setae C2 posterior to genital discs. Remarks. The redescription and drawings are 3. Hysterosomal terminus distinctly bilobate. based on specimens from Orthorhamphus mag­ nirostris. To date, this is the only species of Key to the species of group VI Brephosceles host specific at the family level. (chardrii group) HOSTS 1. Male with pregenital apodemes fused Burhinidae medially forming a trapezoid (Chara- Burhinus vermiculatus Gaud, 1968 driidae) ...... 3 (Cabanis) 1868 Present study Male with pregenital apodemes free Orthorhamphus magnirostris Present study (Haematopodidae) ...... 2 (Vieillot) 1818 2. Female with primary spermathecal duct Esacus recurvirostris Present study expanded apically; secondary ducts (Cuvier) 1829 13u in length ...... turgidus, n. sp., p. 150 Female with primary spermathecal duct Group VI-the charadrii group narrow, not expanded apically; sec- This assemblage of seventeen species com­ ondary ducts 16u in length ...... prises the largest most homogeneous group of ...... haematopi, n. sp., p. 148 the genus Brephosceles. All species are re­ 3. Pregenital apodeme of male with ante­ stricted to the avian order Charadriiformes, rior margin deeply concave, genital representing two families, Charadriidae and organ extending beyond level of setae

Haematopodidae. Within each respective bird c2 ; female with distinct reticular pat- family the males are distinctive. Ectoparasitic tern ...... afribycis, n. sp., p. 153 males on Haematopodidae have the pregenital Pregenital apodeme of male with ante­ apodeme trapezoidal and free (Fig. 120); para­ rior margin straight or shallowly con- sitic males from Charadriidae have a pregenital cave ...... 4 apodeme that is trapezoidal and fused anteri­ 4. Pregenital apodeme with bases widely orly (Fig. 115). Other than the characterization separated; genital organ not extend- stated above, the males of each of these sub­ ing beyond setae C2 ...... groups are impossible to differentiate and over ...... stephanibycis, n. sp., p. 150 1,649 males have been examined during the Pregenital apodeme of male with bases course of this study. A correlation of males and not widely separated; with or without females within a particular species can only be a fusion to the adanal shields...... 5 achieved by using the host bird since the major­ 5. Female with a sclerotized bulbous en­ ity are species specific. Therefore, although a largement of the prmiary spermathec- holotype female and allotype male are desig­ al duct ...... 6 nated, a formal description is given only for Female without bulbous enlargement of males of B. charadrii. primary spermathecal duct...... 7 A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 145

6. Hysterosomal shield of female with dis­ Hysterosomal shield without longitudinal tinct longitudinal striations, with faint striations ...... 14 reticular pattern; without dorsolateral 14. Hysterosomal shield with lateral margins finger-like projections ...... deeply incised at level of setae d3 ...... longistriatus, n. sp., p. 157 ...... constrictus, n. sp., p. 157 Hysterosomal shield of female without Hysterosomal shield not deeply incised

longitudinal striations, with reticular at level of setae d3 •••••.•••••••••••••••••••••••••• 15 pattern, with dorsolateral finger-like 15. Hysterosomal shield without reticular projections ...... tuberi, n. sp., p. 155 pattern; distance between setae d3 7. Female with distinct dorsolateral finger­ and d4 39u; primary spermathecal like projections on hysterosomal duct tapering distally ...... shield ...... 8 ...... inornatus, n. sp., p. 158 Female without finger-like projections Hysterosomal shield with reticular pat- on hysterosomal shield...... 11 tern ...... 16 8. Primary spermathecal duct expanded 16. Hysterosomal shield 267u in length; dis­ proximally and sharply tapering dis­ tance between setae d3 and d4 52u tally; without reticular pattern on hys- (Belonopterus) .... chilensis, n. sp., p. 161 terosomal shield (Hoplopterus) ...... Hysterosomal shield 246u in length; dis­ ...... hoplopteri, n. sp., p. 158 tance between setae d3 and d4 52u Primary spermathecal duct not ex- (Charadrius) .... charadrii Dubinin, p. 145 panded ...... 9 9. Secondary spermathecal ducts with two Brephosce/es charadrii Dubinin bulbous enlargements at the proximal Brephosceles charadrii Dubinin, 1951, Akad. and distal ends (Lobivanel/us) ...... Nauk. S.S.S.R., Zool. Inst., Parasitol. Sborn., ...... /obivanelli, n. sp., p. 162 13: 225-226, Fig. 53. Type host: Charadrius Secondary spermathecal ducts without dubius. C. p/acidus (Charadriidae). expansions ...... 10 Brephosceles charadrii, Gaud and Mouchet, 10. Hysterosomal shield of female with 1957. Ann. Parasitol. hum. comp., 32(5-6): 497- setae d4 and / in a straight line ...... 3 498 ...... collaricus, n. sp., p. 162 Hysterosomal shield of female with Brephosce/es charadrii, Radford, 1958. Rev. Brasil Entomol., 8: 126-127. setae d4 posterior to /3 •••••••••••••.•••••••..•...... virginiensis, n. sp., p. 161 Brephosceles charadrii, Gaud and Till, 1961 . 11. Secondary spermathecal ducts originat- Publ. S. Afr. Inst .Med. Res., II (L): 241. ing from primary spermathecal duct.... 12 Brephosceles charadrii, Peterson and Atyeo, Secondary spermathecal ducts originat- 1968. Bull. Univ. Nebraska St. Mus., 8(4): 218. ing from spermathecal sac ...... 13 This species is most recognizable in the fe­ 12. Secondary ducts 5u in length; primary male by the reticular pattern and the absence duct extremely broad; distance be- of dorsolateral finger-like projections on the tween setae d3 and d4 39u ...... hysterosomal shield. The spermatheca is sac­ ...... vane/Ii Gaud, p. 154 like with the primary duct not expanded prox­ Secondary ducts greater than 10u in imally; secondary ducts are simple, 13u in length; distance between setae d3 and length. d4 52u ...... MALE. Length, excluding postlamellae, 405u; .... histionotus Gaud and Mouchet, p. 160 width, 156u. Dorsal idiosoma: Propodosomal 13. Hysterosomal shield with longitudinal shield 99u in length, 96u in width; without la­ striations; primary spermathecal duct cunae; without external vertical setae; distance tapering distally (Belonopterus) ...... between external scapular setae, 99u. Scapular ...... belonopteri, n. sp:, p. 155 shields well developed. Setae '1 lanceolate, 1Ou 146 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

250u 50u

Figs. 114-118. Brephosce/es charadrii Dubinin, male: dorsal aspect (114); ventral aspect (115); female: dorsal aspect (116); spermatheca and associated ducts (117); ventral aspect (118). A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 147

250u SOu

Sp

WM

Figs. 119-123. Brephosce/es haematopi, new species, male (holotype): dorsal aspect (119); ventral aspect (120); female allotype): dorsal aspect (121); spermatheca and associated ducts (122); ventral aspect (123). 148 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM il"I length, positioned on anterior margin of well HOSTS developed humeral shield; setae sh spiculiform, Charad ri idae 16u in length, posterior to setae h. Hystero­ Charadrius dubius curonicus Present study somal shield 226u in length, 88u in width; ante­ Gmelin, 1789 rior margin strongly concave. Hysterosomal Charadrius dubius dubius Dubinin, 1951 lobes 86u in length, setae d3 positioned near the Scopoli, 1786 Present study apex of lobe, anterior to setae 13 ; setae d4 at Charadrius dubius jerdoni Present study terminus of lobe, approximate to setae pai. ln­ (Legge} 1880 terlobar, postlobar, and extralobar lamellae well Charadrius placidus Dubinin, 1951 developed. All legs subequal. Ventral idiosoma: Gray and Gray, 1863 Present study Epimerites I fused in a Y; surface fields weakly developed. Pregenital apodeme fused, 49u in Brephosceles haematopi, new species length, 62u in width; coxal fields Ill and IV open. This new species can most easily be dis­ Genital organ 18u in length, extending to the tinguished from the following species, Bre­ level of setae c2; subgenital shield present. phosceles turgidus, by the morphology of the Adanal shields not fused medially, not joined to spermatheca and spermathecal duct. The fe­ pregenital apodeme. males of B. haematopi have a spermatheca FEMALE. Length, 410u; width, 137u. Dorsal that is small and sac-like; the duct lacks an idiosoma: Propodosomal shield 96u in length, apical enlargement. In contradistinction, females 96u in width; without lacunae; without external of B. turgidus have a large tubular spermatheca vertical setae; distance between external scap­ and the primary duct is enlarged apically. ular setae 101 u. Setae si 18u in length, sepa­ MALE (holotype}. Length, excluding post­ rated by 78u. Setae '1 lanceolate, 10u in length, lamellae 446u; width, 145u. Dorsal idiosoma: positioned on well developed humeral shield; Propodosomal shield 117u in length, 101 u in setae sh spiculiform, 16u in length, posterior to width; without lacunae; without external vertical setae h. Hysterosomal shield 246u in length, 78u setae; distance between external scapular setae, in width; anterior margin shallowly concave; 96u. Scapular shields well developed. Setae 11 distance between setae d3 and d4 52u; with faint setiform, 11 u in length, positioned on anterior reticular pattern; without dorsolateral finger-like margin of well developed humeral shield; setae projections. Primary spermathecal duct not ex­ sh lanceolate, 13u in length, posterior to setae panded proximally, not tapering posteriorly; h. Hysterosomal shield 277u in length, 78u in secondary ducts simple, 13u in length. Hystero­ width; anterior margin straight. Hysterosomal somal terminus distinctly bilobate; without well lobes 99u in length, setae d3 positioned near developed terminal lamellae. Ventral idiosoma: apex of lobe, anterior to level of setae 13 ; setae Surface fields weakly developed. Preanal apo­ d4 at terminus of lobe, approximate to setae pai. deme absent. lnterlobar, postlobar, and extralobar lamellae Type material. From Charadrius dubius (Cha­ well developed. All legs subequal. Ventral idio­ radriidae}: The primary and secondary types are soma: Epimerites I fused in a Y; surface fields deposited at the Leningrad Academy of Science, weakly developed. Pregenital apodeme free; U.S.S.R. coxal fields Ill and IV open. Genital organ 16u in length, not extending beyond setae c2; sub­ Material examined. (Charadriidae}: 43 males, genital shield present. Adanal shields separate, 38 females from Charadrius dubius curonicus, not joined to pregenital apodeme. China; 5 males, 2 females from Charadrius du­ FEMALE (allotype}. Length, 477u; width, 166u. bius jardoni, Thailand, 2 males, 4 females from Dorsal idiosoma: Propodosomal shield 125u in Charadrius placidus, Japan. length, 119u in width; without lacunae; without Remarks. The drawings of the male, female, external vertical setae; distance between scap­ and spermatheca are from Charadrius dubius ular setae, 112u. Setae 11 setiform, 16u in length, curonicus. positioned on well developed humeral shield; A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 149

250u 50u

Sp

WM

V ·r 127

Figs. 124-128. Brephosceles turgidus, new species, male (holotype): dorsal aspect (124); ventral aspect (125); female allotype): dorsal aspect (126); spermatheca and associated ducts (127); ventral aspect (128). 150 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM setae sh lanceolate, 16u in length, posterior to well developed. All legs subequal. Ventral idio­ setae h. Hysterosomal shield 298u in length, 78u soma: Epimerites I fused in a Y; surface fields in width; anterior margin shallowly concave; weakly developed. Pregenital apodeme free; setae d3 and d4 separated by 75u. Hysterosomal coxal fields Ill and IV open. Genital organ 16u terminus distinctly bilobate, without well devel­ in length, not extending beyond setae c2 ; sub­ oped terminal lamellae, without lacunae, sperm­ genital shield present. Adana! shields separate, athecal duct not enlarged near sac; secondary well developed. ducts 16u in length. Ventral idiosoma: Surface FEMALE (allotype). Length, 498u; width, 182u. fields weakly developed. Preanal apodeme Dorsal idiosoma: Propodosomal shield 125u in absent. length, 127u in width; without lacunae; without Type material. From Haematopus ostralegus external vertical setae; distance between scap­ longirostris (Haematopodidae): holotype male ular setae, 114u. Setae 11 setiform, 21u in length, (USNM), allotype female (USNM), 2 male, 10 positioned on anterior margin of well developed female paratypes, July 15, 1948, Arnhem Penin­ humeral shield; setae sh lanceolate, 18u in sula, Northern Territory, Australia, H. G. Deig­ length, posterior to setae h. Hysterosomal shield nan. Paratypes deposited: Gaud, LAS, SEA, 295u in length, 91 u in width; anterior margin UGA, ZSZM. shallowly concave; setae d3 and d4 separated by Remarks. The drawings of the male and fe­ 73u. Hysterosomal terminus distinctly bilobate, male are from the holotype and allotype respec­ without well developed terminal lamellae, with­ tively. The illustration of the spermatheca is from out lacunae, spermathecal duct enlarged near a paratype. The species is named for the host. sac; secondary ducts 13u in length. Ventral idio­ soma: Surface fields weakly developed. Preanal HOSTS apodeme absent. Haematopodidae Type material. From Haematopus palliatus Haematopus ostralegus longi- Present study palliatus (Haematopodidae): holotype male (NU), rostris Vieillot, 1817 allotype female (NU), 5 male, 2 female para­ types, June 26, 1963, Beaufort, Carteret County, Brephosceles turgidus, new species North Carolina, G. Leidahl. Paratypes deposited: The enlargement of the spermathecal duct AMNH,RNH, UGA,ZSZM. and the absence of a reticular pattern on the Remarks. The drawings of the male and fe­ hysterosomal shield is distinctive for this new male are from the holotype and allotype respec­ species; males are virtually indistinguishable tively. The illustration of the spermatheca is from from B. haematopi except for setae sh which is a paratype. The species is named for the prox­ setiform in B. turgidus, lanceolate in B. hae­ imal swelling on the primary spermathecal duct. matopi. MALE (holotype). Length, excluding post­ HOSTS lamellae, 464u; width, 169u. Dorsal idiosoma: Haematopodidae Propodosomal shield, 122u in length, 106u in Haematopus palfiatus palfiatus Present study width; without lacunae; without external vertical Temminick, 1920 setae; distance between external scapular setae, 99u. Scapular shields well developed. Setae 11 Brephosceles stephanibycis, new species setiform, 13u in length, positioned on anterior Although closely related to Brephosceles afri­ margin of well developed humeral shield; setae bycis, this new species can be distinguished in sh setiform, 13u in length, posterior to setae h. the male by the length of the genital organ and

Hysterosomal shield 280u in length, 104u in in the female by the distance between setae d 3 width; anterior margin shallowly concave. Hys­ and d4. The male genital organ in B. stephani­ terosomal lobes 106u in length; setae d3 posi­ bycis does not extend beyond the insertions of tioned on apex of lobe, anterior to setae 13 ; setae setae C2, the distance between setae d3 and d4 d4 at terminus of lobe, approximate to setae pai. in the females is: B. stephanibycis, 36u; B. afri­ I nterlobar, postlobar and extralobar lamellae bycis, 65u. A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 151

2SOu 50u

Sp

WM

132

Figs. 129-133. Brephosceles stephanibycis, new species, male (holotype): dorsal aspect (129); ventral aspect (130); fe­ male (allotype): dorsal aspect (131); spermatheca and associated ducts (132); ventral aspect (133). 152 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

Figs. 134-138. Brephosce/es afribycis, new species, male (holotype): dorsal aspect (134); ventral aspect (135); female (al­ lotype): dorsal aspect (136); spermatheca and associated ducts (137); ventral aspect (138). A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 153

MALE (holotype). Length, excluding post­ spectively. The spermatheca is from a paratype. lamellae, 407u; width, 161u. Dorsal idiosoma: The species is named for the host. Propodosomal shield 99u in length, 119u in width; without lacunae; without external vertical HOSTS setae; distance between external scapular setae, Stephanibyx melanopterus Present study 103u. Scapular shields well developed. Setae 11 (Cretzschmar) 1829 spiculiform, 10u in length, positioned on anterior Stephanibyx coronatus coro- Present study margin of well developed humeral shield; setae natus (Boddart) 1783 sh spiculiform, 13u in length, posterior to setae Brephosceles afribycis, new species h. Hysterosomal shield 233u in length, 101 u in width; interior margin strongly concave. Hystero­ The males of this new species are distin­ somal lobes 80u in length; setae ds positioned guished by an elongate genital organ which near apex of lobes, anterior to setae l3; setae d: extends beyond the insertions of setae Cs and a at terminus of lobe, approximate to setae pat. strongly concave anterior margin on the pre­ lnterlobar postlobar and extralobar Iamellae genital apodeme. The females are characterized well developed. All legs subequal. Ventral idio­ by the presence of lacunae and the absence of soma: Epimerites I fused in a Y; surface fields dorsolateral finger-like projections on the hys­ weakly developed. Pregenital apodeme fused; terosomal shield. trapezoidal in shape; anterior margin s_traight. MALE (holotype). Length, excluding post­ Genital organ 18u in length, not extending be­ lamellae, 415u; width, 171 u. Dorsal idiosoma: Propodosomal shield 106u in length, 125u in yond setae c2 ; subgenital shield present. Adanal shields well developed; not fused medially, not width; without lacunae; without external vertical joined to pregenital apodeme. setae; distance between external scapular setae, 114u. Scapular shields well developed. Setae 11 FEMALE (allotype). Length, 420u; width, 156u. spiculiform, Bu in length, positioned on anterior Dorsal idiosoma: Propodosomal shield 99u in margin of well developed humeral shield; setae length, 117u in width; without lacunae; without sh spiculiform, 16u in length, posterior to setae external vertical setae; distance between scap­ h. Hysterosomal shield 239u in length, 109u in ular setae, 109u. Setae l1 spiculiform, 7u in witdh; anterior margin strongly concave. Hys­ length, positioned on well developed humeral terosomal lobes 88u in length; setae d3 posi­ shield· setae sh spiculiform, 13u in length, pos­ tioned on anterior one-third of lobe, anterior to terior 'to setae h. Hysterosomal shield 244u in setae '3; setae d4 at terminus of lobe, approxi­ length, 91 u in width; anterior margin strongly mate to setae pai. lnterlobar, postlobar and concave; with lacunae; without dorsolateral extralobar lamellae well developed. All legs sub­ finger-like projections; setae d3 and d4 sepa­ equal. Ventral idiosoma: Epimerites I fused in a rated by 36u. Hysterosomal terminus disti~c!IY Y; surface fields weakly developed. Pregenital bilobate, without terminal lamellae. Ventral tdto­ apodeme fused; trapezoidal in shape, anterior soma: Surface fields weakly developed. Preanal margin strongly concave. Genital organ 29u in apodeme absent. length, extending beyond setae c2; subgenital Type material. From Stephanibyx melanop­ shield apparently absent. Adanal shields not terus (Charadiidae): holotype male (SAIMR), fused medially, not joined to pregenital apo­ allotype female (SAIMR), 4 male, 2 female deme. paratypes, September 20, 1953, Maputo, Mo~am­ FEMALE (allotype). Length, 451 u; width, 182u. bique, T. Santos Dias. Paratypes deposited: Dorsal idiosoma: Propodosomal shield 109u in Gaud, RNH, SAIMR, UGA, USNM. length, 127u in width; without lacunae; without Additional material. (Charadriidae): 21 males, external vertical setae; distance between scapu­ 7 females from Stephanibyx coronatus corona­ lar setae, 114u. Setae '1 spiculiform, 10u in tus, no other data. length, positioned on anterior margin of well Remarks. The drawings of the male and developed humeral shield; setae sh spiculiform, female are from the holotype and allotype re- 18u in length, posterior to setae h. Hysterosomal 154 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM shield 259u in length, 83u in width; anterior unable to recollect the female of this species margin strongly concave; with lacunae, without from the type host. finger-like dorsolateral projections; setae d3 and Brephosceles vane/Ii Gaud d4 separated by 65u. Hysterosomal terminus dis­ tinctly bilobate, without terminal lamellae. Ven­ Brephosceles vane/Ii Gaud, 1959, Bull. Soc. Sci. tral idiosoma: Surface fields weakly developed. Nat. Phys. Maroc, 37(2): 114-115. Type host: Preanal apodeme absent. Vanellus vanel/us (Charadriidae). Type material. From Afribyx lateralis (Chara­ Brephosceles vane/Ii, Gaud and Till, 1961. Publ. driidae): holotype male (USNM), allotype female S. Afr. Inst. Med. Res., 11 (L): 242. (USNM), 2 male, 5 female paratypes, April 23, Brephosceles vane/Ii, Peterson and Atyeo, 1968, 1955, Tete Pan, Portuguese East Africa, collec­ Bull. Univ. Nebraska St. Mus., 8(4): 217-236. tor unknown. Paratypes deposited: Gaud, UGA. Although the males of this species are indis­ Additional material. (Charadriidae): 1 male, 1 tinguishable from a number of related species in female from Afribyx senegal/us, North Rhodesia; the charadrii group, the females are easily rec­ 4 males, 9 females from Afribyx senegal/us, ognized by the spermatheca and associated North Cameroun and 2 males, 1 female, Sudan. ducts, and on the hysterosoma by the absence Remarks. The drawings of the male and fe­ of a well defined reticular pattern and the ab­ male are from the holotype and allotype respec­ sence of dorsolateral finger-like projections. The tively. The spermatheca is from a paratype. The primary spermathecal duct is extremely wide; species is named for the host. secondary ducts short, originating from primary spermathecal duct (Figs. 139 and 140). HOSTS MALE. Indistinguishable (for formal descrip­ Charadriidae tion see B. charadrii, page 145). Afribyx lateralis Present study FEMALE (paratype). Length, 428u; width 171u. (A. Smith) 1839 Dorsal idiosoma: Propodosomal shield 103u in Afribyx senegal/us Present study length, 127u in width; without lacunae; without (L.) 1766 external vertical setae; distance between exter­ nal scapular setae 117u. Setae si 16u in length, Brephosceles furcifer (Trouessart) separated by 91u. Setae '1 setiform, 13u in Proctophyllodes (Pterocolus) ortygometrae fur­ length, positioned on well developed humeral cifer Trouessart, 1885, Bull. Soc. Etud. Sci. shield; setae sh spiculiform, 18u in length, pos­ Angers, 14: 73-74. Ex: Rhinoptilus africanus terior to setae h. Hysterosomal shield 246u in (=Cursorius bicinctus) (Glareolidae). length, 104u in width; anterior margin strongly Pterodectes ortygometrae furcifer, Canestrini concave; distance between setae d 3 and d4 39u; and Kramer, 1899, Tierreich, 7: 122. with faint reticular pattern; without dorsolateral Brephosceles ortygometrae furcifer, Radford, finger-like projections. Primary spermathecal 1953, Parasitol., 42(3,4): 202. duct extremely wide, not tapering posteriorly; Brephosceles furcifer, Gaud and Till, 1961, Publ. secondary ducts Su in length, originating from So. Afr. Inst. Med. Res., 11(L): 241. primary spermathecal duct. Hysterosomal ter­ Brephosceles furcifer, Peterson and Atyeo, 1968, minus distinctly bilobate; without well developed Bull. Univ. Nebraska St. Mus., 8(43): 220. terminal lamellae. Ventral idiosoma: Surface Brephosceles furcifer is known from one male fields weakly developed. Preanal apodeme collected on Rhinoptilus africanus africanus barely visible. (=Cursorius bicinctus) from South Africa. This Type material. From Vanellus vanel/us (Chara­ specimen is indistinguishable from males of 8. driidae): 6 males, 6 females, Africa, J. Gaud, J. charadrii and related species (for a generalized Mouchet. Types are deposited in the personal description see B. charadrii, page 137). B. furci­ collection of Dr. J. Gaud. fer is retained in the charadrii group on the Material examined. (Charadriidae): 23 males, basis of the similarity of the male to the other 11 females from Hoplopterus spinosus, Uganda; included species. To date, the author has been 4 females, Egypt; 10 males, 2 females from A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 155

Hoplopterus armatus, Kenya; 7 males, 2 females cayennensis (Charadriidae}: holotype female from Hemiparra crassirostris leucoptera, Sudan; (USNM}, allotype male (USNM}, 5 male, 6 fe­ 14 males, 5 females from Xiphidiopterus atri­ male paratypes, May 20, 1942, Magdalena ceps, Nigeria. Department, Columbia, N. A. Carriker, Jr.; para­ Remarks. The drawings of the female hystero­ types: 6 males, 3 females, December 26, 1945, soma and spermatheca are from a paratype. Santa Marta, Columbia, M. A. Carriker. Para­ types deposited: AMNH, Gaud, RNH, UGA, HOSTS USNM. Charadriidae Remarks. The drawings of the hysterosomal Vanel/us vanellus Gaud, 1959 shield and the spermatheca are from the holo­ (L.} 1758 Gaud and Till, 1961 type and a paratype respectively. Present study Hoplopterus armatus Present study HOSTS (Burchell} 1822 Charadriidae Hemiparra crassirostris Present study Belonopterus chilensis cay­ Present study leucoptera ennesis (Gmelin} 1789 Reichenow, 1889 Xiphidiopterus atriceps Gaud and Till, 1961 Brephosceles tuberi, new species (Gould} 1834 Present study The spermatheca of Brephosceles tuberi and that of the related species, B. longistriatus, are Brephosceles belonopteri, new species indistinguishable; however, the two species can The distance between setae d 3 and d4 (42u}, be separated on the basis of the reticular pat­ the absence of dorsolateral finger-like projec­ tern on the hysterosomal shield. B. tuberi tions, and a tapering primary spermathecal duct exhibits a faint reticular pattern with 3-4 dorso­ set the female of this species apart from related lateral finger-like projections, whereas in B. species in the charadrii group. Longitudinal stri­ longistriatus the opisthosoma is adorned by dis­ ations on the hysterosomal shield are distinctive tinct longitudinal striations; the finger-like pro­ (Figs. 141 and 142}. jections are absent (Figs 143 and 144}. MALE (allotype}. Indistinguishable (for formal MALE (allotype}. Indistinguishable (for formal description see B. charadrii, page 29}. description see B. charadrii, page 145}. FEMALE (holotype}. Length, 417u; width, 187u. FEMALE (holotype}. Length, 412u; width, Dorsal idiosoma: Propodosomal shield 119u in 150u. Dorsal idiosoma: Propodosomal shield length, 122u in width; without lacunae; without 106u in length, 114u in width; without lacunae; external vertical setae; distance between exter­ without external vertical setae; distance be­ nal scapular setae 127u. Setae si 13u in length, tween external scapular setae 106u. Setae si separated by 99u. Setae '1 spiculiform, 16u in 13u in length, separated by 86u. Setae '1 lance­ length, positioned on well developed humeral olate, 107u in length, positioned on anterior shield; setae sh lanceolate, 21 u in length, pos­ margin of well developed humeral shield; setae terior to setae h. Hysterosomal shield 272u in sh spiculiform, 16u in length, posterior to setae length, 109u in width; anterior margin shallowly h. Hysterosomal shield 239u in length, 86u in

concave; distance between setae d3 and d4 42u; width; anterior margin shallowly concave; dis­ with longitudinal striations in reticular pattern; tance between setae ds and d4 42u; between d4 without dorsolateral finger-like projections. Pri­ and d5 57u; with faint reticular pattern; with 3-4 mary spermathecal duct tapering distally; dorsolateral finger-like projections. Primary secondary ducts thin with slight swelling at mid­ spermathecal duct with proximal sclerotized length, 10u in length. Hysterosomal terminus expansion; secondary duct simple, 13u in length. distinctly bilobate; without well developed ter­ Hysterosomal terminus distinctly bilobate; with­ minal lamellae. Ventral idiosoma: Surface fields out well-developed terminal lamellae. Ventral weakly developed. Preanal apodeme absent. idiosoma: Surface fields weakly developed. Pre­ Type material. From Belonopterus chilensis anal apodeme absent. 156 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

140

5139 141 200u 50u

Sp

HS

~ 146

145 Figs. 139-146. Brephosce/es vane/Ji Gaud, female (paratype): hysterosomal shield (139); spermatheca (140); Brephosceles belonopteri, new species, female (holotype): hysterosomal shield (141); spermatheca (142); Brephosceles tuberi, new species, female (holotype): hysterosomal shield (143); spermatheca (144); Brephosce/es /ongistriatus, new species, female (holotype): hysterosomal shield (145); spermatheca (146). A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 157

Type material. From Charadrius alexandrinus nal scapular setae 99u. Setae si 10u in length, seebohmi (Charadriidae): holotype female separated by 75u. Setae '1 lanceolate, 10u in (USNM), allotype male (USNM), 8 male, 4 fe­ length, positioned on well developed humeral male paratypes, April 16, 1914, Puttalam, North­ shield; setae sh spiculiform, 16u in length, pos­ west Province, Ceylon, S. D. Ripley; 1 male, 2 terior to setae h. Hysterosomal shield 236u in female paratypes, June 2, 1914, Trincomalee, length, 75u in width; anterior margin shallowly

Eastern Province, Ceylon, S. D. Ripley. Para­ concave; distance between setae d3 and d4 57u; types deposited: AMNH, Gaud, RNH, TC, UGA. reticular pattern with prominent longitudinal stri­ Additional material. (Charadriidae): 10 males, ations; without dorsolateral finger-like projec­ 4 females from Charadrius alexandrinus dealba­ tions. Primary spermathecal duct expanded tus, Malaya, 5 males, 2 females Thailand; 3 proximally; secondary ducts simple, 10u in males, 2 females from Charadrius pecuarius, length. Hysterosomal terminus distinctly bilo­ Gold Coast, Africa; 19 males, 9 females from bate; without well developed terminal lamellae. Charadrius peronii, Philippine Islands; 21 males, Ventral idiosoma: Surface fields weakly devel­ 9 females from Charadrius thoracicus, Malagasy oped. Preanal apodeme absent. Republic. Type material. From Charadrius vociferus Remarks. The drawing of the female hystero­ ternominatus (Charadriidae): holotype female soma is from the holotype. The illustration of (USNM) allotype male (USNM), 15 male, 9 the spermatheca is taken from one of the female female paratypes, April 29, 1933, Guanica La­ paratypes. The species is named for the prox­ goon, Puerto Rico, S. T. Danforth; paratypes: imal expansion of the primary spermathecal 1 male, 2 females, July 8, 1912, Manati, Puerto duct. Rico, A. Wetmore; 5 males, 3 females, April 9, 1930, Petit-Trou-de-Nippes, Haiti, W. M. Perrygo; HOSTS 2 females, November 18, 1961, Baffin Bay, Kle­ Charad ri idae berg County, Texas, B. McDaniel. Paratypes Charadrius alexandrinus Present study deposited: AMNH, Gaud, UGA, ZSBS. seebohmi Additional material. (Charadriidae): 4 males, Hartert and Jackson, 1915 3 females from Charadrius melodus melodus, C. alexandrinus dealbatus Present study Louisiana; 2 males, 1 female from Charadrius Swinhoe, 1870 melodus circumcinctus, Mississippi. C. pecuarius Present study Remarks. The drawings of the hysterosoma Temminck, 1823 are from the holotype female. The illustration C. peronii Present study of the spermatheca is from a paratype. The Schlegel, 1856 species is named for the longitudinal striations C. thoracicus Present study on the hysterosomal shield of the female. Richmond, 1896 HOSTS Brephosceles longistriatus, new species Charadriidae This new species is distinguished by a prox­ Charadrius vociferus Present study imally expanded primary spermathecal duct, a ternominatus hysterosoma with longitudinal striations, and by Banks and Kennard, 1766 the absence of dorsolateral finger-like projec­ C. melodus melodus Present study tions. To date, this is the only species which is Ord, 1824 so adorned (Figs. 145 and 146). C. m. circumcinctus Present study MALE (allotype). Indistinguishable (for formal Ord,1824 description see 8. charadrii, page 145). FEMALE (holotype). Length, 404u; width, Brephosceles constrictus, new species 145u. Dorsal idiosoma: Propodosomal shield 96u The new species Brephosceles constrictus in length, 93u in width; without lacunae; without can easily be recognized by the lateral margins external vertical setae; distance between exter- of the hysterosomal shield and the secondary 158 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM ducts of the spermatheca. At approximately the terosoma has three finger-like projections and level of setae d3 the lateral margins of the hys­ the proximal one-third of the primary duct is terosomal shield are strongly incised; secondary expanded (Figs. 147-154). ducts have a fringe-like margin at their apex MALE (allotype). Indistinguishable (for formal (Figs. 147 and 148). description see B. charadrii, page 145). MALE (allotype). Indistinguishable (for formal FEMALE (holotype). Length, 407u; width, description see B. charadrii, page 145). 148u. Dorsal idiosoma: Propodosomal shield FEMALE (holotype). Length, 459u; width, 104u in length, 117u in width; without lacunae; 156u. Dorsal idiosoma: Propodosomal shield without external vertical setae; distance be­ 101 u in length, 119u in width; without lacunae; tween external scapular setae 55u. Setae si 10u without external vertical setae; distance be­ in length, separated by 83u. Setae 11 lanceolate, tween external scapular setae 108u. Setae si 10u in length, positioned on well developed 18u in length, separated by 88u. Setae '1 lance­ humeral shield; setae sh spiculiform, 16u in olate, 13u in length, positioned on well devel­ length, posterior to setae h. Hysterosomal shield oped humeral shield; setae sh spiculiform, 16u 241 u in length, 86u in width; anterior margin in length, posterior to setae h. Hysterosomal shallowly concave; distance between setae ds shield 272u in length, 78u in width; anterior and d 4 , 55u; without reticular pattern; with three margin strongly concave; lateral margin deeply dorsolateral finger-like projections. Proximal incised at level of setae d3; distance between one-third of primary spermathecal duct ex­ setae d8 and d4, 67u; without reticular pattern; panded, heavily sclerotized; secondary ducts without dorsolateral finger-like projections. Pri­ 18u in length. Hysterosomal terminus distinctly mary spermathecal duct not expanded proxim­ bilobate, without well developed terminal lam­ ally; secondary ducts with fringe-like margin on ellae. Ventral idiosoma: Surface fields weakly apex, 10u in length. Hysterosomal terminus dis­ developed. Preanal apodeme absent. tinctly bilobate; without well developed termi­ Type material. From Hoplopterus duvancelli nal lamellae. Ventral idiosoma: Surface fields (Charadriidae): holotype female (USNM), allo­ weakly developed. Preanal apodeme absent. type male (USNM), 12 male, 8 female para­ Type material. From Charadrius bincinctus types, October 28, 1896, Trong, Trong Province, (Charadriidae): holotype female (USNM), allo­ Thailand, W. L. Abott. Paratypes deposited: type male (USNM), 15 male, 10 female para­ BMNH, Gaud, RNH, SAIMR, UGA, USNM. types, July 5, 1884, Manukau Harbor, New Remarks. The drawings of the hysterosoma Zealand, collector unknown. Paratypes depos­ and spermatheca are from the holotype female. ited: BMNH, Gaud, SAIMR, USNM, ZSZM. The species is named for the host. Remarks. The drawings of the hysterosoma and spermatheca are from the female holotype. HOSTS The species is named for the deeply incised lat­ Charadriidae eral margins of the hysterosomal shield. Hoplopterus duvancelii Present study (Lesson) 1826 HOSTS (=H. ventralis) Charadriidae Charadrius bicinctus Present study Brephosceles inornatus, new species Jardine and Selby, 1827 This new species is distinguished by the ab­ sence of projections or a reticular pattern on Brephosceles hoplopteri, new species the hysterosomal shield, the tapering spermath­ The distance between setae ds and d4, the ecal duct, and the distance between setae d3

dorsolateral finger-like projections on the hys­ and d4 (39u) (Figs. 151 and 152). terosoma and the shape of the primary sperm­ MALE (allotype). Indistinguishable (for formal athecal duct sufficiently distinguish this species description see B. charadrii, page 145). from related species in the charadrii group. FEMALE (holotype). Length, 407u; width, Setae d 3 and d 4 are separated by 55u; the hys- 137u. Dorsal idiosoma: Propodosomal shield A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 159

148 150

149 200u 50u 147

Sp

HS

152

Figs. 147-154. Brephosce/es constrictus, new species, female (holotype): hysterosomal shield (147); spermatheca (148); Brephosce/es hop/opteri, new species, female (holotype): hysterosomal shield (149); spermatheca (150); Brephosceles inornatus, new species, female (holotype): hysterosomal shield (151); spermatheca (152); Brephosce/es histionotus Gaud and Mouchet, female (paratype): hysterosomal shield (153); spermatheca (154). 160 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

101 u in length, 96u in width; without lacunae; Gaud and Mouchet, 1957, distinguish the without external vertical setae; distance be­ males of Brephosceles histionotus from B. cha­ tween external scapular setae 101 u. Setae si radrii on the basis of total length, the distance 13u in length, separated by 83u. Setae 11 lance­ between the origin of the genital organ and the olate, Bu in length, positioned on well developed copulatory discs and the distance between the humeral shield; setae sh spiculiform, 13u in anal setae. Recent observations made on hun­ length, posterior to setae h. Hysterosomal shield dreds of males of both species reveal that none 241u in length, 78u in width; anterior margin of these characters or combinations of charac­ strongly concave; distance between setae d3 ters will adequately separate the two males. and d4 39u; without prominent reticular pattern; However, B. histionotus can be recognized in without dorsolateral finger-like projections. Pri­ the female by the reticular pattern on the hys­ mary spermathecal duct tapering distally; terosomal shield and by the position of the secondary ducts simple, 13u in length. Hystero­ secondary ducts of the spermatheca. These sec­ somal terminus distinctly bilobate; without well ondary ducts originate from the primary sperm­ developed terminal lamellae. Ventral idiosoma: athecal duct; a character known to be unique Surface fields weakly developed. Preanal apo­ to the charadrii group (Figs. 153 and 154). deme absent. MALE. Indistinguishable. Type material. From Charadrius leschenau/tii FEMALE (paratype). Length, 415u; width, (Charadriidae): holotype female (USNM), allo­ 153u. Dorsal idiosoma: Propodosomal shield 99u type male (USNM), 9 male, 6 female paratypes, in length, 101 u in width; without lacunae; with­ March 24, 1896, Prahmon, Trong Province, Thai­ out external vertical setae; distance between land, W. L. Abbott. Paratypes deposited: AMNH, external scapular setae 106u. Setae si 16u in BMNH,Gaud, RNH, UGA, USNM. length, separated by 83u. Setae Ii lanceolate, Additional material. (Charadriidae): 11 males, 10u in length, positioned on well developed 6 females from Charadrius mongolus atrifons, humeral shield; setae sh spiculiform, 16u in Sumatra; 14 males, 10 females, Thailand. length, posterior to setae h. Hysterosomal Remarks. The drawings of the hysterosomal shield 244u in length, 86u in width; anterior shield and spermatheca are from the holotype margin shallowly concave; distance between and allotype respectively. The species is named setae da and d4 52u; with prominent reticular for the absence of a reticular pattern on the pattern; without dorsolateral finger-like projec­ hysterosomal shield. tions. Primary spermathecal duct not expanded proximally; secondary ducts originating from primary spermathecal duct; 13u in length. Hys­ HOSTS terosomal terminus bilobate; without well de­ Charadriidae veloped terminal lamellae. Ventral idiosoma: Charadrius leschenaultii Present study Surface fields weakly developed. Preanal apo­ Lesson, 1826 deme absent. Charadrius mongolus atrifons Present study Type material. From Charadrius tricollaris Wagler, 1829 forbesi (Charadriidae): 20 males, 19 females, French Cameroons, J. Gaud and J. Mouchet. Brephosceles histionotus Gaud and Mouchet Primary and secondary types are in the personal Brephosceles histionotus, Gaud and Mouchet, collection of Dr. J. Gaud. 1957, Ann. Parasitol. hum. comp., 32(5-6): 499- Material examined. (Charadriidae): 23 males, 500, Figs. 2(A) and 3(C,D). Type host: Charad­ 12 females from Charadrius tricollaris tricollaris, rius tricol/aris forbesi (Charadriidae). Kenya; 8 males, 3 females from Charadrius tri­ Brephosceles histionotus, Gaud, J. and W. Till. collaris bifrontatus, Madagascar. 1961. Publ. S. Afr. Inst. Med. Res., 11(L): 242. Remarks. The drawings of the hysterosomal Brephosceles histionotus, Peterson and Atyeo, shield are from a paratype. The illustration of 1968, Bull. Univ. Nebraska St. Mus., 8(4): 220. the spermatheca is from identified material. A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 161

HOSTS 6 females from Belonopterus chilensis chilensis, Cha rad ri idae Chile. Charadrius tricollaris forbesi Gaud, 1957 Remarks. The drawings of the hysterosomal (Shelley) 1883 Present study shield and spermatheca are from the holotype Charadrius tricollaris Present study female. The species is named for the host. bifrontatus Cabanis, 1882 Charadrius tricol/aris Present study HOSTS tricollaris Vieillot, 1818 Charadriidae Belonopterus chilensis Present study Brephosceles chilensis, new species lampronotus (Wagner) 1827 Large overall dimensions and the absence of Belonopterus chilensis Present study dorsolateral finger-like projections on the hys­ chilensis (Molina) 1782 terosomal shield of the femals set this species apart from related species in the charadrii Brephosceles virginiensis, new species group. The primary spermathecal duct is not Brephosceles virginiensis, new species, can expanded; secondary ducts are 10u in length be differentiated from the related species by: (Figs. 155 and 156). (1) distance between setae d 3 and d4, 39u, (2) MALE (allotype). Indistinguishable (for formal three-four dorsolateral finger-like projections on description see B. charadrii, page 145). the hysterosoma, (3) primary spermathecal duct FEMALE (holotype). Length, 456u. width, tapering gradually to supranal concavity (Figs. 161 u. Dorsal idiosoma: Propodosomal shield 157 and 158). 112u in length, 125u in width; without lacunae; MALE (allotype). Indistinguishable (for formal without external vertical setae; distance be­ description see B. charadrii, page 145). tween external scapular setae 117u. Setae si FEMALE (holotype). Length, 415u; width, 150u. 21u in length, separated by 91u. Setae l1 spiculi­ Dorsal idiosoma: Propodosomal shield 109u form, 10u in length, positioned on well devel­ in length, 99u in width; without lacunae; without oped humeral shield; setae sh lanceolate, 18u external vertical setae; distance between exter­ in length, posterior to setae h. Hysterosomal nal scapular setae 104u. Setae si 18u in length, shield 267u in length, 100u in width; anterior separated by 83u. Setae '1 lanceolate, 13u in margin shallowly concave; distance between length, positioned on well developed humeral setae d3 and d4 52u; with faint reticular pattern; shield; setae sh spiculiform, 18u in length, pos­ without dorsolateral finger-like projections. Pri­ terior to setae h. Hysterosomal shield 246u in mary spermathecal duct without proximal scler­ length, 78u in width; anterior margin shallowly otized expansion, tapering slightly; secondary concave; distance between setae ds and d4, 39u; ducts 10u in length. Hysterosomal terminus dis­ between setae d4 and d 5, 57u; without reticular tinctly bilobate; without well developed terminal pattern; with 3-4 dorsolateral finger-like projec­ lamellae. Ventral idiosoma: Surface fields tions. Primary spermathecal duct not expanded weakly developed. Preanal apodeme absent. proximally, tapering gradually to supranal con­ Type material. From Belonopterus chilensis cavity; secondary ducts simple, 10u in length. lampronotus (Charadriidae): holotype female Hysterosomal terminus distinctly bilobate; with­ (USNM), allotype male (USNM), 9 male, 7 fe­ out well developed terminal lamellae. Ventral male paratypes, December 15, 1920, Carhue, idiosoma: Surface fields weakly developed. Pre­ Buenos Aires, Argentina, A. Wetmore; para­ anal apodeme absent. types: 6 males, 4 females, September 14, 1898, Type material. From Charadrius wilsonia wi/­ Buenos Aires, Argentina, S. Venturi; 21 males, sonia (Charadriidae): holotype female (NU), 11 females, January, 1897, Buenos Aires, Argen­ allotype male (NU), 18 male, 5 female para­ tina, P. Neuman. Paratypes deposited: AMNH, types, April 19, 1947, Gargatha Inlet, Accomack BMNH, Gaud, SAIMR, SEA, UGA, USNM, ZSBS. County, Virginia, collector unknown. Paratypes Additional material. (Charadriidae): 10 males, deposited: BMNH, Gaud, SIU, UGA. 162 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

Additional material. (Charadriidae): 10 males, Type material. From Lobivanellus indicus 4 females from Charadrius wilsonia cinna­ atronuchalis (Charadriidae): holotype female mominus, Columbia, South America; 7 males, (AMNH), allotype male (AMNH), 10 male, 2 2 females from Charadrius wilsonia beldingi, female paratypes, February 17, 1910, Trong, Mexico; 14 males, 7 females from Charadrius Trong Province, Thailand, collector unknown; wi/sonia rutinucha, Haiti. paratypes: 4 males, 2 females, Assam, India, Remarks. The drawings of the hysterosoma G. G. Crozier; 8 males, 3 females, March 28, are from the female holotypes. The illustration 1936, Doug Me, near Phan Raug, Vietnam, col­ of the spermatheca is from a paratype. The lector unknown. Paratypes deposited: AMNH, species is named for the type locality. BMNH, LAS, SAIMR, UGA, USNM. Remarks. The drawing of the hysterosomal HOSTS shield is from the female holotype. The illustra­ Charadriidae tion of the spermatheca is from a paratype. The Charadrius wilsonia wilsonia Present study species is named for the host. Ord 1814 C. w. beldingi Present study HOSTS Ridgway, 1919 C. w. cinnamominus Present study Charadriidae Ridgway, 1919 Lobivanellus indicus atronu­ Present study C. w. rutinucha Present study chalis Jerdon, 1864 Ridgway, 1874 Brephosce/es collaracus, new species Brephosceles lobivanelli, new species The four dorsolateral finger-like projections The proximal swellings of the secondary ducts on the female hysterosomal shield, the short of the spermatheca, the presence of three dor­ distance between setae d3 and d4, and the rela­ solateral projections, and the distance between tively simple morphology of the spermatheca and associated ducts distinguish this species setae d4 and d5 (44u) sufficiently characterize this species (Figs. 159 and 160). from related species in the charadrii group (Figs. MALE (allotype). Indistinguishable (for formal 161 and 162). description see B. charadrii, page 145). MALE (allotype). Indistinguishable (for formal FEMALE (holotype). Length, 422u; width, description see B. charadrii, page 145). 166u. Dorsal idiosoma: Propodosomal shield 99u FEMALE (holotype). Length, 409u; width, in length, 106u in width; without lacunae; with­ 143u. Dorsal idiosoma: Propodosomal shield out external vertical setae; distance between 104u in length, 127u in width; without lacunae; external scapular setae 112u. Setae si 10u in without external vertical setae; distance be­ length, separated by 88u. Setae 11 spiculiform, tween external scapular setae 106u. Setae si 10u in length, positioned on well developed 13u in length, separated by 86u. Setae '1 lance­ humeral shield; setae sh spiculiform, 16u in olate, 10u in length, positioned on well devel­ length, posterior to setae h. Hysterosomal shield oped humeral shield; setae sh spiculiform, 16u 236u in length, 96u in width; anterior margin in length, posterior to setae h. Hysterosomal strongly concave; distance between setae ds shield 233u in length, 83u in width; anterior and d4, 42u; between setae d4 and d 5, 44u; margin strongly concave; distance between without reticular pattern; with 2-3 distinct dorso­ setae ds and d4, 36u; with faint reticular pattern; lateral finger-like projections. Primary spermath­ with four dorsolateral finger-like projections. ecal duct not expanded proximally; secondary Primary spermathecal duct not expanded prox­ ducts with proximal swellings, 16u in length. imally, not tapering; secondary ducts simple 10u Hysterosomal terminus distinctly bilobate; with­ in length. Hysterosomal terminus distinctly bilo­ out well developed terminal lamellae. Ventral bate; without well developed terminal lamellae. idiosoma: Surface fields weakly developed. Pre­ Ventral idiosoma: Surface fields weakly devel­ anal apodeme absent. oped. Preanal apodeme absent. A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 163

156 158 157 200u 50u

Sp

HS

160

Figs. 155-162. Brephosce/es chi/ensis, new species, female (holotype): hysterosomal shield (155); spermatheca (156); Brephosce/es virginiensis, new species, female (holotype): hysterosomal shield (157); spermatheca (158); Brephosce/es lobivanel/i, new species, female (holotype): hysterosomal shield (159); spermatheca (160); Brephosceles collaracus, new species, female (holotype): hysterosomal shield (161); spermatheca (162). 164 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

Type material. From Charadrius co/laris (Cha­ Pachyptila desolata (Gmelin) 1789 radriidae): holotype female (USNM), allotype Brephosceles prolatus, new species male (USNM), 10 male, 8 female paratypes, Brephosceles pachyptilae, new species March 26, 1957, Tonosi, Los Santos Province, Pterodroma cookii defilippiana (Giglioli Panama, A. Wetmore. Paratypes deposited: and Salvadori) 1869 AMNH, BMNH, RNH, UGA, USNM. Brephosce/es disjunctus, new species Remarks. The drawings of the hysterosomal Brephosceles triangularis, new species shield and the spermatheca are from the holo­ Pterodroma externa cervicalis (Salvin) 1891 type female. The species is named for the host. Brephosce/es disjunctus, new species Brephosce/es lunatus, new species HOSTS Pterodroma externa externa (Salvin) 1876 Charadriidae Brephosce/es /unatus, new species Charadrius co/laris Present study Pterodroma /eucoptera hypo/euca (Salvin) 1888 Vieillot, 1818 Brephosce/es disjunctus, new species Brephosceles pterodromae, new species Pterodroma /eucoptera masafuerae Lonnberg, HOST-PARASITE LIST 1921 Brephosce/es disjunctus, new species ORDER: GAVII FORM ES Puffinus bu/leri Salvin, 1888 Brephosce/es se/enopeltatus, new species Gaviidae Puffinus gravis (O'Reilly) 1818 Gavia arctica (L) 1758 Brephosce/es parvatus, new species Brephosceles forficiger (Megnin and Brephosce/es puffini, new species Trouesart) Puffinus griseus (Gmelin) 1789 Gavia immer (Brunnich) 1764 (=Colymbus Brephosce/es puffini, new species glacialis) Puffinus pacificus chlorohynchus Lesson, 1831 Brephosceles forficiger (Megnin and Brephosce/es parvatus, new species Trouessart) Brephosce/es se/enopeltatus, new species Puffinus pacificus cuneatus Salvin, 1888 Brephosce/es se/enopeltatus, new species ORDER: PROCELLARIIFORMES Puttinus puttinus (Brunnich) 1764 Brephosce/es bilobatus, new species Brephosce/es parvatus, new species Hydrobatidae Putfinus tenuirostris (Temminck) 1835 Hydrobates pelagicus (L.) 1758 Brephosceles parvatus, new species Brephosceles pelagicus (Vitzthum) Brephosceles superbus Dubinin Oceanodroma castro (Harcourt) 1851 Brephosce/es marlae, new species Brephosceles decapus (Gaud) Oceanodroma castro bangsi Nichols, 1914 Brephosce/es /anceolatus, new species Diomedeidae Oceanodroma castro /eucorhoa (Vieillot) Brephosceles decapus (Gaud) Diomedes chlororhynchos Gmelin, 1789 Oceanodroma markhami tristrami Salvin, 1896 Brephosce/es marginiventris (Trouessart) Brephosce/es longipenis, new species Diomedea chrysostoma Forster, 1785 Brephosce/es diomedei, new species Procellariidae Diomedea exulans L., 1758 Brephosceles diomedei, new species Macronectes giganteus (Gmelin) 1789 Diomedea me/anophris Temminck, 1828 Brephosceles marginiventris (Trouessart) Brephosce/es diomedei, new species A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 165

ORDER: ANSERIFORMES Charadrius alexandrinus dealbatus (Swinhoe) 1870 Brephosceles tuberi, new species Anatidae Charadrius a/exandrinus seebohmi Hartert Anas platyrhynchos L., 1758 and Jackson, 1915 Brephosceles anatina Dubinin Brephosceles tuberi, new species Cygnus bewickii Yarrel, 1830 Charadrius bicinctus Jardine and Selby, 1827 Brephosceles discidicus, new species Brephosceles constrictus, new species Nettapus auritus (Boddaert) 1783 Charadrius collaris Vieillot, 1818 Brephosce/es lambda (Trouessart) Brephosceles collaracus, new species Charadrius dubius curonicus Gmelin, 1789 Anhimidae Brephosceles charadrii Dubinin Anhima cornuta (L.) 1766 Charadrius dubius dubius Scopoli, 1786 Brephosce/es anhimae, new species Brephosceles charadrii Dubinin Charadrius dubius jerdoni (Legge) 1880 Brephosce/es charadrii Dubinin ORDER: GRUIFORMES Charadrius leschenaulti Lesson, 1826 Brephosceles inornatus, new species Gruidae Charadrius melodus circumcinctus Ord, 1824 Brephosceles /ongistriatus, new species Balearica regu/orum (Bennet) Charadrius me/odus melodus Ord, 1824 Brephosce/es geranoxenus Peterson Brephosceles longistriatus, new species Charadrius mongolus atrifrons Wagler, 1829 Brephosceles inornatus, new species ORDER: CHARADRIIFORMES Charadrius pecuarius Temminck, 1823 Brephosceles tuberi, new species Burhinidae Charadrius peronii Schlegel, 1856 Brephosceles tuberi, new species Burhinus vermicu/atus (Cabanis) 1868 Charadrius placidus Gray and Gray, 1863 Brephosceles chaubaudi Gaud Brephosce/es charadrii Dubinin Esacus recurvirostris (Cuvier) 1829 Charadrius thoracicus (Richmond) 1896 Brephosceles chaubaudi Gaud Brephosceles tuberi, new species Orthorhamphus magnirostris (Vieillot) 1818 Charadrius tricollaris bifrontatus Cabanis, 1882 Brephosce/es chaubaudi Gaud Brephosceles histionotus Gaud and Mouchet Charadrius tricol/aris forbesi (Shelley) 1883 Charadriidae Brephosceles histionotus Gaud and Mouchet Afribyx lateralis (A. Smith) 1839 Charadrius tricol/aris tricollaris Viei llot, 1818 Brephosceles afribycis, new species Brephosceles histionotus Gaud and Mouchet Afribyx senegallus (L.) 1766 Charadrius vociferus ternominatus Banks and Brephosceles afribycis, new species Kennard, 1766 Belonopterus chilensis cayennensis (Gmelin) Brephosceles longistriatus, new species 1789 Charadrius wilsonia be/dingi (Ridgway) 1919 Brephosce/es belonopteri, new species Brephosceles virginiensis, new species Be!onopterus chilensis chi/ensis (Molina) 1782 Charadrius wilsonia cinnamominus (Ridgway) Brephosceles chilensis, new species 1919 Belonopterus chi/ensis /ampronotus (Wagler) Brephosceles virginiensis, new species 1827 Charadrius wilsonia rutinucha (Ridgway) 1874 Brephosce/es chilensis, new species Brephosceles virginiensis, new species 166 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

Charadrius wi/sonia wi/sonia Ord, 1814 Stephanibyx coronatus coronatus (Boddaert) Brephosceles virginiensis, new species 1783 Hemiparra crassirostris /eucoptera Reichenow, Brephosce/es selenopeltata, new species 1889 Stephanibyx melanopterus (Cretzschmar) 1829 Brephosceles vane/Ii Gaud Brephosce/es se/enopeltata, new species Hoplopterus armatus (Burchell) 1822 Vanellus vanel/us (L.) 1758 Brephosceles vane/Ii Gaud Brephosceles vane/Ii Gaud Hoplopterus duvaucelii (Lesson) 1826 Xiphidiopterus atriceps (Gould) 1834 (=H. ventralis) Brephosce/es vane/Ii Gaud Brephosce/es hoplopterii, new species Lobivanellus indicus atronuchalis Jerdon, 1864 Haematopodidae Brephosce/es /obivanelli, new species Haematopus ostralegus /ongirostris Vieillot, 1817 Rhinoptilus africanus africanus (Temminck) 1807 Brephosce/es haematopi, new species (=Cursorius bicinctus) Haematopus palliatus palliatus Temminick, 1820 Brephosce/es furcifer (Trouessart) Brephosce/es turgidus, new species A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 167

ACKNOWLEDGMENTS I wish to express appreciation to Dr. Warren T. Atyeo for his willingness to help collect material and to arrange for the loans of many type speci­ mens. To those individuals who graciously supplied material for this study, I am also indebted. Type specimens were provided by Dr. Marc Andre (deceased); Dr. Jean Gaud, Laboratoire de Parasitologie, Rennes, France; Dr. Helen Dubininia, Leningrad Academy of Science, Leningrad, Russia; and Dr. Max Vachon, 61 rue de Buffon, 75-Paris (5), France. Numerous specimens were provided by Dr. Thomas Aitkin, Trinidad Regional Virus Laboratory; Mr. Binyon Amerson, U.S. National Museum, Washington, Dis­ trict of Columbia 20560; Dr. John C. Downey, Department of Zoology, Southern Illinois University, Carbondale, Illinois 62901; Dr. Douglas J. Gould and Dr. Elliot McClure, Migratory Animal Pathological Survey, A.P.O. San Francisco, California 96346; Dr. K. E. Hyland, Jr., Department of Zool­ ogy, University fo Rhode Island, Kingston, Rhode Island 02881; Dr. J. Linsley Gressitt, Bernice P. Bishop Museum, Honolulu, Hawaii 96819; and Dr. F. K. E. Zumpt, South African Institute for Medical Research, Johannesburg, Republic of South Africa. For making facilities available and arranging for the examination of ornithological collections, recognition is given to the following institutions: Academy of Natural Sciences of Philadelphia, American Museum of Natural History, British Museum (Natural History), Chicago Museum of Natural His­ tory, Smithsonian Institution, University of Michigan, and the University of Nebraska State Museum. A special acknowledgment is extended to Dr. Richard Zusi of the Divi­ sion of Birds, Smithsonian Institution, for assistance in avian systematics. The major part of this study was financed under the auspices of the National Science Foundation, grant numbers GB-15105 and GB-1620. The author wishes to thank this fine organization for support of this study. 168 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM

REFERENCES FOR AVIAN SYSTEMATICS Mayr, Ernst. 1941. List of New Guinea birds. American Museum of Natural History, New York: 1-260. American Ornithologists' Union. 1961. Check-list of North ---. 1945. Birds of the Southwest Pacific. Macmillan American birds (5th ed.). Port City Press, Inc., Baltimore: Co., New York: 1-316. 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. Jour. 17: New York: 1-316. 75-121. Bannerman, David A. 1953. The birds of west and equato­ McGregor, Richard C. 1909. A manual of Philippine birds. rial Africa. Oliver and Boyd, London: 1-1526. Bureau of Printing, Manila: 1-769. Barrett, Charles, 1947. Australian bird life. Oxford Univ. Munro, George C. 1960. Birds of Hawaii. Bridgeway Press: Press, Melbourne: 1-239. 1-189. Blake, Emmet Reid. 1953. Birds of Mexico. Univ. Chicago Peters, James Lee. 1931-1960. Check-list of birds of the Press, Chicago: 1-644. world. Vols. 1-7, 9, 15. Harvard Univ. Press, Cambridge. Peterson, Roger Tory. 1941. A field guide to western birds. Bond, James. 1956. Check-list of birds of the West Indies. Houghton Mifflin Co., Boston: 1-240. Acad. Nat. Sci. Philadelphia: 1-214. ---. 1954. A field guide to the birds of Britain and Cave, Francis 0., and James D. Macdonald. 1955. Birds of Europe. Collins, London: 1-318. the Sudan. Oliver and Boyd, London: 1-444. ---. 1960. A field guide to the birds of Texas. Hough­ Cory, C. B., B. Conover, and C. E. Hellmayr. 1918-1949. ton Mifflin Co., Boston: 1-304. Catalogue of birds of the Americas. Field Mus. Nat. Hist. Roberts, Austin. 1961. Birds of South Africa (rev. ed.). Zool. Ser. 13 (Part 1, 1942-1949 by C. E. Hellmayr and Cape Times Ltd., Cape Town: 1-504. B. Conover. Part 2, 1918 by C. B. Cory. Parts 3-5, 1924- Smythies, Bertram E. 1953. The birds of Burma (2nd ed.). 1927 by C. B. Cory and C. E. Hellmayr. Parts 6-11, Oliver and Boyd, London: 1-668. 1929-1938 by C. E. Hellmayr). ---. 1960. The birds of Borneo. Oliver and Boyd. Lon­ Delacour, Jean. 1947. Birds of Malaysia. Macmillan Co., don: 1-562. New York: 1-382. Storer, Robert W. 1956. The fossil loon, Colymboides Glenister, A. G. 1955. The birds of the Malay Peninsula, minutus. Condor 58: 413-426. Singapore and Penang. Oxford Univ. Press, London: Van Tyne, Josselyn and Andrew J. Berger. 1961. Funda­ 1-282. mentals of ornithology. John Wiley & Sons, Inc., New Mackworth-Praed, C. W. and C. H. B. Grant. 1952, 1955. York: 1-624. Birds of eastern and northeastern Africa. Longmans, Yudin, K. A. 1965. The phylogeny and classification of the Green and Co., London 1: 1-836; 2: 1-1099. Charadriiformes. Fauna U.S.S.R. 2 (1): 1-257. A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 169

BIBLIOGRAPHY ---. 1953. Sarcoptides plumicoles des oiseaux d'Af­ rique occidentale et centrale. Ann. Parasitol. hum. comp. Atyeo, Warren T., and Norman L. Braasch, 1966. The 28(3): 193-226. feather mite genus Proctophyllodes (Sarcoptiformes: ---. 1957. Acariens plumicoles (Analgesoidea) para­ Proctophyllodidae). Bull. Univ. Nebraska State Mus. 5: sites des oiseaux du Maroc. Bull. Soc. Sci. nat Phys. 1-354. Maroc 37(2): 105-136. Atyeo, Warren T., and J. Gaud. 1966. The chaetotaxy of ---. 1960. Quelques Sarcoptiformes plumicoles du sarcoptiform feather mites (Acarina: Analgoidea). Jour. Congo beige (Analgesoidea). Rev. Zool. Bot. Africa 61 Kansas Entomol. Soc. 39(2): 337-346. (1-2): 131-159. Atyeo, Warren T., and Paul C. Peterson. 1966. The genus ---. 1961. Six genres nouveaux de Sarcoptiformes Dinalloptes (Analgoidea: Proctophyllodidae). Acarologia plumicoles (Analgesoidea). Acarolgia 3(1): 78-95. 8(3): 470-474. ---. 1964. Acariens plumicoles (Analgesoidea). Ann. ---, and ---. 1967a. Antarctic feather mites (Aca­ Mus. Roy. Afrique Cent., Zool. 80(132): 119-130. rina: Sarcoptiformes). Antarctic Res. Ser., Antarctic Ent. Gaud, J., and J. Mouchet. 1957. Acariens plumicoles (Anal­ 10: 97-103. gesoidea) des oiseaux du Cameroun. I. Proctophyllodi­ ---, and ---. 1967b. The feather mite genus Lam­ dae. Ann. Parasitol. hum. comp. 32(5-6): 491-546. inal/optes Dubinin, 1955 (Proctophyllodidae: Alloptinae). Gaud, J., and W. Till. 1961. In Zumpt, F. (ed.). The arthro­ Jour. Kansas Ent. Soc. 40(4): 447-458. pod parasites of vertebrates in Africa south of the Banks, Nathan. 1904. A treatise on the Acarina or mites. Sahara (Ethiopian Region). I. Chelicerata. Publ. S. Afr. Proc. U.S. Nat. Mus. 28: 87-94. Inst. Med. Res. 11(L): 186-301. Bedford, G. A. H. 1932. A synoptic check-list and host-list Hull, J. 1934. Concerning British Analgidae (Feather-Mites). of ectoparasites found on South African Mammalia, Trans. N. Nat. Union 1(3): 200-206. Aves, and Reptilia (2nd ed.). 18th Rept. Dir. Vet. Serv. Megnin, P., and E. L. Trouessart. 1884. Note sur la morph­ Anim. lndust.: 240-267. ologie et sur la classification des Sarcoptides plumi­ ---. 1936. A synoptic check-list and host-list of the coles. Bull. Soc. Etud. Sci. Angers 12: 131-139. ectoparasites found on South African Mammalia, Aves, ---. 1884b. Les Sarcoptides plumicoles. Journ. Micro­ and Reptilia (Suppl. No. 1). Jour. Vet. Sci. Anim. lndust. graph. 8(7): 380-385. 7(1): 70-76. Orwig, Kenneth R. 1968. The genera and species of the Baria, H. F. 1959. Analgesoidea Neotropicais. IV. Sobre feather mite subfamily Trouessartinae (except Trous­ algumas especies novas ou Pouco conhecidas de aca­ sartia) (Acarina: Proctophyllodidae). Bull. Univ. Nebraska rinos plumicolas. Bol. Mus. Nae. Zool. (209): 1-17. State Mus. 8(1): 1-182. Berlese, Antonio. 1885-1897. Acari, Myriopoda et Scorpi­ Oudemans, A. 1910. Notes on Acari. XXth Series. (Acari­ ones. Padova fasc. 27(1); fasc. 77(1). dae.) Deutsch. Ent. Zeitschr. 6: 389-407. Buchholz, R. 1869. Memerkugen uber die Arten der Gat­ Peterson, Paul C., and Warren T. Atyeo. 1968. New genera tung Dermaleichus Koch. Dresden: 1-56. related to Brephosceles (Proctophyllodidae: Alloptinae). Canestrini, G. 1878. Nuove specie do genera Dermaleichus. Bull. Univ. Nebraska State Mus. 8(4): 217-236. Atti del Reale lstituto Veneto di Sci., Lettere, Arti, Ven­ Popp, Egon. 1967. Die Begattung bei den Vogelmilben ezia, ser. 5, 5: 43-70. Pterodectes Robin (Analgesoidea, Acari). Z. Morph. Okol. --. 1886. Famiglia degli Analgesini. Prospetto dell­ Tiere 59: 1-32. 'Acarofauna ltaliana 2: 248, 301-303. Radford, Charles D. 1953. The mites (Acarina: Analgesi­ Canestrini, G., and P. Kramer. 1899. Demodicidae und dae) living on or in the feathers of birds. Parasitol. Sarcoptidae. Das Tierreich 7: 1-193. 43(3,4): 199-230. Dubinin, V. B. 1949. Feather mites of the Procellariiformes ---. 1958. The host-parasite relationships of the feath­ and their particulars. Akad. Nauk S.S.S.R., Zool. Inst., er mites (Acarina: Analgesoidea). Rev. Brasil. Entomol. Parasitol. Sborn. 11: 201-228. 8: 107-170. --. 1950. Feather mites from wintering birds in the Trouessart, E. L. 1885. Note sur la classification des Anal­ low country of Lenkoran. Akad. Nauk S.S.S.R., Baku gesiens et diagnoses d'especes et de genres nouveaux. Inst. Zool., Trudy 14: 58-75. Bull. Soc. Etud. Sci. Angers 14: 46-89. --. 1950. Feather mites of gallinaceous birds of Ka­ ---. 1886. Diagnoses d'especes nouvelles de sarcop­ zakhstan. Akad. Nauk S.S.S.R., Zool. Inst. 8: 62-79. tides plumicoles (Analgesinae). Bull. Soc. Etud. Sci. --. 1951. Feather mites (Analgesidae). Part I. Intro­ Angers 16: 85-156. duction to their study. Fauna S.S.S.R. 6(5): 1-363. ---. 1899. Diagnoses preliminaties d'especes nouvel­ --. 1951. Feather mites of birds of the Barabinskian les d'Acariens plumicoles. Additions et corrections ala Steppes. Akad. Nauk S.S.S.R., Zool. Inst. Parazitol. sous famille des Analgesines. Bull. Soc. Etud. Sci. Sborn. 13: 120-256. Angers 28: 137-199. -. 1952. Feather mites of birds of Wrangel Island. Vassilev, I. D. 1960. Feather mites (Analgesoidea) on some Akad. Nauk S.S.S.R., Zool. Inst., 12: 251-268. birds in Bulgaria. Bulgarian Acad. Sci., Proc. Zool. Inst. Gaud, J. 1952. Sarcoptides plumicoles des oiseaux de 9: 431-437. Madagascar. Mem. Inst. Sci. Madagascar, ser. A 7(1): Vitzthum, Herman. 1921. Acarologische Beobachtungen 81-107. (5. Reihe.) Arch. f. Naturgesch., A 87(4): 1-77. A REVISION OF THE FEATHER MITE GENUS BREPHOSCELES I 171

PARASITE INDEX

afribycls, 144, 153, 165 furclfer, 99, 154, 166 prolatus, 97, 104, 108, 164 Alloptes, 99 geranoxenus, 98, 140, 165 Pterodectes, 99 ALLOPTINAE, 96 haematopl, 98, 144, 148, 166 Pterocolus, 99 anatina, 98, 135, 138, 165 hlstlonotus, 99, 145, 160, 165 pterodromae, 97, 104, 118, 164 anhlmae, 98, 135, 138, 165 hoplopterl, 99, 145, 158, 166 Pterophagus, 99 belonopterl, 98, 145, 155, 165 lnornatus, 99, 145, 158, 165 puffin!, 118, 164 Brephosceles, 97, 98, 99, 100, 113 lambda, 98, 135, 165 selenopeltatus, 104, 123, 164 bllobatus, 97, 98, 124, 131, 164 lanceolatus, 98, 124, 129, 164 stephanlbycls, 144, 150, 166 charadrli, 145, 165 loblvanelll, 99, 145, 162, 166 superbus, 97, 104, 106, 164 chaubaudl, 98, 140, 142, 165 longipenis, 97, 104, 108, 164 marlae, 97, 104, 120, 164 chilensls, 98, 145, 161, 165 longlstriatus, 145, 157, 165 trlangularls, 97, 104, 115, 164 collaracus, 99, 145, 162, 165 lunatus, 97, 104, 113, 164 Trouessartla, 99 constrlctus, 99, 145, 157, 165 marglnlventrls, 98, 124, 164 TROUESSARTINAE, 96 decapus, 97, 104, 164 pachyptllae, 97, 104, 110, 164 tuberl, 145, 155, 165 diomedel, 98, 124, 127, 164 parvatus, 97, 98, 124, 133, 164 turgldus, 98, 144, 150, 166 discidlcus, 98, 135, 165 pelaglcus, 97, 113, 164 vanelll, 145, 154, 166 disjunctus, 97, 98, 124, 129, 164 Proctophyllodes, 90, 92, 96, 99 vlrglnlensls, 99, 145, 161, 165 166 forflclger, 97, 101, 164 PROCTOPHYLLODINAE, 96, 100 172 / BULLETIN OF THE UNIVERSITY OF NEBRASKA STATE MUSEUM HOST INDEX

Afribyx, 154, 165 Dlomedea, 98, 127, 129, 164 Oceanodroma, 97,104,106, 110, 131, africanus africanus, Rhinoptllus, 99, DIOMEDEIDAE, 97, 98, 127, 129, 164 164 154, 166 dublus curonlcus, Charadrl.us, 148, Orthorhamphus, 98, 144, 165 alexandrinus dealbatus, Charadrlus, 165 ostralegus longirostrls; Haematopus, 157, 165 dubius dublus, Charadrius, 148, 165 98, 150, 166 alexandrinus seebohmi, Chara~rius, dublus jerdonl, Charadrius, 148, 165 Pachyptila, 97, 108, 113, 164 157, 165 duvancelil, Hoplopterus, 99, 166 paciflcus chlororhynchus, Puffinus, Anas, 98, 138, 165 Esacus, 98, 144, 165 98, 124, 133, 164 ANATIDAE, 97, 98, 135, 138, 165 externa cervicalls, Pterodroma, 97, paclflcus cuneatus, Puffinus, 123, 164 Anhima, 98, 140, 165 98, 129, 164 palllatus palllatus, Haematopus, 98, ANHIMIDAE, 97, 98, 140, 165 externa externa, Pterodroma, 97, 115, 150, 166 ANSERIFORMES, 97, 98, 165 164 pavonlna regulorum, Balerica, 98 arctica, Gavia, 97, 103, 164 exulans, Diomedea, 98, 129, 164 pecuarius, Charadrlus, 157, 165 armatus, Hoplopterus, 155, 166 Gavia, 97, 101, 103, 164 pelagicus, Hydrobates, 97, 113, 164 atriceps, Xiphiodlopterus, 159, 166 GAVIIDAE, 97, 103, 164 peronll, Charadrius, 157, 165 auritus, Nettapus, 98, 135, 165 GAVIIFORMES, 97, 164 placldus, Charadrius, 148, 165 Balearica, 98, 142, 165 glganteus, Macronectes, 98, 127, 164 platyrhynchos, Anas, 98, 138, 165 Belonopterus, 98, 155, 161, 165 gravls, Puffinus, 98, 120, 133, 164 PROCELLARIIDAE, 97, 98, 108, 113, bewickii, Cygnus, 98, 138, 165 grlseus, Pufflnus, 120, 164 115, 118, 120, 123, 127, 129, 133, biclnctus, Charadrlus, 99, 158, 165 GRUIDAE, 97, 142, 165 164 BURHINIDAE, 97, 144, 165 GRUIFORMES, 97, 98, 165 PROCELLARIIFORMES, 97, 164 Burhlnus, 98, 144, 165 HAEMATOPODIDAE, 98, 144, 150, Pterodroma, 97, 98, 115, 118, 129, bullerl, Pufflnus, 124, 164 166 164 castro bangsl, Oceanodroma, 98, 131, Haematopus, 98, 150 Puffinus, 97, 106, 108, 120, 123, 124, 164 Hemiparra, 155, 166 133, 164 castro leucorhoa, Oceanodroma, 97, Hoplopterus, 99, 155, 158, 166 puffinus, Puffinus, 97, 98, 133, 164 106, 164 Hydrobates, 97, 164 recurvirostris, Esacus, 144, 165 castro, Oceanodroma, 97, 104, 106, HYDROBATIDAE, 97, 104, 106, 110, regulorum, Balearica, 142, 165 164 113, 131, 164 Rhinoptilus, 99, 154, 166 CHARADRIIDAE, 97, 98, 144, 148, immer, Gavia, 97, 101, 103, 164 senegallus, Afribyx, 154, 165 153, 154, 155, 157, 158, 160, 161, lndicus atronuchalls, Loblvanellus, Stephanibyx, 153, 166 162, 164, 165 99, 162, 166 tenuirostris, Puffinus, 97, 98, 106, CHARADRIIFORMES, 97, 98, 165 lateralls, Afribyx, 154, 165 108, 123, 133, 164 Charadrlus, 99, 148, 157, 158, 160, leschenaultii, Charadrius, 160, 165 thoracicus, Charadrius, 157, 165 161, 162, 164, 165, 166 leucoptera hypoleuca, Pterodroma, tricollaris bifrontatus, Charadrius, 99, chllensis cayennensls, Belonopterus, 97, 98, 118, 129, 164 161, 165 98, 155, 165 leucoptera masafuerae, Pterodroma. tricollaris forbesl, Charadrius, 99, chilensis chilensis, Belonopterus, 98, 98, 129, 164 161, 165 161, 165 Loblvanellus, 99, 162, 166 tricollaris tricollaris, Charadrius, 99, chilensls lampronotus, Belonopterus, Macronectes, 98, 127, 164 161, 165 98, 161, 165 magnlrostrls, Orthorhamphus, 144, Vanellus, 155, 166 chlororhynchos, Diomedea, 98, 127, 165 vanellus, Vanellus, 155, 166 164 markhaml tristraml, Oceanodroma, ventralis, Hoplopterus, 99, 158, 166 chrysostoma, Diomedea, 98, 129, 164 97, 110, 164 vermiculatus, Burhinus, 144, 165 collaris, Charadrius, 99, 164, 165 melanophrls melanophris, Diomedea, vociferus ternomlnatus, Charadrlus, cookll deflllppiana, Pterodroma, 98, 98, 129, 164 157, 165 118, 129, 164 wilsonia, Charadrius, 99 melanopterus, Stephanibyx, 153, 166 coronatus coronatus, Stephanlbyx, wilsonia beldingl, Charadrlus, 162, 153, 166 melodus circumcinctus, Charadrius, 165 cornuta, Anhima, 98, 140, 165 157, 165 wilsonia cinnamominus, Charadrlus, crassirostris leucoptera, Hemlparra, melodus melodus, Charadrius, 157, 162, 165 155, 166 165 wilsonia rufinucha, Charadrlus, 162, Cursorius, 154, 166 mongolus, Charadrius, 99 165 Cygnus, 98, 138, 165 mongolus atrifrons, Charadrius, 160, wllsonla wllsonla, Charadrlus, 162, desolata, Pachyptlla, 97, 108, 113, 165 166 164 Nettapus, 98, 135, 165 Xiphidiopterus, 155, 166 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