Proceedings ofthe United StatesNational MuseumSMITHSONIAN INSTITUTION ? WASHINGTON, D.C.
Volume 120 1967 Number 3558
AMBLYCERAN MALLOPHAGA (BITING LICE)FOUND ON THE BUCEROTIDAE (HORNBILLS) '
By Robert E. Elbel ?
Mallophaga of the genera Chapinia and Bucerophagus of the am-blyceran family Menoponidae are found only on hornbills. Thepurpose of this paper is to redescribe and illustrate the known speciesin these genera, describe new species encountered, and comparethe classification of these lice with that of the hornbills, Menoponidaehave been examined from 53 species or subspecies of hornbills (table13). Presented are descriptions and illustrations of 22 species ofMenoponidae of which 17, including 12 new, are species of Chapinia,3 are species or Bucerophagus, and 2 ai*e new species in a new genus,Bucerocolpocephalum.No previous attempt has been made to examine all the Menoponidaefrom the hornbills. Clay (1947) included Chapinia and Bucerophagusin her key to the genera of the Menoponidae, but her figures 8 and 9of the antennae of these genera were transposed accidentally. Thegenus Chapinia was described by Ewing (1927) for his species C.rohusta; later it was described by Bedford (1930) for Menopon bucerotis
' Modified from a doctoral dissertation submitted to the University of Okla-homa, Norman.
' E. and E. Branch, Dugway Proving Ground, Dugway, Utah. 1
2 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 120Kellogg, 1908, and M. lophocerus Bedford, 1920. Hopkins and Clay(1952) included in the genus Chapinia the additional species, Colpo-cephalum hirtum Rudow, 1866, Menopon acutovulvatum Piaget, 1881,and Allomenopon mjobergi Eichler, 1947, but they considered thegeneric position of C hirtum doubtful. In the present study C.hirtum is shown to be a Chapinia, and a neotype is designated;A. mjobergi is shown to be a synonym of M. acutovulvatum. Hopkins(1941) designated a lectotype for Chapinia lophocerus (Bedford),Clay (1949a) designated a lectotype for C. acutovulvata (Piaget), anda lectotype is designated here for C. bucerotis (Kellogg). Piaget(1880) identified a female from Rhyticeros cassidix (Temminck, 1823)as C. hirtum, but Piaget's specimen is shown here to be the new speciesChapinia lydae. The genus Bucerophagus was described by Bedford(1929) for his species B. africanus and for Colpocephalum productumBurmeister, 1838. For the latter species a neotype was erected byConci (1950), and a lectotype was designated by Clay (1951a).Eichler (1947) described a new genus for Menoponforcipatum Nitzsch,1874, but Hopkins and Clay (1952) put M. forcipatum in the genusBucerophagus. A neotype for B. forcipatus (Nitzsch) is designatedhere from Eichler 's material. Clay (1951a) stated that Bucerophagusafricanus and B. productus both infest the two hosts, Bucorvus abys-sinicus (Boddaert, 1783) and B. leadbeateri (Vigors, 1825). Nomorphological or statistical means were found in the present studyto separate the populations of each species on each host so that onlythe two species, Bucerophagus productus and B. africanus, could berecognized.The phylogenetic arrangement of the hornbills (Peters, 1945)shows scant regard for the geographical regions, and the list windsback and forth between the Ethiopian, Oriental, and Australasianregions (table 13). The mallophagan genera studied here, however,fall into definite species-groups confined to the Ethiopian region or tothe Oriental and Australasian regions. It is believed, therefore, thatthe arrangement of the MaUophaga gives more insight into the originof the hornbills than study of the host skins.Classification of the hosts is that proposed by Deignan (1963)except for species not discussed by him, for which Peters (1945) hasbeen followed. Skins of the hosts collected in Thailand are in theU.S. National Museum and were identified by Mr. H. G. Deignan.Collections were made possible by assistance from the U.S. OperationsMission to Thailand and the U.S. National Museum.Grateful appreciation is expressed to Dr. Cluff E. Hopla, OklahomaUniversity, Dr. K. C. Emerson (KCE), Stillwater, Okla., and Dr.Theresa Clay, British Museum (Natural History) (BMNH), fortheir many suggestions and constant help; to Dr. Clay for the loan
NO. 35B8 MALLOPHAGA?ELBEL 3
of specimens in the BMNH, in the Meinertzhagen, in the Piaget,and in the G. H. E. Hopkins (GHEH) collections, the latter from theZoological Museum, Tring, Hertfordshire, England, for obtainingfrom Dr. Eichler and Dr. von Keler the loan of specimens in theZoological Museum, Humboldt University, Berlin, and from Dr.Joao Tendeiro (JT) the loan of specimens in the Centro de Zoologia,Lisboa, Portugal; to Dr. Eric Kjellander for the loan of specimensin the Swedish Museum of Natural History (SMNH); to Dr. RupertL. Wenzel for the loan of specimens in the Chicago Natural HistoryMuseum (CNHM); to Dr. K. C. Emerson and Mr. C. F. W. Muese-beck for the loan of specimens in the U.S. National Museum (USNM),and for mounting mallophagan dried material obtained from thatmuseum. Special thanks are extended to the Chicago Natural His-tory Museum, the Michigan University Museum of Zoology (MMZ),the U.S. National Museum, and Dr. Boonsong Lekagul (BL), Bangkok,Thialand, for permission to examine their collections of hornbill skinsfor Mallophaga; to Dr. Boonsong Lekagul, Mr. H, G. Deignan, Mr.Kitti Thonglongya of Seato Medical Research Laboratory (SMRL),Bangkok, Thailand, Mr. Wanit Songprakob and Mr. Wichit SuwanLaong, both of Songkhla, Thailand, for help in collecting mallophaganfresh material from Thailand; to Dr. Alfred F. Naylor, formerly ofOklahoma University, for help in statistical comparisons of lousepopulations on the hosts, Bucerophagus productus and B. africanus;to the Bernice P. Bishop Museum (BPBM), Honolulu, Hawaii, andthe Prirodoslovni Musej Slovenije (PMS), Ljubljana, Yugoslavia, forthe loan of specimens; and to my wife, Lyda (LE), for help in prepara-tion of the manuscript and in various phases of the study.
Explanation of TermsThe terminology used in this paper agrees with that of Clay (1947)except as noted below."Combs of setae" are rows of short, stout setae, each with thealveoli lying close together and approximately in a straight line, onthe venter of the third femora and the posterolateral areas of oneor more abdominal sternites (figs. 64, 65).
"Brushes of setae" are concentrations of setae on the venter of thethird femora and the posterolateral areas of one or more abdominalsternites. These may take the form of a few widely spaced setaecalled "small scattered brushes" (figs. 23, 24) or a large number ofclosely set setae called "large thick brushes" (figs. 68, 69). Thesetae of the brushes are "normal" if approximately the same lengthand thickness as the surrounding setae or "small" if the majorityare considerably smaller than the surrounding setae.
4 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 120The "preocular slit" is an emargination with approximately equaland parallel margins in the dorsal-lateral margin of the head immedi-ately anterior to the eye (figs. 23, 24).The "preocular notch" is a similar emargination with triangularor semicircular margins (figs. 25, 26, 64-69).The "metasternal plate" on the metastemum was first describedby Bedford (1920) as a 4-sided plate which was normally expandedanteriorly (figs. 26c, 64c, 66c, 68c, 70).The male and female each possess a complete complement ofabdominal segments from the first to the tenth (Cope, 1941).The "pleurites" ("paratergal plates" of some authors) are scleriteswhich pass around the lateral margin of the abdomen and are sepa-rated from the tergites and sternites by clear divisions (Clay, 1954)(figs. 23, 24).The "postspiracular seta" on each side of abdominal tergitesIII-VIII is posterior to the spiracle and is always associated withtwo small setae, the alveoli of the three setae being contiguous (Clay,1954). A similar group of three setae is present on abdominal tergiteII, although there is no functional spiracle on this tergite. In mostAmblycera the postspiracular setae are the most laterad setae of theposterior marginal row (fig. 25p).The "male terminal abdominal tergites" IX and X are almostcompletely fused. "Abdominal sternite VIII" is fragmented intotwo parts which have moved laterad. "Abdominal sternite IX"strengthens the anterior lip of the genital and anal opening (cloaca),and there is no apparent tenth sternite (Cope, 1941).The "basal plate" ("basal apodeme" of Clay, 1956) of the "malegenitaha" supports posteriorly the slender "parameres" laterally andthe "endomeres" centrally (Ewing, 1927).The "sclerite of the male genital sac" is the sclerite on the wallsof the genital sac which is articulated to the basal plate (Clay, 1956)(figs. 18s, 19s).The "lateral horns" of the "male genitaha" are structiu'es on eachside of the large, curved, platelike "endomeres" (Ewing, 1927) (figs.1-3,6,7).The "female terminal abdominal tergites" IX and X are singleplates. "Abdominal sternite VIII" has a pair of gonopods, lyingside by side medially, which are fused at their apices to cover thegenital opening (vulva) between segments VIII and IX, and on eachlateral side of the gonopods is a fragment of sternite VIII (Cope, 1941).The "internal sclerite of female abdominal sternite VIII" appearsto be the sclerite that Clay (1961) calls the internal structure of thefemale genital chamber (figs. 35i, 38i, 39i, 49i).
"Sclerital hooks" are hookhke processes in the female arising on
NO. 3558 MALLOPHAGA?ELBEL 5
either side of the midline of the "ventral sclerite between the vulvaand anus" (fig. 27h).The "anal fringe" ("anal corona" of Ferris, 1923) surrounds thefemale anus on abdominal segment X (figs. 28, 45, 52)
.
"Species-groups" are groups of similar species within a genus.
"Fresh material" indicates that Mallophaga were obtained fromthe host that was collected in the field as contrasted to maUophagan
"dried material" which was obtained from dried museum skins eitherpersonally (REE) or by my wife, Lyda.MethodsDried material was obtained from museum skins by lightly fluffingthe bird feathers, particularly around the neck and lower belly, overa white surface. Emerson (1954) stated that contamination thatoccurred on museum skins was well known and that records of Mal-lophaga so obtained shoiJd be considered questionable. Mallophagathat were obtained from museum skins were considered here to bestragglers unless they belonged to recognized hornbill genera andunless they were represented by other specimens obtained fromadditional skins or fresh material of the same host species. Cor-respondingly, about 20 percent of the dried material was consideredto be stragglers. The mounting procedure was suggested by Dr.K. C. Emerson (in litt.): Mallophaga were placed in 10 percentpotassium hydroxide overnight, transferred to distilled water, andafter one hour the body contents were teased out. Specimenswere placed in fresh 10 percent potassium hydroxide for 6-12 hours,after which they again were transferred to distilled water. Approxi-materly one-half hour later specimens were put into 40 percentethyl alcohol. Fifteen minutes later several drops of carbol fuchsin(Ziehl Nielson) were added and allowed to act for one-half hour.Specimens were placed in 70 percent ethyl alcohol for one-half hour,followed by 95 percent ethyl alcohol for 15 minutes. Next, speci-mens were washed in 100 percent ethyl alcohol for a few minutes andplaced in Beechwood Creosote for one hoiu* to overnight, after whichthey were mounted in Gum Damar or other dried resin media.Drawings were prepared from holo types and allotypes except asnoted in the text. All drawings were prepared with the aid of a300-watt, 35-mm. slide projector as suggested by Dr. K. C. Emerson(in litt.). The monocular microscope with the mounted Mallophagawas turned on its side, the ocular and mirror removed, and the slideprojector placed at the lower end of the microscope so that the fightprojected the image onto a vertical surface, from which the outlinewas traced on Bristol board or drawing velum. Measurements wereobtained by projecting a millimeter scale from a stage micrometer
6 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 120
onto the surface. Details of the Mallophaga were added after themicroscope was uprighted.Measurements for the tables are in millimeters and were made withthe aid of an ocular micrometer.Because of variation in setal number, setae recorded in speciesdescriptions represent the range in numbers of setae from repre-sentative specimens from the material examined.Characters described under genera or species-groups have not beenrepeated for individual specific descriptions. In each genus or species-group the arrangement of the species is based first on morphologicalsimilarity and second on the phylogenetic arrangement of their hosts.Key to Species of Chapinia, Bucerocolpocephalum and Bucerophagus1. Terminal segment of antenna showing definite signs of division either bytransverse line or marginal indentation (figs. 64a, 66a, 68a) 19Terminal segment of antenna without signs of division (figs. 23a, 25a).(Genus Chapinia Ewing). . . 22. Dorsal-lateral margins of head with a preocular slit (figs. 23, 24)
.
(lophocerus species-group) ... 3Dorsal-lateral margins of head with a preocular notch (figs. 25, 26). ... 73. Each lateral margin of abdominal tergites III-VI with a short seta betweenthe spiracle and postspiracular seta (fig. 64 1) 4Abdominal tergites without such short setae (figs. 23, 24).C. robusta Ewing, p. 17.4. Abdominal sternite II with three median rows of setae.C. bucerotis (Kellogg), p. 15.Abdominal sternite II with one median row of setae on posterior margin . 55. Male genitalia with each lateral liorn possessing two sharp posterior points;female anal fringe with fewer than 58 setae (figs. 2, 28).C. lophocerus (Bedford), p. 13.Male genitalia with eacli lateral liorn possessing one or two rounded posteriorpoints; female anal fringe with more than 58 setae 66. Male genitalia with each lateral horn possessing two rounded posteriorpoints; female anal fringe with more than 70 setae (fig. 1).C. fasciati, new species, p. 12.Male genitalia with each lateral horn possessing one rounded posterior point
;
female anal fringe witli 60-64 setae (fig. 3) . C. camuri, new species, p. 15.7. Each lateral margin of abdominal tergites III-VI with a short seta betweenthe spiracle and postspiracular seta (fig. 64 1) 8Abdominal tergites without sucli short setae 118. Male genitalia with endomeres possessing small inner plate and paired outerrims, each with serrulations on posterior inner margin; female with eachlateral projection of ventral sclerite between vulva and anus possessingmore than eight thick, posteriorly directed setae (figs. 71, 72sp).C. waniti, new species, p. 22.Male genitalia with endomeres possessing small or large inner plate and pairedouter rims without serrulations; female with each lateral projection ofventral sclerite between vulva and anus possessing fewer than eight thick,posteriorly directed setae (fig. 43sp) 9
>40. 3558 MALLOPHAGA?ELBEL 7
9. Male genitalia with endomeres possessing paired outer rims and largeshieldlike inner plate with paired lateral flanges and central terminalpoint; female abdominal sternite VIII with more than 32 setae on posteriormargin (figs. 12, 37) C. malayensis, new species, p. 25.Male genitalia with endomeres possessing paired outer rims and small innerplate that is not shieldlike; female abdominal sternite VIII with fewer than32 setae on posterior margin 10[0. Male genitalia with endomeres possessing on each outer rim a triangularinternal knob wider than long; female terminal abdominal tergite with atmost five long setae each side of midline on posterior margin (figs. 8, 33)
.
C. clayae, new species, p. 21.Male genitalia with endomeres possessing on each outer rim a triangularinternal knob longer than wide; female terminal abdominal tergite withat least six long setae each side of midline on posterior margin (figs. 9, 35)
.
C. acutovulvata (Piaget), p. 23.
.1. Abdominal sternite II with at least two median rows of setae 16Abdominal sternite II with one median row of setae on posterior margin.{acutovulvata species-group) ... 1212. Male abdominal sternite II with more than 46 total setae; female analfringe with more than 48 setae 13Male abdominal sternite II with fewer than 44 total setae; female anal fringewith fewer than 46 setae 14L3. Male genitalia with endomeres consisting only of paired outer rims whichare curved inwardly; female abdominal sternite VIII with internal scleritealmost as wide as long (figs. 13, 38i) . . . . C. hoplai, new species, p. 26.Male genitalia with endomeres consisting only of paired outer rims whichare straight and nearly parallel; female abdominal sternite VIII withinternal sclerite much longer than wide (figs. 14, 39i).C. boonsongi, new species, p. 27.L4. Male genitalia with endomeres consisting of paired plates having posteriorextension split; female venter of third femora and abdominal sternitesIV-VI without brushes (figs. 17, 25) . C. traylori, new species, p. 31.Male genitalia with endomeres either lacking paired plates or, if present,posterior extension not split; female venter of third femora and pos-terolateral margins of abdominal sternites IV-VI each with brushes ofnormal setae 15L5. Male venter of third femora and posterolateral margins of abdominalsternites IV-VI each with brushes of normal setae; female abdominalsternite VIII without internal sclerite (fig. 40).C. wenzeli, new species, p. 28.Male venter of third femora and abdominal sternites IV-VI without brushes;female abdominal sternite VIII with internal sclerite having slenderposteriorly divergent margins (fig. 43) . . C. blakei, new species, p. 29.16. Venter of third femora and posterolateral margins of abdominal sternitesIV-VI each with brushes of normal setae 17Venter of third femora and abdominal sternites IV-VI without brushes(figs. 25, 26) C. traylori, new species, p. 31.17. Male genitalia with parameres enlarged anteriorly; female terminal ab-dominal tergite with the two median setae on posterior margin as widelyspaced as four times the distance between each of the 3-4 long setae eachside of midline (figs. 20, 46) C. lydae, new species, p. 32.Male genitalia with parameres slender anteriorly.(hirta species-group) ... 18
8 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 12018. Sclerite of male genital sac large, nearly twice as long as wide; female un-known (fig. ISs) C. muesebecki, new species, p. 34.Sclerite of male genital sac small, about as wide as long; female terminalabdominal tergite with 5 long setae on posterior margin each side of mid-line, these 10 setae being evenh' spaced (figs. 19s, 4S).C. hirta (Rudow), p. 35.19. Venter of third femora and posterolateral margins of fourth abdominalsternite each with three fuU rows of combs (figs. 64, 65).(Genus Bucerocolpocephalum, new genus). . 20Venter of third femora and fourth abdominal sternite without combs.(Genus Bucerophagus Bedford). . . 2120. Male genitaha with endomeres having a pair of posterior points; femaleabdominal sternite VIII with internal triangular sclerite (figs. 49, 57).Bucerocolpocephalum emersoni, new species, p. 37.Male genitalia with endomeres lacking a pair of posterior points; femaleabdominal sternite VIII without internal sclerite (figs. 50, 58).Bucerocolpocephalum deignani, new species, p. 38.21. Venter of third femora and posterolateral margins of abdominal sternitesIV and V, each with brushes of normal setae (figs. 68, 69) 22Venter of third femora and abdominal sternites IV and V without brushes(figs. 66, 67) Bucerophagus forcipatus (Nitzsch), p. 42.22. INIetasternal plate triangular, expanded anteriorly (fig. 70).Bucerophagus productus (Burmeister), p. 44.Metasternal plate trapezoidal, expanded anteriorly (fig. 6Sc).Bucerophagus africanus Bedford, p. 46.Genus Chapinia E^vingFigures 23-26Chapinia Ewing, 1927, p. 88. [Genotype: Chapinia robusta Ewing, 1927.]Allomenopon Bedford, 1930, p. 153. [Genotype: Menopon hucerotis Kellogg,1908.]Head triangular, width 1^ to 2 times that of length. Foreheadmuch narrower anteriorly. Temples expanded. Antennae 4-jointed,third segment constricted at base, and terminal segment capitatewithout signs of division. Antennary fossa deep, covered above byexpansion of lateral margin of head, posterior margin of which bearseye with double cornea. Dorsal-lateral margin of forehead anteriorto eye with preocular sUt or shallow notch. Gidar region with 3-7setae varying in length on each lateral margin. Pronotum expandedanteriorly with posterior marginal row of long setae. Metanotumexpanded posteriorly with posterior marginal row of long setae andtwo short setae on each lateral margin. Metanotum separated frommesonotum and from pleurites. Thoracic sternal plates as shown infigures 23b, 25b, and 26c. Metasternal plate trapezoidal, expandedanteriorly, with 6-22 setae. Venter of third femora and posterolateralmargins of abdominal sternites IV-VI, each with or without brushesof normal setae. Abdominal segments consist of tergites, sternites,and pleurites, the latter without prolongation of postero ventral angles.
NO- 3558 MALLOPHAGA?ELBEL 9Abdominal tergites each with a posterior marginal row of setae, themost laterad being the postspiracular seta. Each lateral margin ofabdominal tergites II-VI with or without a short seta between thespiracle and postspiracular seta. Stemites and pleurites each with aposterior marginal row of long setae and with numerous shorter setae.Male terminal abdominal sternites VIII and IX fused with partialdivision only from sternite VII (figs. 4, 10, 22, 24, 26). Male genitahaas illustrated for each species, vrith. parameres either expanded an-teriorly or spht posteriorly or both. Female terminal abdominalsegments as illustrated for each species, ^si.ih lateral processes arisingfrom ventral sclerite between vulva and anus, with long stout setaebut never strong spines. Females larger than males, usually \\-ithmore abdominal sternal setae, but general shape and chaetotaxysimilar to that of males except for terminal abdominal segments.Both Ewdng (1927) and Bedford (1930) stated that the ptero-thorax was undivided. As noted by Cope (1941), the sclerotizedmedian button behind the prothorax (fig. 25m) is a vestige of themesonotum; the supposed mesonotum, the narrow sclerotized bandposterior to this button, is a mere extension of the subcoxae. Ewing(1927) stated that the abdomen consisted of 9 segments in the femaleand 10 segments in the male, but as shown by Cope (1941),the abdomen of both the male and female has 10 segments each.Chapinia resembles most closely Bucerophagus (figs. 66-69) butdiffers in several characters: The terminal segment of the antennashows no sign of division in Chapinia, but there are definite signs ofdivision into two parts either by transverse fine or marginal indenta-tion in Bucerophagus. The venter of the third femora may havebrushes of normal setae in Cltapinia and Bucerophagun; similar brushesare present on posterolateral margins of abdominal sternites IV-VIin Chapinia but abdominal sternites IV and V in Buceroploagus. Eachlateral margin of abdominal tergites II-VI may have a short setabetween the spiracle and postspiracular seta in Cliapinia, but 1-5short setae may be present on margins of abdominal tergites II-\TIIin Bucerophagus. Male terminal abdominal sternites VIII and IXare fused in Chapinia with partial division only from abdominalsternite VII, but abdominal sternites VIII and IX may be fused inBucerophagus with a complete division from abdominal sternite VII.Male genitaha have parameres slender or expanded anteriorly inChapinia but branched anteriorly in Bucerophagus. Lateral processesarising from the ventral sclerite between the female vulva and anushave long stout setae in Chapinia but long stout setae and strongspines in Bucerophagus.The male genitalia and details of the male and female terminalabdominal segments are the best characters for separating species
10 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 120
of Chapinia. Other characters useful in species separation are: Thepresence or absence of brushes of normal setae on the venter of thethird femora and posterolateral margins of abdominal sternitesIV-VI; the presence or absence of a short seta on each lateral marginof abdominal tergites III-VI between the spiracle and postspiracularseta; the number of median rows of setae, and the total number ofsetae on abdominal sternite II. The number and length of setae onthe lateral margins of the gular region are too variable to be of muchuse in separating species.For convenience of classification the species of Chapinia have beenarranged into species-groups.Hosts : Species of Chapinia have been found on the genera Tockus,Anorrhinus, Penelopides, Rhyticeros, Anthracocerus, Bycanistes,Ceratogymna, and Buceros of the avian family Bucerotidae.The lophocerus Species-GroupSpecies similar in shape to Chapinia rohusta (figs. 23, 24). Dif-fering from other species-groups in the following combination ofcharacters: Dorsal-lateral margins of head with preocular slit; venterof third femora and posterolateral margins of abdominal sternitesIV-VI each with brushes of normal setae; each lateral margin of ab-dominal tergites II-VI with or without a short seta between thespiracle and postspiracular seta; females with more abdominal sternalsetae than males; abdominal sternite II with either one or threemedian rows of setae; male genitalia with lateral horns on each sideof endomeres and with parameres enlarged anteriorly, not split pos-teriorly; females with sclerital hooks on each side of midhne of ventralsclerite between vulva and anus; female abdominal sternite VIIIwith most of setae similar in size to setae on posterior margin.Hosts: Species of the lophocerus species-group have been foundon the genera Tockus, Bycanistes, and Ceratogymna of the avianfamily Bucerotidae.Species of the lophocerus species-group are aU similar in size exceptthat both sexes of Chapinia camuri are smaller than correspondingsexes of other species, and males of C. rohusta are larger than othermales. The small size of C. camuri might be expected since its host,the 15-inch Tockus camurus, is the smallest known hornbill. Theventer of the third femora and posterolateral margins of abdominalsternites IV-VI each have large thick brushes of normal setae inC. hucerotis but small scattered brushes of normal setae in other speciesof the lophocerus species-group, although the brushes are slightlythicker in C. rohusta. Each lateral margin of abdominal tergitesII-VI has a short seta between the spiracle and postspiracular seta in
NO. 3558 MALLOPHAGA?ELBEL H
all species except in O. robusta. Abdominal sternite II has more totalsetae in both sexes of C. bucerotis and O. robusta than in correspondingsexes of C.fasciati, C. lophocerus, and O. camuri; this sternite has onemedian row of setae in all species except C. bucerotis, which has threemedian rows. The male genitalia have each lateral horn possessingposterior points in all species except C. robusta, which has one sharpmedian point crossing the broad endomeres; the posterior points aresharp in C. bucerotis and C. lophocerus but rounded in C. fasciati andC. camuri] C. camuri has only one posterior point instead of two as inthe other species; the two points do not reach the slender endomeresin C. lophocerus, but one point crosses the broad endomeres in C.bucerotis (figs. 1-7). In the female the ventral sclerite between thevulva and anus is elevated medially between the sclerital hooks inC. fasciati, C. bucerotis, and C. robusta, is elevated only slightly inO. camuri, and is not elevated in C. lophocerus. The female anal fringehas the most setae in C. fasciati and the fewest in C. lophocerus.Clay (1958) treated populations of the ischnoceran genus Degeeriellaas subspecies when the male genitalia were apparently identical ordiffered only in a minor degree and when there were other minormorphological differences. Because of similarity of the genitalia,Chapinia fasciati and C. camuri could be considered subspecies ofG. lophocerus. This would express the similarity of their six hostspecies, which are all members of the Ethiopian genus Tockus. Clay(1958) pointed out that the genitalia, particularly in the Amblycera,might show only minor differences throughout a genus or species-groupand great differences in other groups. She therefore concluded thatdifferentiation of the genitalia has taken place at different rates indifferent groups. Similarly, Johnson (1960) stated that evolution andmorphological divergence would not be expected to proceed at thesame rate for all free-living species. It would seem that the similarityin the genitalia of C. fasciati, C. lophocerus, and 0. camuri wouldindicate either that evolution has not proceeded as rapidly in thesespecies or that they have not been isolated as long as other species ofChapinia. Clay (1958) stated that if subspecies were populations thatwould interbreed under natural conditions if they occurred sym-patrically, any morphological differences which might prevent inter-breeding should be considered as specific characters. Johnson (1960)believes that there is little possibility of finding interbreeding popu-lations among lice which are isolated on their hosts. She stated thatit would be desirable to treat all stable recognizable forms of Anopluraand MaUophaga as species. Clay (1962) consequently stated thatthe subspecific category might be useful in some of the ischnocerangenera, but its application in the Amblycera is less satisfactory and
12 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 120
should not be used until more is known about the relationships betweenpopulations in this superfamily.The members of the lophocerus species-group are arranged accordingto the phylogeny of their hosts, since this order agrees with the morpho-logical similarities or the parasites.Chapinia fasciatif new speciesFigure 1Male: Smaller than Chapinia robusta in all measurements; smallerthan C. lophocerus in all measurements except width of metathorax(table 1). Venter of third femora and posterolateral margins ofabdominal sternites IV-VI each with small scattered brushes ofnormal setae. Each lateral margin of abdominal tergites II-VI witha short seta between the spiracle and postspiracular seta. Abdominalsternite II with 32^2 total setae and one median row of setae onposterior margin. Terminal abdominal segments similar to those ofC. lophocerus. Genitalia as shown in figure 1, each lateral horn withtwo rounded posterior points.Female: Larger than Chapinia robusta in all measurements exceptwidth of metathorax; smaller than C. lophocerus in aU measurementsexcept total length and width of prothorax (table 2). Resembles themale except that abdominal sternite II has 54 total setae. Terminalabdominal segments similar to those of C. lophocerus. Ventralsclerite between vulva and anus elevated medially between scleritalhooks. Anal fringe with 72-86 setae.Discussion : Chapinia fasciati resembles most closely C. lophocerus.The male genitaUa have each lateral horn possessing two posteriorpoints which are rounded in C. fasciati but sharp in C. lophocerus.The ventral sclerite between female vulva and anus is elevatedmedially between the sclerital hooks in C. fasciati but not elevated inC. lophocerus. The female anal fringe has more than 70 setae inC. fasciati but fewer than 58 setae in C. lophocerus.Material examined: 27 males and 34 females from fresh and driedmaterial collected in the Ethiopian region.Type host: TocJcus fasciatus fasciatus (Shaw, 1811).Type material: Holotype male and allotype female from Eden,French Cameroons, Africa, collected by J. Mouchet, BMNH. Para-types: 18 males and 15 females from French Cameroons, Africa,collected by J. Mouchet, BMNH; 1 male from Kasongo, BelgianCongo, Africa, Nov. 13, 1959, collected by P. L. G. Benuit, JT;3 males and 10 females from CNHM skins from Entebbe, Uganda,Africa, 1895-1916, collected by F. J. Jackson and others, REE; 2males and 1 female from CNHM skins from Bitya, Cameroons,
NO. 3558 MALLOPHAGA?ELBEL 13Africa, 1924-1927, collected by O. L. Bates, REE; 1 female fromCNHM skin from Yokadouma, French Cameroons, Africa, Oct. 19,1946, collected by A. I. Good, REE; 1 female from CNHM skinsfrom Bwamba, Ruwenzori, Uganda, Africa, 1944-1946, collected byV. Someren, REE; 1 male from CNHM skins from Ebolowa, FrenchCameroons, Africa, 1952-1953, collected by A. I. Good, REE; 1 maleand 3 females from USNM skins from Congo, Africa, 1917, collectedby C. R. Aschemeier, REE; from Tockus alhoterminatus suahelicus(Neumann, 1905): 1 female from CNHM skins from Sokoke Forest,Kenya, Africa, June 1932, collected by V. Someren, REE; 1 femalefrom USNM skins from Nairobi, Kenya, Africa, 1909, collected byLoring and Mearns, REE.
Chapinia lophocerus (Bedford)Figures 2, 4, 27, 28Menopon lophocerus Bedford, 1920, p. 717, pis. 1 (fig. 1), 3 (fig. 1). [Type host:Lophoceros leucomelas= Tockus flavirostris leucomelas (Lichtenstein, 1842).]Chapinia lophocerus (Bedford)?Hopkins and Clay, 1952, p. 67.Bedford did not designate a type from his material which containeda pair of MaUophaga from Lophoceros leucomelas= Tockus flavirostrisleucomelas (Lichtenstein, 1842), a pair from L. epirhinus? Tockusnasutus cafler (SundevaU, 1851), and two males and one female fromL. erythrorhynchos= Tockus e. erythrorhynchus (Temminck, 1823).A lectotype was designated by Hopkins (1941) from the host,Lophoceros leucomelas, since the male from that host agreed best withBedford's figure of the male genitaha.Male: Smaller than Chapinia robusta in all measurements (table 1).Venter of third femora and posterolateral margins of abdominalsternites IV-VI each with small scattered brushes of normal setae.Each lateral margin of abdominal tergites II-VI with a short setabetween the spiracle and postspiracular seta. Abdominal sternite IIwith 28-42 total setae and one median row of setae on posteriormargin. Terminal abdominal segments as shown in figm-e 4. Genitahaas shown in figure 2, each lateral horn with two sharp posteriorpoints which do not reach the slender endomeres.Female: Larger than Chapinia robusta in aU measurements exceptwidth of prothorax and width of metathorax (table 2). Resemblesthe male except that abdominal sternite II has 34-54 total setae.Terminal abdominal tergite with 22-30 setae, short setae alternatingwith long, on posterior margin; ventral sclerite between vulva andanus not elevated medially between sclerital hooks (fig. 27). Analfringe with 42-56 setae (fig. 28).
14 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 120Discussion: Bedford (1920) gave the followingmeasurements (in mm).male femalelength of head 0.25 0.33width of head 0.53 0.60width of prothorax 0.38 0.43width of metathorax 0.55 0.71width of abdomen 0.85 1.15total length 1.74 2.36Except for females being larger in abdominal width and total length,these measurements faU within the ranges of Chapinia lophocerus(tables 1, 2). C. lophocerus resembles most closely C. fasciati.The venter of the third femora and posterolateral margins of abdominalsternites IV-VI each have smaU scattered brushes of normal setae inC. lophocerus, C. jasciati, and C. camuri but large thick brushes ofnormal setae in C. bucerotis. Abdominal sternite II of both sexeshas fewer than 58 total setae and one median row of setae in C.lophocerus, C. fasciati, and C. camuri but more than 58 total setaeand three median rows of setae in C. bucerotis. The male genitaliahave each lateral horn possessing posterior points that do not reachthe slender endomeres in C. lophocerus, C. Jasciati, and C. camuri butone sharp point that crosses the broad endomeres in C. bucerotis;the posterior two points are sharp in C. lophocerus but rounded inC. Jasciati, and there is only one rounded posterior point in C. camuri.The ventral sclerite between female vulva and anus is not elevatedmedially between the sclerital hooks in C. lophocerus, is elevatedshghtly in C. camuri, but is more elevated in C. fasciati and C. bucerotis.The female anal fringe has fewer than 58 setae in C. lophocerus, at least72 setae in C. fasciati, and 58-72 setae in C. camuri and C. bucerotis.Material examined: 6 males and 19 females from fresh and driedmaterial collected in the Ethiopian region; lectotype male and syntypefemale from Transvaal, South Africa, September 1917, collected byG. A. H. Bedford, GHEH; from the type host: 1 male and 4 femalesfrom Pretoria Zoo, South Africa, Feb. 10, 1938, collected by G. A. H.Bedford, GHEH; from Tockus n. nasutus (Linnaeus, 1766): 1 maleand 1 female from Maroua, North French Cameroons, Africa, 1959,collected by J. Mouchet, BMNH 1960-105; 1 male and 2 females fromMansoa, Portuguese Guinea, Africa, Feb. 14, 1951, collected by J.Tendeiro, JT; from Tockus e. erythrorhynchus (Temminck, 1823): 1male and 4 females from Somaliland, Africa, February 1949, Meinertz-hagen 18708, BMNH; 1 male from USNM skins from Ethiopia,Africa, 1912, collected by Childs Frick, KEE; from Tockus f. fia-virostris (Riippell, 1835): 1 female from USNM skins from Ethiopia,Africa, 1912, collected by Childs Frick, REE; from Tockus deckeni(Cabanis, 1869): 6 females from Koka, Ethiopia, Africa, Dec. 13,1960, coUected by Savo Brelih, PMS.
NO. 3558 MALLOPHAGA?ELBEL 15Drawings were made of the lectotype male and a female collected inthe Pretoria Zoo. Specimens in GHEH.Chapinia camuri, new speciesFigure 3Male: Smaller than Chapinia rohusta in all measurements; smallerthan C. lophocerus in all measurements except length of head and widthof prothorax (table 1). Venter of third femora and posterolateralmargins of abdominal sternites IV-VI each with small scatteredbrushes of normal setae. Each lateral margin of abdominal tergitesII-VI with a short seta between the spiracle and postspiracular seta.Abdominal sternite II with 37 or 38 total setae and one median row ofsetae on posterior margin. Terminal abdominal segments similar tothose of C. lophocerus. Genitalia as shown in figure 3, each lateralhorn with one rounded posterior point.Female: Smaller than other Chapinia in aU measurements exceptlength of head and width of abdomen (table 2). Resembles the maleexcept that abdominal sternite II has 44-56 total setae. Terminalabdominal segments similar to those of C. lophocerus. Ventralsclerite between vulva and anus slightly elevated medially betweensclerital hooks. Anal fringe with 60-64 setae.Discussion: Although smaller in size, Chopinia camuri resemblesmost closely C. lophocerus. The male genitalia have each lateral hornpossessing one rounded posterior point in C. camuri but two sharp pos-terior points in C. lophocerus. The ventral sclerite between femalevulva and anus is elevated medially only sUghtly between the scleritalhooks in C. camuri and is not elevated in C. lophocerus. The femaleanal fringe has more than 58 setae in C. camuri but fewer than 58 setaein C. lophocerus.Material examined : 2 males and 2 females from fresh material col-lected in the Ethiopian region.Type host: Tockus camurus camurus Cassin, 1857.Type material: Holotype male and allotype female from Ambam,French Cameroons, Africa, 1955, collected by J. Mouchet, BMNH.Paratypes : 1 male and 1 female with same data.Chapinia bucerotis (Kellogg)Figures 5, 6, 29, 30Menopon bucerotis Kellogg, 1908, p. 54, pi. 7 (fig. 12) .?Bedford, 1920, pi. 3 (fig.2) (male genitalia). [Type host: Bycanistes cristatus= Bycanistes brevisomissus Peters, 1945.]Chapinia bucerotis (Kellogg).?Hopkins and Clay, 1952, p. 67.Kellogg did not designate a type from his material, which contained1 male and 1 female syntypes on slides and about 40 syntypes in
X6 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 120
alcohol. The slide specunens were remounted and the male is desig-nated hereby as a lectotype; the slide has been so labeled. Approxi-mately one-half of the syntype material formerly in alcohol wasmounted.Male: Smaller than C^a^im'd ro6ti?to in all measurements (table 1).Venter of third femora and posterolateral margins of abdominalsternites IV-VI each with large thick brushes of normal setae. Eachlateral margin of abdominal tergites II-VI with a short seta betweenthe spiracle and postspiracular seta. Abdominal sternite II with60-82 setae and three median rows of setae. Terminal abdominalsegments as shown in figure 5. Genitalia as shown in figure 6, eachlateral horn with two sharp posterior points, one of which crosses thebroad endomeres.Female: Approximately the same size as Chapinia robusta (table 2).Resembles the male except that abdominal sternite II has 68-104total setae. Terminal abdominal tergite with 22-32 setae, shortsetae alternating with long, on posterior margin; ventral scleritebetween vulva and anus elevated medially between sclerital hooks(fig. 29). Anal fringe with 58-72 setae (fig. 30).Discussion: Kellogg (1908) gave the foUomngmeasurements (inmm).
NO. 3558 MALLOPHAGA?ELBEL 17
as much as in C. robusta but is not elevated in C. lophocerus. Thefemale anal fringe has at least 58 setae in C. bucerotis but at most56 setae in C. lophocerus.Material examined: 34 males, 29 females, and approximately 20specimens in alcohol from fresh and dried material collected in theEthiopian region; lectotype male and syntypes, 12 males, 13 females,and about 20 syntypes in alcohol from Kilimanjaro, Tanganyika,Africa, collected by Sjostedt, SMNH; from Bycanistes bucinatorsharpii (Elliot, 1873): 1 female from CNHM skin from MountTandan, Mouila, Gabon, Africa, June 9, 1951, collected by H. A.Beatty, REE; 1 female from USNM skins from Congo, Africa, 1917-1918, collected by C R. Aschemeier, REE; from Bycanistes bucinatorduboisi X sharpii: 1 female from CNHM skin from Yaounde, FrenchCameroons, Africa, July 12, 1948, collected by A. I. Good, REE;from Bycanistes bucinator duboisi W. Sclater, 1884: 1 male fromCNHM skins from Elat, French Cameroons, Africa, collected byRev. M. Fraser, REE; from Bycanistes b. bucinator (Temminck,1824): 1 male and 1 female from Pietermaritzburg, South Africa,1917, GHEH; 3 males and 2 females from CNHM skins from Kenya,Africa, 1918-1922, collected by V. Someren, REE; from Bycanistesc. cylindricus (Temminck, 1831): 1 male and 1 female from CNHMskins from Liberia, Africa, February-June 1948, collected by H. A.Beatty, REE; from Bycanistes cylindricus albotibialis (Cabanis andReichenow, 1877) : 2 males and 1 female from Mbalmayo, FrenchCameroons, Africa, collected by J. Mouchet, BMNH; 1 male fromCNHM skin from French Cameroons, Africa, July 8, 1907, REE;1 male from CNHM skin from French Cameroons, Africa, June 25,1940, collected by A. I. Good, REE; 1 male from CNHM skin fromUganda, Africa, July 15, 1945, collected by V. Someren, REE; fromBycanistes subcylindricus subqtiadratus Cabanis, 1880: 6 males and 6females from Uganda, Africa, April 1936, Meinertzhagen 7674,7708, 7709, BMNH; 1 male from CNHM skin from Kampala, Uganda,Africa, Sept. 2, 1918, collected by V. Someren, REE; 1 male and 2females from USNM skins from Uganda, Africa, June 1920, collectedby H. C. Raven, REE; from Bycanistes b. brevis Friedmann, 1929:2 males from CNHM skin from Mount Kenya, Kenya, Africa, Novem-ber 1946, collected by V. Someren, REE.Drawings were made of the lectotype male and the syntype femalemounted on the same slide. Specimens in SMNH.Chapinia robusta EwingFigures 7, 23, 24, 31, 32Chapinia robusta Ewing, 1927, p. 89. [Type host: Ceratogymna atrata (Temminck,1835).]Chapinia robusta Ewing,?Hopkins and Clay, 1952, p. 68.221-522?67 2
18 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 120Through the courtesy of Dr. K. C. Emerson, BMNH specimensfrom the type host, here examined, were determined to be conspecificwith the USNM holotype and allotype (USNM 40137, Nytonga,Congo, Africa, Nov. 3, 1917, collected by E. A. Chapin).Male: As illustrated in figure 24. Larger than other species ofthe lophocerus species-group in all measurements except width ofabdomen; approximately the same size as Chapinia traylori (table 1).Venter of third femora and posterolateral margins of abdominalsternites IV-VI each with small scattered brushes of normal setae.Each lateral margin of abdominal tergites II-VI without a shortseta between the spiracle and postspiracular seta. Abdominalsternite II with 48-66 total setae and one median row of setae onposterior margin. Terminal abdominal segments as shown in figure24c. Genitalia as shown in figure 7, each lateral horn with one largesharp point which crosses the broad endomeres.Female: As illustrated in figure 23. Larger than Chapinia camuriin all measurements except length of head; approximately the samesize as other species of the lophocerus species-group; slightly smallerthan C. traylori in all measurements except length of head (table 2).Resembles the male except that abdominal sternite II has 56-68 totalsetae. Terminal abdominal tergite with 20-24 setae, short setaealternating with long, on posterior margin; ventral sclerite betweenvulva and anus elevated medially between sclerital hooks (fig. 31).Anal fringe with 56-62 setae (fig. 32).Discussion: Ewing (1927) gave the follo^^dng measurements (in mm).male
NO. 3558 MALLOPHAGA?ELBEL 19
medially between the sclerital hooks slightly more in C. rohustathan in G. hucerotis.Material examined: 4 males and 5 females from fresh and driedmaterial collected in the Ethiopian region; from the type host: 3males and 3 females from Ambam, French Cameroons, Africa, 1955,collected by J. Moiichet, BMNH; 1 female from CNHM skin fromFougamou, Gabon, Africa, Aug. 4,, 1951, collected by H. A. Beatty,REE; from Ceratogymna elata (Temminck, 1831): 1 male and 1female from Konn, French Cameroons, Africa, Apr. 26, 1947, collectedby V. Aellen, BMNH 1954-487.Drawings were made of a male and a female from the type hostcollected in Ambam, French Cameroons, Africa. Specimens inBMNH. The acutovulvata Species-GroupSpecies similar in shape to Chapinia traylori (figs. 25, 26) . Differingfrom other species-groups in the following combination of characters:Dorsal-lateral margins of head with a preocular notch; venter of thirdfemora and posterolateral margins of abdominal sternites IV-VI eachwith or without brushes of normal setae; each lateral margin of ab-dominal tergites II-VI with or without a short seta between thespiracle and postspiracular seta; females usually with more abdominalsternal setae than males; abdominal sternite II usually with onemedian row of setae on posterior margin; male genitalia withoutlateral horns on each side of endomeres and with parameres enlargedanteriorly, split posteriorly; females without sclerital hooks on eachside of midline of ventral sclerite between vulva and anus; femaleabdominal sternite VIII with most of setae much shorter than thoseon the posterior margin.Hosts: Species of the acutovulvata species-group have been foundon the genera Tockus, Anorrhinus, Penelopides, Rhyticeros, Anthra-coceros, and Buceros of the avian family Bucerotidae.Species of the acutovulvata species-group are all similar in sizeexcept that males of Chapinia wenzeli are smaller than other males,and females of C. traylori are larger than other females. The venterof the thu'd femora and posterolateral margins of abdominal sternitesIV-VI each have large thick brushes of normal setae in both sexes ofC. waniti, C. acutovulvata, C. malayensis, and C. hoplai; small scatteredbrushes of normal setae in females of C. hlakei and in both sexes ofC. clayae, C. hoonsongi, C. wenzeli, and C. lydae; brushes are absentin the male of C. hlakei and in both sexes of C. traylori. Each lateralmargin of abdominal tergites II-VI has a short seta between thespiracle and postspiracular seta in C clayae, C. waniti, C. acutovulvata,
20 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 120and C. malayensis. Abdominal sternite II has more total setae inboth sexes of C. waniti, C. acutovulvata, and C. hoplai than in corre-sponding sexes of C. malayensis, G. wenzeli, C. hlakei, and C. traylori;more total setae in females than in males except in C. lydae; andsetae are arranged in one median row except for one or two medianrows in C. waniti, two median rows in C. traylori, and three medianrows in C. lydae. The male genitalia have endomeres with an innerplate and paired outer rims in C. clayae, C. waniti, C. acutovulvata,C. malayensis, and C. lydae, the paired outer rims possessing internalknobs only in C. clayae and C. acutovulvata and serridations only inC. waniti; the inner plate possessing paired lateral flanges and centralterminal point only in C. malayensis; only paired outer rims in C.hoplai, C. hoonsongi, and C. wenzeli, the posterolateral margin beingsplit in C. wenzeli; and only paired plates in C. hlakei and C. traylori.The female terminal abdominal tergite has on the posterior marginat most 24 setae except for C. waniti, C. acutovulvata, and C. hoplaiwith at least 24 setae; of these setae approximately two-thirds arelong and one-third are short in C. waniti, C acutovulvata, C. malayensis,C. hoonsongi, and C. lydae; approximately one-half are long and one-half are short in C. clayae, and C. hoplai; these setae are arrangedwith at least five long setae on each side of the midlme in C. acutovulvataand C. malayensis but at most six setae in all other species of theacutovulvata species-group. In the female the ventral sclerite be-tween the vulva and anus is curved sharply on the anterior marginin C. clayae and C. acutovulvata but is only slightly curved in the otherspecies of the group; on each lateral projection of this sclerite thereare 4-6 thick, posteriorly directed setae except in C. waniti whichhas 10 or 11 and in C. traylori which has 2 or 3 (figs. 43 sp, 44 sp, and72 sp). The female abdominal sternite VIII has on the posteriormargin the most setae in 0. malayensis with more than 34 and thefewest in C. hlakei with at most 18. The female anal fringe has atleast 44 setae in C. waniti, C. acutovulvata, C. malayensis, C. hoplai,C. hoonsongi, and C. lydae but at most 44 setae in C. clayae, C. wenzeli,C. hlakei, and C. traylori.Were the species of the acutovulvata species-group arranged accordingto the phylogeny of their hosts (Peters, 1945), the order would be:C. clayae, C. waniti, C. wenzeli, C. hlakei, C. lydae, C. hoonsongi,C. malayensis, C. acutovulvata, C. hoplai, and C. traylori, rather thanG. clayae, C. waniti, C CLCutovulvata, C. malayensis, C. hoplai, C.hoonsongi, C. wenzeli, C. hlakei, C. traylori, and C. lydae, which isbased on morphological similarities of the lice.
NO. 3558 MALLOPHAGA?ELBEL 21Chapinia clayae, new speciesFigures 8, 33, 34Both sexes are smaller than corresponding sexes of Chapinia trayloriin all measurements except length of head (tables 1, 2).Male: Venter of third femora and posterolateral margins of abdom-inal sternites IV-VI each with small scattered brushes of normal setae.Each lateral margin of abdominal tergites II-VI with a short setabetween the spiracle and postspiracular seta. Abdominal sternite IIwith 38-50 total setae and one median row of setae on posteriormargin. Terminal abdominal segments similar to those of C. acuto-vulvata. Genitalia as shown in figure 8, endomeres with small innerplate and paired outer rims.Female: Resembles the male except that abdominal sternite II has66-76 total setae. Terminal abdominal tergite with 12 long and 12short setae on posterior margin; abdominal sternite VIII with 22-30setae on posterior margin (fig. 33). Anal fringe with 38-44 setae(fig. 34).
^Discussion: Although smaller in size, Chapinia clayae resemblesmost closely C. acutovulvata. The venter of the third femora andposterolateral margins of abdominal sternites IV-VI each have smallscattered brushes of normal setae in C. clayae but large thick brushesof normal setae in C. acutovulvata. The male genitaha have endomerespossessing on each outer rim a triangular internal knob that is widerthan long in C clayae but longer than wide in C. acutovulvata. Thefemale terminal abdominal tergite has on the posterior margin at most24 setae, of which approximately one-half are long and one-half areshort, in C. clayae, but at least 26 setae, of which approximatelytwo-thirds are long and one-third are short, in C. acutovulvata. Inaddition this margin has on each side of the midline at most four longsetae in C. clayae but at least six long setae in C. acutovulvata. Thefemale anal fringe has at most 44 setae in C. clayae but at least 44setae in C. acutovulvata.Material examined: 19 males and 29 females from fresh and driedmaterial collected in India and Nepal.Type host: Tockus birostris (Scopoli, 1786).Type material: Holotype male, allotype female, and 2 male and 4female paratypes on same slide from Rajputana, India, March 1937,Memertzhagen 8855-8856, BMNH. The holotype and allotypeare each the second from the right in the rows of males and femalesas seen under the microscope. Paratypes: 5 males and 12 femaleswith same data except Meinertzhagen 8932; 1 male from Nepal,
22 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 120December 1935, Meinertzhagen 4859, BMNH; 4 males and 5 fe-males from Nepal, February 1936, Meinertzhagen 4858, BMNH;1 female from CNIIM sldns from Kotla, Kangra, East Pmijab, India,1946 and 1948, collected by W. Koelz, REE; 1 male and 1 femalefrom CNHM skins from Bheragliat, Central Provinces, India, March-April 1946, collected by W. Koelz and R. Chand, REE; 1 male fromCNHM skins from Behvani, KisU, Central Provinces, India, J\dy-August 1946, collected by W. Koelz, REE; 1 female from CNHMsldn from Kanha, Central Provinces, India, Aug. 29, 1946, collectedby R. Chand, REE; 2 males from CNHM sldns from Kalnali, UnitedProvinces, India, February 1947, collected by W. Koelz, REE; 1 maleand 1 female from CNHM skins from Nichland, United Provinces,India, February 1947, collected by W. Koelz, REE; 1 female fromCNHM skins from Simra, Nepal, Mar. 4, 1947, collected by W. Koelzand R. Chand, REE; 1 female from CNHM sldns from Baihar,Balaghat, India, January-February 1949, collected by R. L. Flem-ming, REE; 1 female from USNM skins from India, 1898, 1946-1948,REE; from Tockus g. griseus (Latham, 1790): 1 male from CNHMskins from Nilambus, Madras, India, February-March 1937, collectedby W. Koelz, REE.Chapinia clayae is named for Dr. Theresa Clay of the BritishMuseum (Natiu-al History) in appreciation for her continuous assist-ance throughout the study, for the loan of hornbiU Menoponidaefrom the BMNH, and for helping to obtain the loan of hornbillMenoponidae from other museums.Chapinia tvaniti, new speciesFigures 71, 72Male: Larger than Chapinia traylori in aU measurements exceptwidth of prothorax and width of metathorax (table 1). Venter ofthird femora and posterolateral margins of abdominal sternitesIV-VI each with large thick brushes of normal setae. Each lateralmargin of abdominal tergites II-VI with a short seta between thespiracle and postspiracular seta. Abdominal sternite II with 68-78total setae and one or two median rows of setae. Terminal abdominalsegments similar to those of C. acutovulvata. Genitalia as shown infigiu-e 71, endomeres Avith small inner plate and paired outer rims.Female: Smaller than Chapinia traylori in all measurements exceptlength and w^dth of head (table 2). Resembles the male except thatabdominal sternite II has 92-104 total setae. Terminal abdominaltergite with 14-18 long and 10-12 short setae on posterior margin;abdominal sternite VIII Avith 28-34 setae on posterior margui (fig. 72)
;
each lateral projection of the ventral sclerite between vulva and anus
NO. 35D8 MALLOPHAGA?ELBEL 23
with 10-11 thick, posteriorly directed setae (fig. 72 sp). Anal fringesimilar to that of C. acutovulvata, with 50-64 setae.Discussion: Chaxnnia vjaniti resembles most closely C. acutovulvata.Abdominal sternite II of both sexes has more total setae in C. wanitithan in corresponding sexes of C. acutovulvata. The male genitaliahave endomeres possessing on each outer rim serrulations on theposterior inner margin in C. waniti but a triangular internal knob inC. acutovulvata. Each lateral projection of the ventral scleritebetween female vulva and anus has more than eight thick, posteriorlydirected setae in C. waniti but fewer than eight in all other Chajnnia.The female anal fringe has at least 56 setae in C. waniti but at most54 setae in C. acutovulvata.Material examined: 8 males and 7 females from fresh materialcollected in Thailand.Type host: Anorrhinus galeritus carinatus (Blyth, 1845).Type material: Holotype male and allotype female from Chong,Muang, Trang, Thailand, Mar. 4, 1903, collected by Wichit SuwanLaong, USNM. Paratypes: 6 males and 3 females with same data;1 male and 1 female from Lamo, Muang, Trang, Thailand, Mar. 3,1903, collected by Wichit Suwan Laong, USNM; 2 females fromNa Wong, Muang, Phatthalung, Thailand, Mar. 0, 1963, collectedby Wichit Suwan Laong, USNM.Chapinia waniti is named for Mr. Wanit Songprakob, Songkhla,Thailand, in appreciation for mounting Mallophaga and for directingthe activities of the field collector, Wichit Suwan Laong. After mydeparture from Thailand in April 1963, both boys collected for theBemice P. Bishop Museum.Chapinia acutovulvatM (Piaget)Figures 9, 10, 35, 36Menopon aculovulvatum Piaget, 1881, p. 5, pi. 1 (fig. 4). [Type host: Bucerosmalaharicus= Anthracoceros a. albirostris (Shaw, 1808).]Menopon aculovulvatum Piaget, 188.5, p. 106, pi. 11 (fig. 8).Allomenopon rnjobergi Eichler, 1947, pp. 2, 20, figs. 1, 2 (new synonym). [Typehost: Anthracoceros convexus (Temminck, 1831).]Chapinia rnjobergi (Eichler)?Hopkins and Clay, 1952, p. 68.Chapinia acutovulvata (Piaget)?Hopkins and Clay, 1952, p. 67.Dr. Eichler's specimens of Chapinia rnjobergi from Anthracocerosconvexus in the Zoological Museum, Humboldt University, Berlin,were loaned through the courtesy of Dr. von Keler. Comparisonof these lice with specimens of Chapinia acutovulvata from the typehost discloses no morphological differences between the two series.A lectotype male for Chapinia acutovulvata was designated by Clay(1949a) from the Piaget collection, now in BMNH, BM 777, with6 syntype females, BM 774 and 776.
24 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 120Both sexes are smaller than corresponding sexes of Chapiniatraylori in all measurements except length and width of head (tables1,2).Male: Venter of third femora and posterolateral margins of abdom-inal sternites IV-VI each with large thick brushes of normal seta.Each lateral margin of abdominal tergites II-VI with a short setaebetween the spiracle and postspiracular seta. Abdominal sternite IIwith 42-54 total setae and one median row of setae on posterior margin.Terminal abdominal segments as shown in figure 10. Genitalia asshown in figure 9, endomeres with small inner plate and paired outerrims.Female: Kesembles the male except that abdominal sternite IIhas 76-86 total setae. Terminal abdominal tergite with 16-22 longand 10-12 short setae on posterior margin; abdominal sternite VIIIwith 18-30 setae on posterior margin (fig. 35). Anal fringe with44-54 setae (fig. 36).Discussion: Although largerm size, Chapinia acutovulvata resemblesmost closely C. clayae. The venter of the third femora and postero-lateral margins of abdominal sternites IV-VI each have large thickbrushes of normal setae in C. acutovulvata but small scattered brushesof normal setae in C. clayae. Abdominal sternite II of both sexesof C. acutovulvata has fewer total setae than in corresponding sexes ofC. waniti but has more total setae than in corresponding sexes ofC. malayensis. The male genitalia have endomeres possessing oneach outer rim a triangular internal knob that is longer than wide inC. acutovulvata but wider than long in C. clayae. These internal knobsare absent in other Chapinia. The female terminal abdominal ter-gite has on the posterior margin at least 26 setae in C. acutovulvata,of which approximately two-thirds are long and one-third are short,but at most 24 setae in C. clayae and C. malayensis, of which approx-imately one-half are long and one-half are short in C. clayae, butapproximately two-thirds are long and one-third are short in C.malayensis. In addition this margin has on each side of midline atleast six long setae in C. acutovulvata but at most five long setae inC. clayae. The female abdominal sternite VIII has on the posteriormargin fewer than 32 setae in C. acutovulvata but more than 34 setaein C. malayensis. Each lateral projection of the ventral scleritebetween female vulva and anus has fewer than eight thick, posteriorlydirected setae in C. acutovulvata, but more than eight m C. waniti(fig. 72 sp.). The female anal fringe has 44-54 setae in C. acuto-vulvata, at most 44 setae in C. clayae, and at least 56 setae in C. waniti.Material examined: 33 males and 35 females from fresh and driedmaterial collected in the Oriental region; from the type host: 2females, Piaget, BMNH 1953-21; 2 males and 2 females from Nepal,
NO. 3558 MALLOPHAGA?ELBEL 25December 1935, Meinertzhagen 4872, BMNH; from Anthracocerosalbirostris leucogaster (Blyth, 1841): 2 males and 2 females fromMyitkyina, Burma, Mar. 26, 1945, collected by the U.S. Typhus Com-mission, BMNH; 3 males, 1 female with same data, USNM; 2 malesand 1 female from StiUwell Road, Myitkyina, Burma, Sept. 26, 1945,collected by H. S. Fuller, BMNH 1947-321 (164); 2 males, 2 femaleswith same data, USNM; 2 males from Hin Laem, Tha Khanun,Kanchanabm-i, Thailand, Nov. 27, 1952, collected by Robert E. Elbeland H. G. Deignan, USNM; 2 males from Ban Khlua Klang, PrachuapKhiri Khan, Thailand, December 1952, collected by Robert E. Elbeland H. G. Deignan, USNM ; 2 males and 1 female from Ban Nam Phu,Phu Khieo, Chaiyaphum, Thailand, Dec. 22, 1952, collected by RobertE. Elbel, USNM; 1 male and 1 female from Ban Thung Chuak, SalokBat, Khanu, Kamphaeng Phet, Thailand, June 25, 1953, collected byRobert E. Elbel, USNM; 1 female from Tha Dm Daeng, Pa Bon, PakPha Yun, Phatthalung, Thailand, July 30, 1962, collected by WichitSuwan Laong, USNM; 6 males and 6 females from Muang Kluang,Kapoe, Ranong, Thailand, 1962-1963, collected by Wichit SuwanLaong, USNM; 6 males and 11 females from Pa Dong Lan, Chumphae,Khon Kaen, Thailand, Dec. 2, 1962, collected by Kitti Thonglongya,SMRL; from Anthracoceros coronatus (Boddaert, 1783): 2 femalesfrom CNHM skins from Kanha, Central Provinces, India, August1946, collected by Rup Chand, REE; 2 females from CNHM skinsfrom Nawadeh, Bihas, India, Nov. 11, 1947, collected by W. Koelz,REE; 1 male from USNM skins from India and Ceylon, 1874 and1944, collected by B. H. Swales and S. D. Ripley, REE; from Anthra-coceros convexus (Temminck, 1831): 1 male and 1 female from lot 1584(TMRN), Zoological Museum, Humboldt University, Berlin; accord-ing to Eichler (1947) the Mallophaga WEC 2268 from this later namedhost were collected in Sumatra by E. Mjoberg; from Anthracocerosmarchei Oustalet, 1885: 1 male from Puerto Princesa, Palawan,Philippines, May 12, 1962, collected by Max Thompson, USNMBPM-PI 2313.Drawings were made of a male and a female from Anthracocerosalbirostris leucogaster collected in Myitkyina, Burma. Specimens inBMNH. Chapinia malayensisy new speciesFigures 12, 37Male: Smaller than Chapinia traylori in all measurements exceptwidth of abdomen (table 1). Venter of third femora and postero-lateral margins of abdominal sternites IV-VI each with large thickbrushes of normal setae which are not as numerous on abdominalsternite VI. Each lateral margin of abdominal tergites II-VI witha short seta between the spiracle and postspiracular seta. Abdominal
26 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 120
sternite II with 34 total setae and one median row of setae on posteriormargin. Terminal abdominal segments similar to those of C. acuto-vulvata. Genitalia as shown in figure 12, endomeres with large innerplate and paired outer rims.Female: Slightly smaller than Chapinia traylori in all measurementsexcept length of head (table 2), Resembles the male except thatabdominal sternite II has 64 total setae. Terminal abdominal tergitewith 20 long and 4 short setae on posterior margin; abdominal sterniteVIII with 24 long and 12 short setae on posterior margin (fig. 37).Anal fringe similar to that of C. acutovulvata with 50 setae.Discussion: Chapinia malayensis resembles most closely C. acuto-vulvata. Abdominal sternite II of both sexes has fewer total setae inC. malayensis than in corresponding sexes of C. acutovulvata. Malegenitalia have endomeres possessing a large inner plate with pairedlateral flanges and a central terminal point in C. malayensis but asmall inner plate in C. acutovulvata. The female terminal abdominaltergite has on the posterior margin at most 24 setae in C. malayensisbut at least 26 setae in C. acutovulvata. The female abdominalsternite VIII has on the posterior margin more than 34 setae in C.malayensis but fewer than 32 setae in C. acutovulvata.Material examined: 1 male and 1 female from fresh materialcollected in Borneo.Type host: Anthracoceros malayanus (Raffles, 1822).Type material: Holotype male and allotype female from Borneo,Aleinertzhagen 10910, BMNH.Chapinia hoplai, new speciesFigures 13, 38Both sexes are slightly smaller than corresponding sexes of Chapiniatraylori in all measurements except length of head (tables 1,2).Male: Venter of third femora and posterolateral margins of ab-dominal sternites IV-VI each with large thick brushes of normalsetae. Each lateral margin of abdominal tergites II-VI without ashort seta between the spiracle and postspiracular seta. Abdominalsternite II with 66-68 total setae and one median row of setae onposterior margin. Terminal abdominal segments similar to those ofC. acutovulvata. Genitalia as shown in figure 13, endomeres withpaired outer rims only.Female: Resembles the male except that abdominal sternite II has80-88 total setae. Terminal abdominal tergite with 14 long and 12short setae on posterior margin; abdominal sternite VIII with 16long and 4 short setae on posterior margin (fig. 38). Anal fringesimilar to that of C. acutovulvata with 50-54 setae.Discussion: Although slightly larger in size, Chapinia hoplai re-
NO. 3558 MALLOPHAGA?ELBEL 27sembles most closely C. hoonsongi. The venter of the third femoraand posterolateral margins of abdominal sternite IV-VI each havelarge thick brushes of normal setae in C. hoplai but small scatteredbrushes of normal setae in C hoonsongi. Abdominal sternite II ofboth sexes has more total setae in C. hoplai than in correspondingsexes of C. hoonsongi. The male genitalia have endomeres with pairedouter rims that are curved inwardly in C. hoplai but straight andnearly parallel in C. hoonsongi. The female terminal abdominaltergite has on the posterior margin more than 24 setae of whichapproximately one-half are long and one-half are short in C. hoplaibut fewer than 22 setae of which approximately two-thirds are longand one-third are short in C. hoonsongi.Material examined: 3 males and 2 females from dried materialcollected in the Philippines.Type host: Anthracoceros montani (Oustalet, 1880).Type material: Holotype male and allotype female from USNMskins from Sulu, and Tawitawi, Philippines, 1891, collected byD. C. Worchester and F. S. Bourns, EEE in USNM. Paratypes:2 males and 1 female with same data.Chapinia hoplai is named for Dr. Cluff E. Hopla, Department ofZoology, University of Oklahoma, in appreciation for his thoughtfuladvice while directing this study.Chapinia hoonsongi, new speciesFigures 11, 14, 39Both sexes are smaller than corresponding sexes of Chapinia tray-lori in all measurements except length of head (tables 1,2).Male: Venter of third femora and posterolateral margins of ab-dominal sternites IV-VI each with small scattered brushes of normalsetae which are not as numerous on abdominal sternite VI. Eachlateral margin of abdominal tergites II-VI without a short setabetween the spiracle and postspiracular seta. Abdominal sterniteII with 48-62 total setae and one median row of setae on posteriormargin. Terminal abdominal segments as shown in figure 11.Genitalia as shown in figure 14, endomeres with paired outer rimsonly.Female: Resembles the male except that abdominal sternite IIhas 62-70 total setae. Terminal abdominal tergite with 14-16 longand 4 short setae on posterior margin; abdominal sternite VIII with18-20 setae on posterior margin (fig. 39). Anal fringe similar to thatof C. acutovulvata with 50-56 setae.Discussion: Although slightly smaller in size, Chapinia hoonsongiresembles most closely C. hoplai. The venter of the third femoraand posterolateral margins of abdominal sternites IV-VI each have
28 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 120
small scattered brushes of normal setae in C. hoonsongi but large thickbrushes of normal setae in C. hoplai. Abdominal sternite II of bothsexes has fewer total setae in C hoonsongi than in corresponding sexesof C. hoplai. The male genitalia have endomeres with paired outerrims that are straight and nearly parallel in C. hoonsongi but curvedinwardly in C. hoplai. The female terminal abdominal tergite hason the posterior margin fewer than 22 setae of which approximatelytwo-thirds are long and one-third are short in C. hoonsongi but morethan 24 setae of which approximately one-half are long and one-halfare short in C hoplai.Material examined: 5 males and 8 females from fresh and driedmaterial collected in Thailand.Type host: Rhyticeros undulatus ticehursti Deignan, 1941.Type material: Holotype male from USNM skin from Ban HaiHuai, Thailand, June 15, 1936, collected by H. G. Deignan, EEEin USNM. Additional types from Rhyticeros u. undulatus (Shaw,1811): Allotype female from Khao Phap Pha Mt., Ban Na, Muang,Phatthalung, Thailand, Feb. 7, 1955, collected by Boonsong Lekagul,USNM. Paratypes: 3 males and 3 females with same data; 1 malefrom BL skin from Nong Ko, Siracha, Chon Buri, Thailand, August1953, collected by Boonsong Lekagul, REE; 4 females from Lamo,Muang, Trang, Thailand, Mar. 5, 1963, collected by Wichit SuwanLaong, USNM.Chapinia hoonsongi is named for Dr. Boonsong Lekagul, Bangkokphysician and naturalist, in appreciation for the fresh material hecollected from Thailand and for permission to examine his hornbillskins for MaUophaga.Chapinia wenzeli, new speciesFigures 15, 40, 41Both sexes are smaller than corresponding sexes of Chapinia tray-lori in all measurements except length of head in females (tables 1, 2).Male: Venter of third femora and posterolateral margins of abdom-inal sternites IV-VI each with small scattered brushes of normalsetae which are not as numerous on abdominal sternite VI. Eachlateral margin of abdominal tergites II-VI without a short setabetween the spiracle and postspiracular seta. Abdominal sterniteII with 30-36 total setae and one median row of setae on posteriormargin. Terminal abdominal segments similar to those of C. hoon-songi. Genitalia as shown in figure 15, endomeres with paired outerrims only, the posterolateral margins of which are split.Female: Resembles the male except that abdominal sternite IIhas 50-60 total setae. Terminal abdominal tergite with 12-14 longand 6-8 short setae on posterior margin; abdominal sternite VIII
NO. 3558 MALLOPHAGA?ELBEL 29
with 18-24 setae on posterior margin (fig. 40). Anal fringe with40-44 setae (fig. 41).Discussion: Chapinia wenzeli resembles most closely C. blakei.The venter of the third femora and posterolateral margins of abdom-inal sternites IV-VI each have small scattered brushes of normalsetae in both sexes of C. wenzeli but only in the female of C. blakei.The male genitalia have endomeres with paired outer rims in C.wenzeli but paired plates in C. blakei. However, the paired platesmay be split in C. blakei, giving the appearance of paired outer rimsand inner plate, but the inner plate in this case is unsymmetrical.The female abdominal sternite VIII has on the posterior margin atleast 18 setae in C. wenzeli but at most 18 setae in C. blakei, and theinternal sclerite, absent in C. wenzeli, is present in C. blakei. Thefemale anal fringe has at least 40 setae in C. wenzeli and at most 40setae in C. blakei.Material examined : 20 males and 23 females from fresh and driedmaterial collected in the Philippines.Type host: Penelopides panini samarensis Steere, 1890.Type material: Holotype male and allotype female from CNHMskins from Sandayong, Sierra BuUones, Bohol Island, Philippines,April 1955, collected by D. S. Kabor, REE in CNHM. Paratypes:2 females with same data; 2 males from CNHM skins from Matu-guinao, Samar Island, Philippines, April 1957, collected by D. S.Rabor, REE; from Penelopides panini manilloe (Boddaert, 1783):1 male and 1 female from CNHM skin from Bataan, Luzon Island,Philippines, Jan. 17, 1905, collected by Celestino and Canton, REE;from Penelopides panini mindorensis Steere, 1890: 1 male fromCNHM skin from Balete, Rio Baca, Mindanao, Philippines, Apr. 1,1905, collected by McGregor, Celestino, and Canton, REE; fromPenelopides panini affinis Tweeddale, 1877: 8 males and 2 femalesfrom CNHM skins from Mindanao, Philippines, 1946 and 1947,collected by Werner and Alcasid, REE; 3 males and 8 females fromDavao, Mindanao, Philippines, Jan. 18, 1947, KCE; 4 males and9 females from Mindanao, Phihppines.Chapinia wenzeli is named for Dr. Rupert L. Wenzel, Curator ofInsects, Chicago Natural History Museum, in appreciation for theloan of hornbill Menoponidae from that museum.Chapinia blakei^ new speciesFigures 16, 42, 43Male: Slightly smaller than Chapinia traylori in all measurementsexcept length of head and width of metathorax (table 1). Venterof third femora and abdominal sternites IV-VI each without brushes.Each lateral margin of abdominal tergites II-VI without a short
30 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 120
seta between the spiracle and postspiracular seta. Abdominalsternite II with 32-42 total setae and one median row of setae onposterior margin. Terminal abdominal segments similar to thoseof C. boonsongi. Genitalia as shown in figure 16, endomeres withpaired plates.Female: Smaller than Chapinia traylori in all measurements exceptlength of head (table 2). Resembles the male except that the venterof the third femora and posterolateral margins of abdominal sternitesIV-VI each have small scattered brushes of normal setae which arenot as numerous on abdominal sternite VI. Abdominal sternite IIwith 52-64 total setae. Terminal abdominal tergite with 10-16long and 8-14 short setae on posterior margin; abdominal sterniteVIII with 14-18 setae on posterior margin (fig. 43) and with internalsclerite having slender posteriorly divergent margins. Anal fringewith 34-40 setae (fig. 42).Discussion : Chapinia blakei resembles most closely C. wenzeli. Theventer of the third femora and posterolateral margins of abdominalsternites IV-VI each have small scattered brushes of normal setae infemales of C. blakei and in both sexes of C. wenzeli. The male genitaliahave endomeres with paired plates in C. blakei but paired outer rimsin C. wenzeli. However, the paired plates may be split in C. blakei,giving the appearance of paired outer rims and inner plate, but theinner plate in this case is unsymmetrical. The male genitalia ofC. traylori also have endomeres with paired plates, but the posteriorextension of the endomeres is split in C. traylori and not split inC. blakei. The female abdominal sternite VIII has on the posteriormargin at most 18 setae in C. blakei but at least 18 setae in C. wenzeli,and the internal sclerite with slender posteriorly divergent margins,present in G. blakei, is absent in both C. wenzeli and C. traylori. Thefemale anal fringe has at most 40 setae in C. blakei and at least 40setae in C. wenzeli.Material examined: 13 males and 16 females from fresh and driedmaterial collected in the Philippines.Type host: Rhyticeros I. leucocephalus (Vieillot, 1816).Type material: Holotype male and allotype female from CNHMskins from Zamboanga, Mindanao Island, Philippines, 1948 and 1956,collected by D. S. Rabor, REE in CNHM. Paratypes: 11 males and12 females from Mutya, Canon, Mindanao Island, Philippines,December 1961, collected by Rabor and Gonzales, BPBM; 1 femalefrom Davao, Tagum, Mindanao Island, Philippines, Oct. 13, 1946,collected by H. Hoogstraal, CNHM; from Rhyticeros leucocephaluswaldeni (Sharpe, 1877) : 1 male and 2 females from CNHM skins fromTolong, Negros Island, Philippines, November-December 1948, col-lected by D. S. Rabor, REE.
NO. 3558 MALLOPHAGA?ELBEL 31Chapinia blakei is named for Dr. Emmet R. Blake, Curator of Birds,Chicago Natural History Museum, in appreciation for permission toexamine hornbill skins for Mallophaga in that museum.Chapinia traylori, new speciesFigures 17, 25, 26, 44, 45Male: As illustrated in figure 26. Slightly larger than Chapiniaclayae, C. acutovulvata, C. malayensis, C. hoplai, C. boonsongi, C.wenzeli, C. blakei, and C. lydae in all measurements except length ofhead in C. clayae, C. acutovulvata, C. hoplai, C. boonsongi, C. blakei,and C. lydae, width of head in C. acutovulvata, width of metathorax inC. blakei, and width of abdomen in C malayensis; smaller than C.waniti in all measurements except width of prothorax and width ofmetathorax; approximately the same size as C. robusta (table 1).Venter of third femora and abdominal sternites IV-VI each withoutbrushes. Each lateral margin of abdominal tergites II-VI without ashort seta between the spiracle and postspiracular seta. Abdominalsternite II with 24-42 total setae and two median rows of setaealthough the anterior row has widely separated setae. Terminalabdominal segments as shown in figure 26e. Genitalia as shown infigure 17, endomeres with paired plates.Female: As illustrated in figure 25. Larger than Chapinia wanitiand Chapinia acutovulvata in all measurements except length and widthof head; slightly larger than other Chapinia in all measurements exceptlength of head (table 2). Resembles the male except that abdominalsternite II has 38-58 total setae. Terminal abdominal tergite with10 long and 6-10 short setae on posterior margin; abdominal sterniteVIII with 14-20 setae on posterior margin and with small triangularinternal sclerite (fig. 44) ; each lateral projection of the ventral scleritebetween vulva and anus with 2 or 3 thick, posteriorly directed setae(fig. 44sp). Anal fringe with 34-40 setae (fig, 45).Discussion: Chapinia traylori resembles most closely C. blakei.The venter of the third femora and posterolateral margins of abdomi-nal sternites IV-VI lack brushes in both sexes of C. traylori and in themale of C. blakei, but these margins have small scattered brushes ofnormal setae in females of C. blakei. The male genitalia have endo-meres with paired plates in both C. traylori and C. blakei, but theposterior extension of the endomeres is split in C. traylori and notsplit in C. blakei; the paired plates may be spht in C. blakei, givingthe appearance of paired outer rims and inner plate, but the innerplate in this case is unsymmetrical. The female abdominal sterniteVIII has a small triangular internal sclerite in C. traylori but a muchlarger sclerite with slender posteriorly divergent margins in C. blakei.Each lateral projection of the ventral sclerite between female vulva
32 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 120and anus has fewer than four thick, posteriorly directed setae in C.traylori but more than four in all other Chapinia.Material examined: 73 males and 52 females from fresh and driedmaterial collected in the Philippines.Type host: Buceros hydrocorax semigaleatus Tweeddale, 1878.Type material: Holotype male, allotype female, and paratypefemale on sameshde from CNHM skins from San Isidro, Samar Island,Philippines, April-May 1957, collected by D. S. Rabor, REE inCNHM. The allotype female is the largest female and is next tothe male. Paratypes: 1 male and 1 female from CNHM skinsfrom Cantaub, Sierra Bullones, Bohol Island, Philippines, April-May1955, collected by D. S. Rabor, REE; 2 males from CNHM skinsfrom Matuguinao, Samar Island, Phihppines, April 1957, collectedby D. S. Rabor, REE; 15 males and 9 females from CNHM skinsfrom Mount Capato-an, Samar Island, Philippines, May 1957,collected by D. S. Rabor, REE; from Buceros hydrocorax mindanensisTweeddale, 1877: 11 males and 12 females from Mutya, Canon,Mindanao Island, Phihppines, Dec. 23, 1961, collected by Rabor andGonzales, BPBM; 9 males and 7 females from Mount McKinley,Davao, Mindanao Island, Philippines, August 1946, collected by H.Hoogstraal, CNHM; 2 males and 3 females from CNHM skins fromTaglawig, Tagum, Davao, Mindanao Island, Philippines, October1946, collected by Celestino, REE; 13 males and 7 females from MountApo, Todaya, Mindanao Island, Philippines, Oct. 25, 1946, collectedby Hoogstraal and Hey, CNHM; 2 males and 1 female from CNHMskins from Kidapawan, Cotabata, Mindanao Island, Philippines,Dec. 2, 1946, collected by Alcasid, REE; 1 male and 1 female fromCNHM skins from Burungkot Upi, Cotabata, Mindanao Island,January 1947, collected by Werner and Alcasid, REE; 9 males and 4females from CNHM skins from Mount Malindang, Zamboanga,Mindanao Island, Philippines, March-May 1956, collected by D. S.Rabor, REE; 2 males and 1 female from USNM skins from MindanaoIsland, Philippines, Augustr-September 1903, collected by E. A.Mearns, REE; 5 males and 4 females from Mindanao Island, Philip-pines.Chapinia traylori is named for Dr. M. A. Traylor, Division ofBirds, Chicago Natural History Museum, in appreciation for assistancein examination of hornbill skins for Mallophaga in that museum.Chapinia lydae, new speciesFigures 20, 46
"Colpocephalum hirtum Rudow, 1866."?Piaget, 1880, p. 530, pi. 44 (fig. 3).[Not Rudow, 1866; type host: Buceros cassidix= Rhyticeros cassidix (Tem-minck, 1823).]Clay (1951b) stated that it was impossible to say whether or not
NO. 3558 MALLOPHAGA?ELBEL 33Piaget's specimens of a headless female and 2 nymphs from Buceroscassidix?Rhyticeros cassidix (Temminck, 1823) were conspecific withRudow's hirtum from Buceros ruJicollis=Rhyticeros plicatus ruficollis(Vieillot, 1816). Through the courtesy of Dr. Clay, Piaget's speci-mens have been examined, and they do not appear to be conspecificwith Chapinia hirta (Rudow, 1866). They are described herewithas part of the type material from Rhyticeros cassidix.Both sexes are smaller than corresponding sexes of Chapinia tray-lori in all measurements except length of head, but this measurementin the male is larger than in males of other Chapinia (tables 1,2).Male: Venter of third femora and posterolateral margins of ab-dominal sternites IV-VI each with small scattered brushes of normalsetae which are not as numerous on abdominal sternite VI. Eachlateral margin of abdominal tergites II-VI without a short setabetween the spiracle and postspkacular seta. Abdominal sternite IIwith 62-64 total setae and three median rows of setae. Terminalabdominal segments similar to those of C. boonsongi. Genitalia asshown in figure 20, endomeres with inner plate and paired outer rims.Female: Resembles the male except that abdominal sternite II has54-68 total setae. Terminal abdominal tergite with 14 long and 8short setae on posterior margin; abdominal sternite VIII with 20-24setae on posterior margin (fig. 46). Anal fringe similar to that of Chirta, with 46-50 setae.Discussion: Chapinia lydae superficially resembles C. hirta. Ab-dominal sternite II in both species has approximately the same numberof total setae in females as in males and three median rows of setae.Abdominal sternite II of other members of the acutovulvata species-group has more setae in the females than in the males and 1 or 2 medianrows of setae. The male genitalia in C. lydae are wider than in C.hirta, and the parameres are enlarged anteriorly only in C lydae; theendormeres have an inner plate and paired outer rims in C. lydae, butendomeres apparently are absent in C. hirta. The female terminalabdominal tergite has on the posterior margin in C. lydae 3 or 4 longsetae on each side of the midline, the two median setae being as widelyspaced as four times the distance between each of the 3 or 4 long setae,but in C. hirta 5 long setae on each side of the midhne yield a total of10 setae that are evenly spaced.Material examined: 3 males and 5 females from dried skins col-lected in the Celebes.Type material: Holotype male and allotype female from USNMskins from Palaleh River, Celebes, Aug. 9, 1914, collected by H. C.Raven, LE in USNM. Paratypes : 2 males and 3 females with samedata; 1 female, Piaget, BMNH, 1928-325.Chapinia lydae is named for my wife, Lyda, in appreciation for the221-522?67 3
34 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 120dried material that she obtained froni Rhyticeros cassidix and otherhosts in the USNM and for much help in preparation of the manu-script. The hirta Species-GroupSpecies similar in shape to Chapinia traylori (figs. 25, 26). Differ-ing from other species-groups in the following combination of charac-ters: Dorsal-lateral margins of head with a preocular notch; venterof third femora and posterolateral margins of abdominal sternitesIV-VI each with small scattered brushes of normal setae which arenot as numerous on abdominal sternite VI; each lateral margin ofabdominal tergites II-VI without a short seta between the spiracleand postspiracular seta; females with approximately the same numberof abdominal sternal setae as males; abdominal sternite II withthree median rows of setae; male genitalia much narrower than forother species-groups, without lateral horns on each side of endomeres,and with parameres not enlarged anteriorly but split posteriorly;females without sclerital hooks on each side of midline of ventralsclerite between vulva and anus; female abdominal sternite VIIIwith most of setae much shorter than those on posterior margin.Hosts: Species of the hirta species-group have been found only onthe genera Penelopides and Rhyticeros of the avian family Bucerotidae.Chapinia muesebecki, new speciesF1GUKE8 18, 21Male: Smaller than Chapinia traylori in all measurements exceptlength of head (table 1). Abdominal sternite II with 58-62 totalsetae. Terminal abdominal segments as shown in figure 21. Gen-italia as shown in figure 18.Female: Unknown.Discussion: Chapinia muesebecki resembles most closely C. hirta.Male terminal abdominal segments are shorter in C. muesebecki thanin C. hirta, and the partial division between abdominal sternitesVII and VIII is not as pronounced in C. muesebecki as in C. hirta.The sclerite of male genital sac is nearly twice as long as wide inC. muesebecki but approximately as wide as long in C. hirta, and thissclerite is approximately three times as long in C. muesebecki as inC. hirta.Material examined: 2 males from dried skins collected in theCelebes.Type host: Penelopides e. exarhatus (Temminck, 1823).Type material: Holotype male from USNM skins from Celebes,1914-1916, collected by H. C. Raven, REE in USNM. Paratypemale with same data.
NO. 3558 MALLOPHAGA?ELBEL 35Chapinia muesebecki is named for Mr. C. F. W. Muesebeck, Divisionof Insects, U.S. National Museum, in appreciation for the loan ofMallophaga from that museum.Chapinia hirta (Rudow)Figures 19, 22, 47, 48Colpocephalum hirtum Rudow, 1866, p. 474. [Type host: Buceros ruficollis= Rhyticeros plicatus ruficollis (Vieillot, 1816).]Colpocephalum hirtum Rudow, 1869, p. 399.Chapinia hirta (Rudow).?Hopkins and Clay, 1952, p. 67.Hopkins and Clay state that the generic position of hirtum is doubt-ful. Examination of specimens from the type host shows them to beChapinia. Therefore, the male, BM 13376, is designated hereby asneotype of C. hirta. The slide has been so labeled.Both sexes are smaller than corresponding sexes of Chapinia trayloriin all measurements except length of head (tables 1,2).Male: Abdominal sternite II with 60-64 total setae. Terminalabdominal segments as shown in figure 22. Genitalia as shown infigure 19.Female: Resembles the male except that terminal abdominal seg-ments have a tergite with 12 long and 10 short setae on posteriormargin; abdominal sternite VIII has 18-22 setae on posterior margin(fig. 48). Anal fringe with 46^8 setae (fig. 47).Discussion: Chapinia hirta resembles most closely C. muesebecki.Also, C. hirta superficially resembles C. lydae. Abdominal sternite IIin these three species has approximately the same number of totalsetae in females as in males, and three median rows of setae. Maleterminal abdominal segments are longer in C. hirta than in C. muese-becki, and the partial division between abdominal sternites VII andVIII is more pronounced in C. hirta than in C. muesebecki. Thesclerite of male genital sac is approximately as wide as long in C. hirtabut nearly twice as long as wide in C. muesebecki, and this sclerite isapproximately one-third as long in C. hirta as in C. muesebecki. Themale genitalia in C. hirta are narrower than in C. lydae; the parameres,straight-sided in C. hirta, are enlarged anteriorly in C. lydae; theendomeres apparently are absent in C. hirta, but have an inner plateand paired outer rims in C. lydae. The female terminal abdominaltergite has on the posterior margin in C. hirta 5 long setae on eachside of the midline, these 10 setae being evenly spaced; however, inC lydae 3 or 4 long setae on each side of the midline have the twomedian setae as widely spaced as four times the distance betweeneach of the 3 or 4 long setae.Material examined: 3 males and 2 females from fresh and driedmaterial collected in the Oriental and Australasian regions; neotypemale, BM 13376, and female, BM 13375, from New Guinea, BMNH;221-522?67 4
36 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 120from Rhyticeros plicatus suhruficollis (Blyth, 1843) : 1 female fromUSNIM skins from Domellsland, Mergui Archipelago, 1904, collectedby W. L. Abbott, REE; from Rhyticeros plicatus mendanae (Hartert,1924) : 1 male from CNHM skins from Guadalcanal, Solomon Islands,August-October 1944, collected by W. J. Beecher, REE; 1 male fromMMZ skin from Guadalcanal, Solomon Islands, Jan. 20, 1944, collectedby K. W. Prescott, REE.Drawings were made of the neotype male and the female, BM 13375.Specimens in BMNH.Bucerocolpocephalum, new genusFigures 64, 65Head triangular, width 1% to 1% times that of length. Foreheadshghtly narrower anteriorly. Temples expanded. Antennae 4-jointed, third segTaent constricted at base, and terminal segmentcapitate with definite signs of division into two parts, either by trans-verse line or marginal indentation. Antennary fossa deep, coveredabove by expansion of lateral margin of head, posterior margin ofwhich lacks an eye. Dorsal-lateral margin of forehead above anten-nary fossa with shallow notch. Gular region narrow with a ridge oneach lateral margin from which 8-11 setae extend. Pronotum ex-panded anteriorly with posterior marginal row of long setae. Meta-notum expanded posteriorly with posterior marginal row of long setaeand 4-7 short setae on each lateral margin. Metanotum separatedfrom mesonotum and from pleurites. The sclerotized median buttonbehind the prothorax (fig. 25m) is a vestige of the mesonotum; thesupposed mesonotum, the narrow sclerotized band posterior tothis button, is a mere extension of the subcoxae (Cope, 1941).Thoracic sternal plates as shown in figures 64b and 64c. Metasternalplate oval with 14-24 setae. Venter of third femora and postero-lateral margins of abdominal sternite IV each with combs of setae.Abdominal segments consist of tergites, sternites, and pleurites, thelatter without prolongation of posteroventral angles. Abdominaltergites each with a posterior marginal row of setae, the most lateradbeing the postspiracular seta. Each lateral margin of abdominaltergites II-VIII with 1-4 short setae between the spu-acle and post-spiracular seta. Sternites and pleurites each with a posterior mar-ginal row of long and short setae and with numerous short, usuallythick setae. Male terminal abdominal sternites VIII and IX fusedwith complete division from sternite VII (fig. 65). Male genitaliaas illustrated for each species with parameres anteriorly either en-larged or curved inwardly. Female terminal abdominal segments asillustrated for each species, with lateral processes arising from ventral
NO. 3558 MALLOPHAGA?ELBEL 37
sclerite between vulva and anus, with long stout setae and strongspines. Females similar to males in size, general shape, and chaeto-taxy except for terminal abdominal segTnents.Bucerocolpocephalum resembles most closely Bucerophagus (figs.66-69) but dilTers in several characters: The posterior margin ofthe expansion of the lateral margin of the head covering the antennaryfossa lacks an eye in Bucerocolpocephalum, but an eye with a doublecornea is present in Bucerophagus. The gular region has on eachlateral ridge 8-11 setae in Bucerocolpocephalum, but the ridge isabsent, and each lateral margin has at most eight setae in Bucero-phagus. The matasternal plate is oval in Bucerocolpocephalum buttrapezoidal or triangular in Bucerophagus. The venter of the thirdfemora has combs of setae in Bucerocolpocephalum but may havelarge thick brushes of normal setae in Bucerophagus; similar combsare present on posterolateral margins of abdominal sternite IV inBucerocolpocephalum, and similar brushes are present on postero-lateral margins of abdominal sternites IV and V in Bucerophagus.Male genitalia of Bucerocolpocephalum are shorter than in Buceropha-gus. The female anal fringe is weak in Bucerocolpocephalum andprominent in Bucerophagus.The male genitalia and details of the male and female terminalabdominal segments are the best characters for separating speciesof Bucerocolpocephalum.Hosts: Species of Bucerocolpocephalum have been found only onthe genera Ptilolaemus and Anorrhinus of the avian family Bucero-tidae. [Genotype: Bucerocolpocephalum emersoni, new species.]Bucerocolpoceplialum emersoni, new speciesFigures 49, 57, 64, 65Both sexes are approximately the same size as corresponding sexesof Bucerocolpocephalum deignani (table 3).Male: As illustrated in figure 65. Metasternal plate with 16-20setae. Abdominal sternite II with 40-48 total setae. Terminalabdominal segments as shown in figure 65e. Genitalia as shown infigure 57.Female: As illustrated in figure 64. Kesembles the male exceptthat metasternal plate has 16-24 setae. Terminal abdominal tergitewith 30-36 setae on posterior margin; abdominal sternite VIII with32-42 setae on posterior margin and with internal triangular sclerite;anal fringe with 44-48 weak setae (fig. 49).Discussion: Bucerocolpocephalum emersoni resembles most closelyB. deignani. Male terminal abdominal sternites VIII and IX havea lateral notch in B. emersoni but not in B. deignani. Male genitaliahave parameres anteriorly slender, curved inwardly with each lateral
38 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 120point reaching endomeres in B. emersoni and anteriorly enlarged, notcurved inwardly, in B. deignani; endomeres have a pair of posteriorpoints in B. emersoni which are absent in B. deignani. The femaleterminal abdominal tergite has thick setae along the entire posteriormargin in B. emersoni, but thick setae are absent medially in B.deignani. The female abdominal sternite VIII has an internal tri-angular sclerite in B. emersoni that is absent in B. deignani.Material examined: 13 males and 19 females from fresh and driedmaterial collected in the Oriental region.Type host: Ptilolaemus tickelli austeni (Jerdon, 1872).Type material: Holotype male and allotype female from Phu LornLo Mt., Kok Sathon, Dan Sai, Loei, Thailand, Mar. 23, 1954, col-lected by Robert E. Elbel, USNM. Paratypes: 8 males and 15females with same data; from Ptilolaemus tickelli indochinensisDelacour and JabouiUe, 1928: 4 males and 3 females from CNHMskins from Muong Yo, Laos, and Muong Maun, Tonkin, Indochina,March-May 1929, collected by Van Tyne, REE.Bucerocolpocephalum emersoni is named for Dr. K. C. Emerson,Stillwater, Okla., in appreciation for his untiring help and advicethroughout this study, particularly in the preparation of the manu-script and illustrations, and in other studies on Oriental MaUophaga.Bucerocolpocephalum deignani, new speciesFigures 50, 58, 59Both sexes are approximately the same size as corresponding sexesof Bucerocolpocephalum emersoni (table 3).Male: Metasternal plate with 14-18 setae. Abdominal sternite IIwith 44-46 total setae. Terminal abdominal segments as shown infigure 59. Genitalia as shown in figure 58.Female: Resembles the male except that abdominal sternite II has42-58 total setae. Terminal abdominal tergite with 28-34 setae onposterior margin; abdominal sternite VIII with 34-42 setae on poste-rior margin; anal fringe with 24-40 weak setae (fig. 50).Discussion: Bucerocolpocephalum deignani resembles most closelyB. emersoni. Male terminal abdominal sternites VIII and IX lackthe lateral notch in B. deignani that is present in B. emersoni. MalegenitaUa have parameres anteriorly enlarged, not curved inwardly,in B. deignani and anteriorly slender, curved inwardly with eachlateral point reaching endomeres in B. emersoni; endomeres lack thepair of posterior points in B. deignani that are present in B. emersoni.The female terminal abdominal tergite lacks thick setae medially onthe posterior margin in B. deignani, but thick setae are present alongthis entire margin in B. emersoni. The female abdominal sternite
NO. 3558 MALLOPHAGA?ELBEL 39VIII lacks the internal triangular sclerite in B. deignani that is presentin B. emersoni.Material examined: 39 males and 28 females from fresh and driedmaterial collected in the Oriental region.Type host: Anorrhinus g. galeritus (Temminck, 1831).Type material: Holotype male from CNHM skin from Kinabatan-gan, North Borneo, May 18, 1950, collected by D. D. Davis, REEin CNHM. Paratypes: 2 males with same data. Additional typesfrom Anorrhinus g. carinatus (Blyth, 1845): Allotype female fromUSNM skins from Trang, Thailand, 1896 and 1899, collected by W.L. Abbott, REE in USNM. Paratypes: 1 male and 1 female withsame data; 27 males and 18 females from Lamo and Chong, Muang,Trang, Thailand, March 1963, collected by Wichit Suwan Laong,USNM; 8 males and 7 females from Na Wong, Ban Na, Muang,Phatthalung, Thailand, Mar. 6, 1963, collected by Wichit SuwanLaong, USNM: 1 female from BL skin from Khao Phap Pha Mt.,Ban Na, Muang, Phatthalung, Thailand, Sept. 4, 1954, collected byB. Lekagul, REE.Bucerocolpocephalum deignani is named for Mr. H. G. Deignan,Division of Birds, U.S. National Museum, in appreciation for thefresh material he collected in Thailand, for supplying identificationsand information on hosts, and for permission to examine hornbillskins for Mallophaga in the USNM.Genus Bucerophagus BedfordFigures 66-69Bucerophagus Bedford, 1929, p. 509, figs. 11, 12. [Genotype: Bucerophagusafricanus Bedford, 1929.]Antimenopon Eichler, 1947, p. 3, figs. 3-5. [Genotype: Menopon forcipatumNitzsch, 1874.]Head triangular, width Iji to 1% times that of length. Foreheadnarrower anteriorly. Temples expanded. Antenna 4-jointed, thirdsegment constricted at base, and terminal segment capitate withdefinite signs of division into two parts, either by transverse line ormarginal indentation. Antennary fossa deep, covered above by ex-pansion of lateral margin of head, posterior margin of which bears eyewith double cornea. Dorsal-lateral margin of forehead anterior toeye with shallow notch, Gular region mth 2-8 setae varying in lengthon each lateral margin. Pronotimi expanded anteriorly with posteriormarginal row of long setae. Metanotum expanded posteriorly withposterior marginal row of long setae and 2-5 short setae on each lateralmargin. Metanotum separated from mesonotum and from pleurites.The sclerotized median button behind the prothorax (fig. 25m) is a
40 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 120
vestige of the mesonotmn; the supposed mesonotuin, the narrowsclerotized band posterior to this button, is a mere extension of thesubcoxae (Cope, 1941). Thoracic sternal plates as shown in figures66b, 66c, 68b, 68c, and 70. Metasternal plate trapezoidal or triangular,expanded anteriorly, with 6-34 setae. Venter of third femora andposterolateral margins of abdominal sternites IV and V each with orwithout large thick brushes of normal setae. Abdominal segmentsconsist of tergites, sternites, and pleurites, the latter without prolonga-tion of posteroventral angles. Abdominal tergites each with aposterior marginal row of setae, the most laterad being the post-spiracular seta. Each lateral margin of abdominal tergites II-VIIIwith or without 1-5 short setae between the spiracle and postspiracularseta. Sternites and plemites each with a posterior marginal row oflong setae and with numerous shorter setae. Male terminal abdom-inal sternites VIII and IX either fused or not but with completedivision from abdominal sternite VII (figs. 67, 69). Male genitaliaas illustrated for each species with parameres branched anteriorly andeither split or unsplit posteriorly. Female terminal abdominal seg-ments as illustrated for each species with lateral processes arising fromventral sclerite between vulva and anus with long stout setae andstrong spines. Females larger than males, usually with more abdom-inal sternal setae but general shape and chaetotaxy similar to that ofmales except for terminal abdominal segments.Bucerophagus resembles both Chapinia (figs. 23-26) and Bucerocol-pocephalum (figs. 64, 65) but differs in several characters: The termi-nal segment of the antenna shows definite signs of division into tw^oparts either by transverse line or marginal indentation in Bucerophagusand Bucerocolpocephaluw,, but there is no sign of division in Chapinia.The posterior margin of the expansion of the lateral margin of thehead covering the antennary fossa has an eye with a double corneain Bucerophagus and Chapinia, but an eye is absent in Bucerocol-pocephalum. The gular region lacks a lateral ridge and each lateralmargin has at most 8 setae in Bucerophagus and Chapinia, but eachlateral ridge has 8-11 setae in Bucerocolpocephalum. The metasternalplate is trapezoidal or triangular in Bucerophagus and Chapinia butoval in Bucerocolpocephalum. The venter of the third femora mayhave brushes of normal setae in Bucerophagus and Chapinia but hascombs of setae in Bucerocolpocephalum; similar brushes are presenton posterolateral margins of abdominal sternites IV and V in Bucero-phagus and abdominal sternites IV-VI in Chapinia, but combs ofsetae are present on posterolateral margins of abdominal sternite IV inBucerocolpocephalum. Each lateral margin of abdominal tergitesII-VIII may have 1-5 short setae between the spiracle and post-spiracular seta in Bucerophagus and Bucerocolpocephalum, but one
NO. 3558 MALLOPHAGA?ELBEL 41
short seta may be present on margins of abdominal tergites II-VI inChapinia. Male terminal abdominal sternites VIII and IX may befused in Bucerophagus and Bucerocolpocephalum, with a completedivision from abdominal sternite VII, but abdominal sternites VIIIand IX are fused in Chapinia with a partial division only from ab-dominal sternite VII. Male genitalia of Bucerophagus are longer thanin Bucerocolpocephalum; parameres are branched anteriorly in Bucero-phagus but are slender or expanded anteriorly in Chapinia. Lateralprocesses arising from the ventral sclerite between the female vulvaand an\is have long stout setae and strong spines in Bucerophagus andBucerocolpocephalum, but only long stout setae in Chapinia. Thefemale anal fringe prominent in Bucerophagus and Chapinia is weakin Bucerocolpocephalum.The male genitalia and details of the male and female terminalabdominal segments are the best characters for separating species ofBucerophagus. Other characters useful in species separation are:The shape of the metasternal plate and the number of setae present;the presence or absence of brushes of normal setae on the venter ofthe third femora and posterolateral margins of abdominal sternitesIV and V; the number present or absent of short setae on each lateralmargin of abdominal tergites III-VIII between the spiracle andpostspiracular seta; the total number of setae on each of abdominalsternites I and II. The number and length of setae on the lateralmargins of the gular region are too variable to be of much use inseparating species.Eichler (1947) believed that the lack of brushes, the rounded pro-jected lobe on the posterior end of the male abdomen, the specific malegenital apparatus, and the female anal ring of setae were enough toplace Menopon Jorcipatum Nitzsch in a separate genus. Hopkins andClay (1952) correctly placed M.Jorcipatum in the genus Bucerophagus.Since there are several characters separating B. forcipatus from thecomplex B. productus and B. africanus, it is believed here that therelationship can be shown best by species-groups.Hosts : Species of Bucerophagus have been found only on the generaBuceros, Rhinoplax, and Bucorvus of the avian family Bucerotidae.The forcipatus Species-GroupAs illustrated in figures 66, 67. Differing from the productusspecies-group in the following combination of characters: Headwidth iy2 to 1% times that of length; metanotum with two short setaeon each lateral margin and without setae on anterior margin; meta-sternal plate with less than 14 setae; venter of third femora andabdominal sternites IV and V without brushes; each lateral margin ofabdominal tergites II-VIII without short setae between the spiracle
42 PROCEEDINGS OF THE NATIONAL MUSEUM ^ol. 120and postspiracular seta; females with approximately the same numberof abdominal sternal setae as males; both sexes having abdominalsternite I with fewer than 20 total setae and abdominal sternite IIwith fewer than 44 total setae; male terminal abdominal sternite IXprojecting posteriorly as rounded lobe and with complete divisionfrom abdominal sternite VIII; male genitaUa with parameres poste-riorly split and curved inwardly; female terminal abdominal tergitewith fewer than 12 setae on posterior margin; female abdominalsternite VIII with fewer than 24 setae on posterior margin.Hosts: Bucerophagus Jorcipatus has been found only on the generaBuceros and Bhinoplax of the avian family Bucerotidae.Bucerophagus forcipatus (Nitzsch)Figures 51, 52, 60, 66, 67Menopon forcipatum "Nitzsch."?Giebel, 1874, p. 289, pi. 15 (figs. 7, 8.) [Typehost: Buceros rhinoceros=^Buceros rhinoceros sumatranus Schlegel and Muller,1840.]Antimenopon forcipatum "Nitzsch in Giebel."?Eichler, 1947, pp. 3, 20, figs. 3-5.Bucerophagus forcipatus (Nitzsch).?Hopkins and Clay, 1952, p. 64.Eichler's description and figures are not recognizable. He des-ignated specimens from Buceros rhinoceros from Sumatra as neotypematerial, but he did not select a neotype. His shde specimens havebeen remounted and examined; the male, 2275 ji, is designated herebyas neotype. The slide has been so labeled. The female, 2275 jf,is mounted on the same slide with the neotj^pe.Male: As illustrated in figure 67. Smaller than Bucerophagusafricanus in all measurements except length of head and width ofmetathorax (table 12). Metasternal plate trapezoidal, expandedanteriorly, with 6-12 setae (fig. 66c). Abdominal sternite I with6-18 total setae and abdominal sternite II with 30-36 total setae.Terminal abdominal segments as shown in figure 67d. GenitaUa asshown in figm-e 60.Female: As illustrated in figure 66. Smaller than Bucerophagusafricanus in all measurements except width of metathorax (table 12).Resembles the male except that abdominal sternite II has 34-42total setae. Terminal abdominal tergite with 8 long and 2 shortsetae on posterior margin; abdominal sternite VIII with 18-22 longand 4 short setae on posterior margin (fig. 51). Anal fringe with44-54 setae (fig. 52).Material examined: 49 males and 66 females from fresh and driedmaterial collected in the Oriental region; neotype male and specimensfrom the same series, 1 male and 4 females, from WEC 2275, SMNH.According to Eichler (1947), WEC 2275 was collected in Sumatra byE. Mjoberg; from the type host: 23 males and 29 females from
NO. 3558 MALLOPHAGA?ELBEL 43USNM skins from Tarussan Bay, West Sumatra, 1904-1905, collectedby W. L, Abbott, REE; from Buceros rhinoceros borneoensis Schlegeland Miiller, 1840: 1 male and 6 females from Serabang Bay, Sarawak,Borneo, Jan. 11, 1958, BMNH 1958-737; 4 females from Borneo,Meinertzhagen 10890, BMNH; 2 males and 5 females from CNHMskin from Sapagayo Forest Reservation, Sandakan, North Borneo, July27, 1950, collected by R. F. Inger and D. D. Davis, REE; fromBuceros bicornis homrai Hodgson, 1 832 : 7 males and 5 females fromBan Khlua Klang, Praclmap Khiri Khan, Thailand, December 1952,collected by Robert E. Elbel, and H. G. Deignan, USNM; 11 malesand 8 females from Khlong Khlung, Kamphaeng Phet, Thailand,Apr. 7, 1953, collected by Robert E. Elbel and H. G. Deignan, USNM;1 male and 3 females from Ban Muang Khai, Tha Li, Loei, Thailand,Dec. 7, 1953, collected by Robert E. Elbel, USNM; 2 males and 1female from Banghin, Kapoe, Ranong, Thailand, Feb. 6, 1963,collected by Wichit Suwan Laong, USNM; from Rhinoplax vigil(J. R. Forster, 1781): 1 female from Borneo, Meinertzhagen 10888,BMNH.Drawings were made of a male and a female from Buceros bicornishomrai collected in Khlong Khlung, Thailand. Specimens in USNM.The productus Species-GroupSpecies similar in shape to Bucerophagus africanus (figs. 68,69).Differing from the forcipatus species-group in the following combina-tion of characters: Head width 1}^ to IK times that of length; metano-tum with 3-5 short setae on each lateral margin and with 6-8 setaeon anterior margin; metasternal plate with more than 20 setae;venter of third femora and posterolateral margins of abdominalsternites IV and V each ^vith thick brushes of normal setae; each lateralmargin of abdominal tergites II-VIII with 1-5 short setae betweenthe spiracle and postspiracular seta; females with more abdominalsternal setae than males; both sexes having abdominal sternite Iwith more than 20 total setae and abdominal sternite II with morethan 60 total setae; male terminal abdominal sternites VIII and IXneither projecting posteriorly as rounded lobe nor with division butwith complete division from abdominal sternite VII; male genitaliawith parameres straight and not split posteriorly; female terminalabdominal tergite with more than 24 setae on posterior margin;female abdominal sternite VIII with more than 28 setae on posteriormargin.Hosts: Species of the productus species-group have been foundonly on the genus Bucorvus of the avian family Bucerotidae.
44 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 120Bucerophagus productus (Burmeister)Figures 53, 54, 61, 62, 70Colpocepfialum productum Burmeister, 1838, p. 439. [Type host: Buceros dbys-sinicus^Bucorvus abyssinicus (Boddaert, 1783).]Colpocephalum vittatus Giebel, 1866, p. 394 (nomen nudum),Colpocephalum productum "Nitzsch."?Giebel, 1874, p. 266, pi. 14 (figs. 2, 3).Colpocephalum eurijgaster Piaget, 1888, p. 162, pi. 4 (fig. 5). [Type host: Lep-toptilus argala eTror=^Bucorvus abyssinicus.]Bucerophagus productus "Nitzsch in Burmeister"?Conci, 1950, p. 78, figs. 1-7.Bucerophagus productus (Burmeister).?Hopkins and Clay, 1952, p. 64.A neotype, in the Zoologischen Institute der Universitat Halle,Germany, was erected by Conci, who redescribed and figuredBucerophagus productus from Bucorvus abyssinicus collected in eastAfrica by Prof. E. Zavattari. A male from the same series is in thecollection of Mr. G. II. E. Hopkins, Zoological Museum, Tring,Hertsfordshire, England.A lectotype male was designated by Clay (1951a) from the Col-pocephalum eurygaster syntypes in the Piaget collection, and it isnow in the British Museum (Natual History), BM 1157a, wdth 3syntype males and 1 syntype female, BM 1157 and 1158.Clay (1951a) stated that Bucerophagus productus also seemed toocciu' naturally on Bucorvus leadbeateri ("Vigors).Male: Smaller than Bucerophagus ajricanus in all measurementsexcept length of head; approximately the same size as B. Jorcipatus(table 12). Metasternal plate triangular, expanded anteriorly, with22-26 setae (fig. 70). Each lateral margin of abdominal tergitesIII-VIII with 2-4 short setae between the spiracle and postspiracularseta except for tergite VII "with 1 or 2 setae and tergite VIII with oneseta. Abdominal sternite I with 22-24 total setae and abdominalsternite II with 66-68 total setae. Terminal abdominal segmentsas shown in figure 61. Genitalia as shown in figure 62, with branchconnecting parameres posterior to endomeres pointed medially.Female: Approximately the same size as Bucerophagus ajricanus;larger than B. forcipatus in all measurements except width of headand width of metathorax (table 12). Resembles the male exceptthat metasternal plate has 26-34 setae. Abdominal sternite I with28 total setae and abdominal sternite II with 74-76 total setae. Ter-minal abdominal tergite with 26-38 setae on posterior margin; ab-dominal sternite VIII with 30-36 setae on posterior margin (fig. 53).Anal fringe with 42-56 setae (fig. 54)
.
Discussion: Bucerophagus productus resembles most closely B.ajricanus which, however, is much more pigmented than B. productus.Males of B. productus are much smaller than females, but males ofB. ajricanus are only slightly smaller than females. The metasternalplate is triangular in B. productus and trapezoidal in B. ajricanus;
NO- 3558 MALLOPHAGA?ELBEL 45this plate has fewer setae in both sexes of B. productus than in cor-responding sexes of B. africanus. Each lateral margin of abdominaltergites III-VI between the spiracle and postspu-acular seta has moreshort setae in males and fewer in females of B. j^roductus than incorresponding sexes of B. africanus. Abdominal sternite I hasapproximately one-half the number of setae in both sexes of B.productus as in corresponding sexes of B. africanus; abdommal ster-nite II has slightly more setae in males and slightly fewer setae infemales of B. productus than of B. africanus. The male abdominalsternite VIII lacks the central T-shaped plate in B. productus whichis present in B. africanus (fig. 69d). The male genitalia has the in-ternal branch connecting the parameres posterior to the endomerespomted medially in B. productus but rounded in B. africaiius. Thefemale anal fringe has fewer than 58 setae in B. productus but morethan 58 setae in B. africanus.Comparison on different hosts: No morphological differences werefound between specimens of Bucerophagus productus found on thetwo hosts, Bucorvus ahyssinicus and B. leadbeateri, so standard meas-urements were tested against the null hypothesis that there wereno differences in measurements (tables 4-7).Terminology and formulae are as follows:
XI = mean measurement of B. productus specimens on host 1, Bucorvus ahyssinicusX2= mean measurement of B. productus specimens on host 2, Bucorvus leadbeateriD= difference in mean measurements, (xj? X2) or (x2? xi)
,2 . S(X-X)2 Sx2-(Sx)2/NS2= variance =?^^;^?-^= --^^
?
??N?
1
N?
1
s?= variance of measurements of B. productus specimens on host 1, B. ahyssinicus83= variance of measurements of B. productus specimens on host 2, B. leadbeateriF= the ratio of the larger variance divided by the smaller=^ or ^=?si^ Si NdSE-Standard Error of p^ /S(x-xi)2+ S(x-X2)2 /J_ , JL^V No+ Nd-2 VN^^Nd/CL=- Confidence Limits for D =D? (SE) (t.05)Since it is not possible by measurements alone to decide fromwhich host specimens came, populations from two hosts are con-sidered conspecific.Material examined: 39 males and 44 females from fresh and driedmaterial collected in the Ethiopian region; from the type host: 4males and 2 females from Gula, Uganda, Africa, July 10, 1936, col-lected by G. H. E. Hopkins, GHEH; 6 females from Ethiopia, Africa,March 1909, BMNH 3673; 1 female from CNHM skin from Africa,Jan. 30, 1946, REE; 3 males and 1 female from USNM skins fromSirre, Ethiopia, Africa, Feb. 13, 1912, collected by Childs Frick,REE; 3 males from USNM skins from Uganda, Africa, January-221-522?67 5
46 PROCEEDINGS OF TPIE NATIONAL MUSEUM vol. 120February 1910, collected by E. A. Mearns, llEE; from Bucorvusleadbeateri (Vigors, 1825): 9 males and 18 females from CNHMskins from Chitau, Bihe, Angola, Africa, 1932-1934, collected byJean Bodaly, REE; 15 males and 14 females from CNHM skin fromPondi, Benguela, Angola, Africa, Sept. 14, 1936, collected by K. H.Prior, REE; 5 males and 2 females from CNHM skins from Kari Pan,Makari, Becliuanaland, Africa, August 1930, collected by Vernay,Lang, and Roberts, REE.Drawings were made of a male and a female from the type hostcollected in Gula, Uganda, Africa. Specimens in GHEH.Bucerophagus africanus BedfordFigures 55, 56, 63, 68, 69Bucerophagus africanus Bedford, 1929, p. 509, figs. 11, 12. [Type host: Bucorvusschlegeli Roberts= Bucorvus leadbeateri (Vigors, 1825).]Bucerophagus africanus Bedford?Hopkins and Clay, 1952, p. 64.Clay (1951a) stated that according to Mr. G. H. E. HopkinsBucerophagus africanus also occurred on Bucorvus abyssinicus.Male: As illustrated in figure 69. Larger than either Bucerophagusforcipatus or B. productus except for length of head (table 12). Meta-sternal plate trapezoidal, expanded anteriorly, with 30-34 setae(fig. 68c). Each lateral margin of abdominal tergites III-VIII withtwo short setae between the spiracle and postspiracular seta exceptfor tergites III and VIII each with one short seta. Abdominalsternite I with 40 total setae and abdominal sternite II with 62total setae. Abdominal sternite VIII with central T-shaped plate(fig. 69d). Genitalia as shown in figure 63.Female: As illustrated in figure 68. Approximately the same sizeas Bucerophagus productus: larger than B. forcipatus in all measure-ments except width of metathorax (table 12). Resembles the maleexcept that metasternal plate has 36-42 setae. Each lateral marginof abdominal tergites III-VIII with 3-5 short setae between thespiracle and postspiracular seta except for tergite VIII with one seta.Abdominal sternite I with 50 total setae and abdominal sternite IIwith 86 total setae. Terminal abdominal tergite with 32-40 setae onposterior margin; abdominal sternite VIII with 28-38 setae on pos-terior margin (fig. 55). Anal fringe with 60-66 setae (fig. 56).Discussion: Bucerophagus africanus resembles most closely B. pro-ductus which, however, is not as pigmented as B. africanus. Males ofB. africanus are only shghtly smaller than females, but males ofB. productus are much smaller than females. The metasternal plateis trapezoidal in B. africanus and triangular in B. productus; this platehas more setae in both sexes of B. africanus than in corresponding
NO. 8558 MALLOPHAGA?ELBEL 47
sexes of B. productus. Each lateral margin of abdominal tergitesIII-VI between the spiracle and postspiracidar seta has fewer shortsetae in males and more in females of B. africanus than in correspond-ing sexes of B. productus. Abdominal sternite I has approximatelytwice the number of setae in both sexes of B. africanus as in corre-sponding sexes of B. productus; abdominal sternite II has slightly-fewer setae in males and slightly more setae in females of B. africanusthan of B. productus. The male abdominal sternite VIII has a centralT-shaped plate in B. africanus which is absent in B. productus. Themale genitalia has the internal branch connecting the parameresposterior to the endomeres rounded medially in B. africanus butpointed in B. productus. The female anal fringe has more than 58setae in B. africanus but fewer than 58 setae in B. productus.Comparison on different hosts: No morphological differences werefound between specimens of Bucerophagus africanus found on the twohosts, Bucorvus abyssinicus and B. leadbeateri, so standard measure-ments were tested against the null hypothesis that there were nodifferences in measurements (tables 8-11).Terminology and formulae were the same as those used for Bucer-ophagus productus except as follows:
xi= mean measurement of B, africanus specimens on host 1, Bucorvus abyssinicusX2= mean measurement of B. africanus specimens on host 2, Bucorvus leadbeateriSi= variance of measurements of B. africanus specimens on host 1, Bucorvusabyssinicussi= variance of measurements of B. africanus specimens on host 2, BucorvusleadbeateriSince it is not possible by measurements alone to decide fromwhich host specimens came, populations from two hosts are con-sidered conspecific.Material examined: 13 males and 13 females from fresh and driedmaterial collected in the Ethiopian region; from the type host: 1 femalefrom Mafa, South-West Africa, February 1923, BMNH; 9 males and7 females from Cameroons Zoo, Africa, November 1936, BMNH 8127;1 female from CNHM skins from Kari Pan, Makari, Bechuanaland,Africa, August 1930, collected by Vernay, Lang, and Roberts, REE;from Bucorvus abyssinicus (Boddaert, 1783): 1 male and 1 female fromKoubadge, French Cameroons, Africa, July 1947, collected by V.Aellen, BMNH 1954-487; 2 males and 2 females from USNM skinsfrom Sirre, Ethiopia, Africa, Feb. 13, 1912, collected by Childs Frick,REE; 1 male and 1 female from Nyala, Sudan, Africa, Feb. 12, 1949,KCE.Drawings were made of a male and a female from the type hostcollected in the Cameroons Zoo. Specimens in BMNH.
48 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 120Aviparasitological RelationshipsSince Mallophaga are obligatory, usually highly host-specific,external parasites, their distribution is dependent on the distributionof their hosts (Emerson and Ward, 1958). The arrangement of theMallophaga based on morphological similarities does not followexactly Peters' (1945) phylogenetic arrangement of the hornbillhosts (table 13). In the genus Chapinia, species of the lophocerusspecies-group infest hosts in the genera Tockus, Bycanistes, andCeratogymna of the Ethiopian region, but species of the acutovulvataand hirta species-groups infest hosts in the genera Tockus, Anorrhinus,Penelopides, Rhyticeros, Anthracoceros, and Buceros of the Orientaland Australasian regions. Species of Bucerocolpocephalum infesthosts in the genera Ptilolaemus and Anorrhinus of the Oriental region.In the genus Bucerophagus, the species B. jorcipatus of the forcipatusspecies-group infests hosts in the genera Buceros and Rhinoplax ofthe Oriental region, but species of the productus species-group infesthosts in the genus Bucorvus of the Ethiopian region.Tockus is the only hornbill genus with amblyceran lice that hasmembers in both the Ethiopian and Oriental regions (table 13).Yet the amblyceran, Chapinia clayae, from the Oriental species,Tockus birostris and Tockus g. griseus, does not resemble members ofthe lophocerus species-group which infest other species of Tockus.Instead, C. clayae resembles most closely C. acutovulvata from theOriental species of Anthracoceros. It would appear that there hasbeen more recent contact between the Indian Tockus and Anthraco-ceros whose ranges overlap than between the more nearly relatedIndian Tockus and African Tockus. Kellogg (1896) was the first tomention that Mallophaga live their entire lives on the host bird andthat infestation of new hosts is accomplished by the actual migrationof individuals from one bird to another, during copulation, nesting,or roosting. Clay (1949b) stated that normally bu-ds of differentspecies did not come into close enough contact for lice to be trans-ferred from host to host but that interchange of lice could take placebetween predator and prey, nestling and foster parent in broodparasites, by the use of common dust baths (according to Hoyle,1938), and by phoresy, which is the transfer of lice by Hippoboscidflies. In the case of brood parasites she stated that for the Eiu-opeanCuckoo, Cuculus canorus, lice of the foster parents had never beenestablished on the cuckoo. She further stated that establishmenton the new host might be prevented by competition of the alreadyadapted resident louse population, by the host specificity of theimmigrant louse making feeding and development on the new hostimpossible, or by the fact that only males or unfertilized females hadbeen introduced. Clay (1962) described natural straggling as
NO. 8558 MALLOPIIAGA?ELBEL 49
occurring between hosts that happened to be nesting in close prox-imity; she stated that establishment on the new host might be facili-tated by the absence of a resident louse.It would appear that both lophocerus and acutovulvata species-groupsshared a common ancestor of Chapinia on Tockus before the Indianand African Tockus became separated. Once separated, the Chapiniaevolved as did the birds to the recognized species within each species-group. Natural straggling may have accounted for establishmenton some of the hosts.Clay (1949b) mentioned that the chief factor influencing theproduction of allopatric species and genera of Mallophaga has beenthe successive sphtting of the host populations during the evolutionof the birds, thus leaving isolated louse populations. The lousepopulation is considered as comprising all individuals that can inter-breed because their hosts can interbreed (Clay, 1958). Kellogg(1896) stated that with the spreading of the ancestral bird species,geographical races arose within the limits of the species. With timeand isolation, these races became distinct species which were oftendistinguished only by superficial differences in color, etc. TheMallophaga remained practically unaffected since their environmentwas essentially unchanged. The environment of the Mallophaga,the physical and chemical composition of the feathers and blood,changes more slowly than do other factors leading toward speciationof the bird; until this environment changes, the Mallophaga wouldremain unchanged (Clay, 1949b). For example, in the acutovulvataspecies-group, the hosts Anthracoceros coronatus and A. convexus arenow considered to be full species distinct from A. a. albirostris, A. a.leucogaster, and A. marchei, yet all these hosts bear the same speciesof Mallophaga, Chapinia acutovulvata. Similarly, in the lophocerusspecies-group, Chapinia bucerotis infests eight subspecies in fourspecies of Bycanistes, and C. lophocerus infests six subspecies in fourspecies of Tockus. In the forcipatus species-group, Bucerophagusjorcipatus infests three subspecies in three species and two genera.Although Anthracoceros marchei is restricted to the Philippines, itis host to Chapinia acutovulvata which infests other Anthracocerosspecies with wider distribution in the Oriental region. A. montanialso is restricted to the Philippines, but its amblyceran parasite,C. hoplai, resembles most closely C. boonsongi from Rhyticeros undu-latus, which is distributed elsewhere in the Oriental region. Chapiniawenzeli from Penelopides panini and C. traylori from Buceros hydro-corax both resemble most closely C. blakei from Rhyticeros leucoce-phalus. Although the host genus Buceros is not considered to be asrelated to Rhyticeros as is Penelopides, the host species from whichC. wenzeli, C. blakei, and C. traylori were obtained are all restricted
50 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 120to the Philippines. Thus, C. traylori may have evolved as a result ofnatural straggling from the stock that gave rise to C. wenzeli andC. blakei and subsequently became estabUshed on the host B. hydro-corax.Chapinia lydae from Rhyticeros cassidix has the characters of theacutomlvata species-group but superficially resembles members of thehirta species-group of which C. muesebecki, from Penelopides e. exar-hatus, like C. lydae, is restricted to the Australasian region (table 13).Chapinia hirta, the other member of the hirta species-group, infestssubspecies of Rhyticeros plicatus in both the Oriental and Australasianregions.The hornbill genus Buceros is host to both Chapinia and Bucero-phagus (table 13); however, these Mallophaga do not infest the samehosts. Buceros hydrocorax, the host of Chapinia traylori, is restrictedto the Philippines, but B. rhinoceros and B. bicornis, the hosts ofBucerophagus jorcipatus, are distributed elsewhere in the Orientalregion. On the other hand, the mallophagan species, Chapiniawaniti and Bucerocolpocephalum deignani, do infest the same host,Anorrhinus galeritus carinatus.In the productus species-group both Bucerophagus productus and B.ajricanus infest the two hosts, Bucorvus abyssinicus and B. lead-beateri. The population of each mallophagan species on each hostcould not be separated morphologically or statistically. Thus, onlythe two species, Bucerophagus productus and B. ajricanus, could berecognized. Similarly, Clay (1955) recognized only the one species,Bucorvellus docophorus, although specimens from Bucorvus lead-beateri showed a tendency to be smaller in size than specimens fromB. abyssinicus. She further stated that it would be expected fromHarrison's rule (1915) that specimens from B. leadbeateri, the smallerhost, would be smaller than specimens from the larger B. abyssinicus.However, Mackworth-Praed and Grant (1952) stated that B. lead-beateri was the largest of the HornbUls; they gave wing measurementsfor B. leadbeateri as 509-595 mm. and for B. abyssinicus as 495-595mm.Harrison (1915) stated the rule that bears his name: that in general,when a mallophagan genus is well distributed over a considerablenumber of nearly related hosts, the size of the parasite is roughlyproportional to the size of the host. Chapinia camuri, the smallestspecies of Chapinia, infests the smallest hornbill, Tockus camurus,but Chapinia traylori, the largest species of Ch,apinia, does not infestRhyticeros undulatus, the largest host for species of Chapinia.
NO. 3568 MALLOPHAGA?ELBEL 51SummaryAmblyceran Mallophaga of the family Menoponidae were examinedfrom 53 species or subspecies of hornbills. Descriptions and illustra-tions are presented for 22 species in three genera of hornbiU Meno-ponidae of which 14 species are new. The genus Chapinia nowcontains three species-groups and 17 species of which 12 are new.The genus Bucerophagus now contains two species-groups and threespecies. The new genus Bucerocolpocephalum type emersoni iserected here for two new comb-bearing species. The new speciesare as follows: Chapinia fasciati, C. camuri, C. clayae, C. waniti,C. malayensis, C. hoplai, C. boonsongi, C. wenzeli, C. hlakei, C. tray-lori, C. lydae, C. muesebecki, Bucerocolpocephalum emersoni, and B.deignani. New synonymy is C. acutovulvata (Piaget, 1881) (=C.mjobergi (Eichler, 1947)). New type designations are: A neotype forChapinia hirta (Kudow, 1866), a neotype for Bucerophagus forcipatus(Nitzsch, 1874), and a lectotype for C. bucerotis (Kellogg, 1908).Differential characters are listed for genera, species-groups, andspecies, and a key is provided for separating the species.The amblyceran species-groups are confined to the Ethiopianregion or to the Oriental and Australasian regions as shown in atable of the hornbill hosts, their distribution, and amblyceran para-sites. Tockus is the only hornbill genus with amblyceran lice thatis present in both the Ethiopian and Oriental regions; however,Chapinia clayae of the acutovulvata species-group from the Orientalspecies of Tockus resembles C. acutovulvata from Oriental species ofAnthracoceros more closely than species of the lophocerus species-group from Ethiopian species of Tockus. It would appear, thatthere has been more recent contact between the Oriental species ofTockus and Anthracoceros, whose ranges overlap, than between themore closely related Oriental and Ethiopian species of Tockus.
Literature CitedBedford, G. A. H.1920. Anoplura from south African hosts, pt. 2. Rept. Vet. Res. SouthAfrica, vols. 7, 8, pp. 709-741, 7 pis.1929. Anoplura (Siphunculata and Mallophaga) from south African hosts.Rept. Vet. Res. South Africa, vol. 15, pp. 501-549, 34 figs.1930. New genera and species of Mallophaga from south African hosts.Rept. Vet. Res. South Africa, vol. 16, pp. 153-173, 16 figs.BiTRMEISTER, C. H. C.1838. Mallophaga. In Handbuch der Entomologie, vol. 2, pp. 418-443.
52 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 120Clat, Theresa1947. A preliminary key to the genera of the Menoponidae (Mallophaga)
.
Proc. Zool. Soc. London, vol. 117, pp. 457-477, 40 figs.1949a. Systematic notes on the Piaget collections of Mallophaga, pt. 1.Ann. Mag. Nat. Hist. ser. 12, vol. 2, pp. 811-838, 895-921, 6 figs.1949b. Some problems in the evolution of a group of ectoparasites. Evolu-tion, vol. 3, pp. 279-299, 4 figs.1951a. Systematic notes on the Piaget collections, pt. 2. Ann. Mag. Nat.Hist. ser. 12, vol. 4, pp. 173-182.1951b. Systematic notes on the Piaget collections of Mallophaga, pt. 3.Ann. Mag. Nat. Hist. ser. 12, vol. 4, pp. 1159-1168, 15 figs., 1 pi.1954. The postspiracular seta and sensillus in the Mallophaga (Insecta).Ann. Mag. Nat. Hist. ser. 12, vol. 7, pp. 716-718, 2 figs.1955. A new genus of Ishnocera (Mallophaga). Proc. Roy. Ent. Soc. Lon-don (B), vol. 24, pp. 1-6, 7 figs., 1 pi.1956. Phthiraptera. In Tuxen, Taxonomist's glossary of genitalia in in-sects, pp. 145-148, 4 figs.1958. Revisions of Mallophaga genera: Degeeriella from the Falconiformes.Bull. British Mus. (Nat. Hist.) Ent., vol. 7, pp. 5-217, 164 figs.,9 pis.1961. A new genus and species of Menoponidae (Mallophaga Insecta) fromApteryx. Ann. Mag. Nat, Hist. ser. 13, vol. 3, pp. 571-577, 12figs., 1 pi.1962. A key to the species of Aclornithophilus Ferris with notes and descrip-tions of new species. Bull. British Mus. (Nat. Hist.) Ent., vol. 11,pp. 191-252, 72 figs., 8 pis.CoNci, Cesarb1950. Mallophaga. Riv. Biol. Colon., vol. 10, pp. 77-82, 7 figs.Cope, Oliver B.1941. The morphology of a species of the genus Tetrophthalmus (Mallophaga:Menoponidae). Microentomology, vol. 6, pp. 71-92, 10 figs.Deignan, H. G.1963. Checklist of the birds of Thailand. U.S. Nat. Mus. Bull., vol. 226,pp. 1-263, 1 fig.Eichler, von Wolfdietrich1947. Dr. E. Mjoberg's zoological collections from Sumatra, 15: Mallo-phaga. Ark. Zool., vol., 39 A, pp. 1-21, 40 figs.Emerson, K. C.1954. Review of the genus Menopon Nitzsch, 1818 (Mallophaga). Ann.Mag. Nat. Hist. ser. 12, vol. 7, pp. 225-232, 12 figs.Emerson, K. C, and Ward, Ronald A.1958. Notes on the Philippine Mallophaga, 1: Species from Ciconiiformes,Anseriformes, Falconiformes, Galliformes, Gniiformes, and Chara-driiformes (Philippine Zoological Expedition 1946-1947). Fieldi-ana, vol. 42, pp. 49-61, 4 figs.EwiNG, H. E.1927. Descriptions of new genera and species of Mallophaga together withkeys to some related genera of Menoponidae and Philopteridae.Journ. Washington Acad. Sci., vol. 17, pp. 86-96.Ferris, G. F.1923. The mallophagan family Menoponidae. Parasitology, vol. 16, pp.55-65.
NO. 3558 MALLOPHAGA?ELBEL 53GlEBEL, C.1866. Die im Zoologischen Museum der Universitat Halle aufgestelltenEpizoen nebst Beobachtungen iiber dieselben. Zeitschr. Ges.Naturwiss., vol. 28, pp. 353-397.1874. Insecta Epizoa, 329 pp., 20 pis.Harrison, L.1915. Mallophaga from Apteryx and their significance with a note on thegenus Rallicola. Parasitology, vol. 8, pp. 88-100, 6 figs,Hopkins, G. H. E,1941. Stray notes on Mallophaga, 15: Notes on the types of the Mallo-phaga described by Bedford, Ann, Mag. Nat. Hist, ser 11, vol. 7,pp. 274-294,Hopkins, G. H. E., and Clat, Theresa1952. A checklist of the genera and species of Mallophaga, 362 pp.HOYLE, W. L,1938. Transmission of poultry parasites. Trans. Kansas Acad. Sci., vol.41, pp. 379-384.Johnson, Phyllis T.1960. The Anoplura of African Rodents and Insectivores. U.S. Dept,Agric. Tech. Bull, vol. 1211, pp. 1-116, 180 figs.Kellog, V, L,1896, New Mallophaga, 1: With special reference to a collection made frommaritime birds of the bay of Monterey, California. Proc. Cali-fornia Acad. Sci., vol. 6, pp. 31-182, 4 figs., 14 pis.1908. Mallophaga. No. 4 of Corrodentia, no. 15 in Sjostedt, Wissenschaft-liche Ergebnisse der Schwcdischen Zoologischen Expedition nachdem Kilimandjaro, dem Meru und den umgebenden MassaisteppenDeutsch-Ostafrikas, 1905-1906 . , ., pp. 43-58, 1 pi.Mackworth-Praed and Grant, G, H, B.1952. Birds of East and Northeast Africa, ser. 1 ,vol. 1, 836 pp.NiTzscH, L.
?
see Giebel, 1874.Peters, J, L.1945. Bucerotidae. In Checklist of birds of the world, vol. 5, pp. 254-272.PlAGET, E.1880. Les Pediculines: Essai monographique, xxxix, + 714 pp., 56 pis.1881. Quatre Nouvelles Pediculines. Tijdschr, Ent., vol. 24, pp. 1-6, 1 pi.1885, Les Pediculines: Supplement, xii+201 pp., 17 pis.1888. Quelques nouvelles Pediculines. Tijdschr, Ent., vol. 31, pp. 147-168,2 pis.RuDOW, F,1866. Charakteristik neuer Federlinge. Zeitschr. Ges. Naturwiss., vol. 27,pp. 465-477.1809. Neue Mallophagen. Zeitschr. Ges. Naturwiss., vol. 34, pp. 387-407
54 PROCEEDINGS OF THE NATIONAL MUSEUMTable 1.
?
Measurements in mm. of Chapinia males (from specimens from whichdrawings were made and from hosts, in parentheses, other than type hosts)
MALLOPHAGA?ELBEL 55Table 2,
?
Measurements in mm. of Chapinia females (from specimens fromwhich drawings were made and from hosts, in parentheses, other than type hosts)
56 PROCEEDINGS OF THE NATIONAL MUSEUMTable 3.
?
Measurements in mm. of Bucerocolpocephalum (from specimens fromwhich drawings were made and from hosts, in parentheses, other than type hosts)
MALLOPHAGA?ELBEL 57Table 5.
?
Measurements in mm. and computations for Bucerophagus productusmales on host 2, Bucorvus leadbeateri (see p. 45 for explanation of formulae)
58 PROCEEDINGS OF THE NATIONAL MUSEUMTable 7.
?
Measurements in mm. and computations for Bucerophagus productus(see p. 45 for explanation of formulae)Females on host2, BucoTvusleadbeateri
MALLOPHAGA?ELBEL 59Table 8.
?
Measurements in mm. and computations for Bucerophagus africanusmales on host 1, Bucorvus abyssinicus (see p. 47 for explanation of formulae)
60 PROCEEDINGS OF THE NATIONAL MUSEUMTable 11.
?
Measurements in mm. and computations for Bucerophagus africanus(see p.47 for explanation of formulae)Females on host 2,BucoTvnsleadbeateri
MALLOPHAGA-?ELBEL 61Table 13.
?
Bucerotidae arranged phylogenetically (Peters, 1945) with geographicaldistribution and amblyceran parasites (blank indicates no lice found on host)
Host
62 PROCEEDINGS OF THE NATIONAL MUSEUMTable 13.
?
Bucerotidae arranged phylogenetically {Peters, 1945) with geographicaldistribution and amblyceran parasites (blank indicates no lice found on host)
?
Continued
Host
MALLOPHAGA?ELBEL 63
Figures 1-7.?^The lophocerus species-group. Male genitalia, ventral view: 1, Chapiniafasciati, new species, holotype; 2, C. lophocerus (Bedford), lectotype; 3, C. camuri, newspecies, holotype; 6, C. bucerotis (Kellogg), lectotype; 7, C. robusta Ewing. Male terminalabdominal segments, dorsal-ventral view: 4, C. lophocerus (Bedford), lectotype; 5, Cbucerotis (Kellogg), lectotype.
64 PROCEEDINGS OF THE NATIONAL MUSEUM
Figures 8-14.?The acutovulvata species-group. Male genitalia, ventral view: 8, Chapiniaclayae, new species, holotype; 9, C acutovulvata (Piaget); 12, C. malayensis, new species,holotype; 13, C. hoplai, new species, holotype; 14, C. boonsongi, new species, holotype.Male terminal abdominal segments, dorsal-ventral view: 10, C. acutovulvata (Piaget);11, C boonsongi, new species, holotype.
MALLOPHAGA?ELBEL 65
Figures 15-22.?The acutovuhata and hirta species-groups. Male genitalia, ventral view:15, Chapinia wenzeli, new species, holotype; 16, C. blakei, new species, holotype; 17,C. traylori, new species, holotype; 18, C. muesebecki, new species, holotype; 19, C. hirta(Rudow), neotype; 20, C. lydae, new species, holotype. Male terminal abdominal seg-ments, dorsal-ventral view: 21, C muesebecki, new species, holotype; 22, C. hirta (Rudow),neotype. (s= sclerite of genital sac.)
66 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 120
Figures 23, 24.
?
Ckapinia rohusta Ewing, dorsal-ventral view: 23, female; 24, male,(a^antenna of femalej b^prosternal plate of female; c^terminal abdominal segmentsof male.)
MALLOPHAGA?ELBEL 67
Figures 25, 26.
?
Chapinia iraylori, new species, dorsal-ventral view: 25, allotype female;26, holotype male. (a= antenna of female; b= prosternal plate of female; c= metasternalplate of male; d= first abdominal sternite of male; e= terminal abdominal segmentsof male; m=mesonotum; p=postspiracular seta.)
68 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 120
Figures 27-34.?The lophocerus and acutovulvata species-groups. Female terminal abdomi-nal segments, dorsal-ventral view: 27, Chapinia lophocerus (Bedford); 29, C. bucerotis(Kellogg); 31, C. robusia Ewing; 33, C. clayae, new species, allotype. Female analfringes, ventral view: 28, C. lophocerus (Bedford); 30, C. bucerotis (Kellogg); 32, C.robusta Ewing; 34, C. clayae^ new species, allotype. (h=sclerital hooks.)
MALLOPHAGA?ELBEL 69
Figures 35-39.?The acutovulvata species-group. Female terminal abdominal segments,dorsal-ventral view: 35, Chapinia acutovulvata (Piaget); 37, C. malayensis, new species,allotype; 38, C. hoplai, new species, allotype; 39, C. boonsongt, new species, allotype.Female anal fringes, ventral view: 36, C. acutovulvata (Piaget). (i==internal sclerite ofabdominal sternlte VIII.)
70 PROCEEDINGS OF THE NATIONAL MUSEUM
Figures 40-45.?The acutovulvata species-group. Female terminal abdominal segments,dorsal-ventral view: 40, Chapinia wenzeli, new species, allotype; 43, C. blakei, newspecies, allotype; 44, C. traylori, new species, allotype. Female anal fringes, ventralview: 41, C. wenzeli, new species, allotype; 42, C. blakei, new species, allotype; 45, C,traylori, new species, allotype. (i= internal sclerite of abdominal sternite VIII; sp=posteriorly directed setae on lateral projection of ventral sclerite between vulva andanus.)
MALLOPHAGA?EliBEL 71
Figures 46-52.?Female terminal abdominal segments, dorsal-ventral view: 46, Chapinialydae, new species, allotype; 48, C. hirta (Rudow); 49, Bucerocolpocephalum emersoni,new species, allotype; 50, B. deignani, new species, allotype; 51, Bucerophagus forcipatus(Nitzsch). Female anal fringes, ventral view: 47, C. hirta (Rudow); 52, Bucerophagusforcipatus (Nitzsch). (i= internal sclerite of abdominal sternite VIII.)
72 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 120
Figures 53-63.?Female terminal abdominal segments, dorsal-ventral view: 53, Bucero-phagus productus (Burmeister); 55, B. africanus Bedford. Female anal fringe, ventralview: 54, B. productus (Burmeister); 56, B. africanus Bedford. Male genitalia, ventralview: 57, Bucerocolpocephalum emersoni, new species, holotype; 58, B. deignani, newspecies, holotype; 60, Bucerophagus forcipatus (Nitzsch); 62, B. productus (Burmeister);63, B. africanus Bedford. Male terminal abdominal segments, dorsal-ventral view: 59,Bucerocolpocephalum deignani, new species, holotype; 61, Bucerophagus productus (Bur-meister).
MALLOPHAGA?ELBEL 73
Figures 64, 65.
?
Bucerocolpocephalum emersoni, new species, dorsal-ventral view: 64allotype female; 65, holotype male. (a= antenna of female; b= prosternal plate of female;c=metasternal plate of female; d= first abdominal sternite of female; e= terminal ab-dominal segments of male; 1= short lateral setae between spiracle and postspiracular seta.)
74 PROCEEDINGS OF THE NATIONAL MUSEUM
fk] t#
Figures 66, 67.
?
Bucerophagus forcipatus (Nitzsch), dorsal-ventral view: 66, female;67, male. (a== antenna of female; b=prosternal plate of female; c= metasternal plate offemale; d= terminal abdominal segments of male.)
MALLOPHAGA?ELBEL 75
Figures 68-70.
?
Bucerophagus africanus Bedford, dorsal-ventral view: 68, female; 69, male,(a= antenna of female; b=prosternal plate of female; c=metasternal plate of female;d= central T-shaped plate of male sternlte VIII.) B. productus (Burmeister): 70.metasternal plate of female.
76 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 120
Figures 71, 72.
?
Chapinia watiiti, new species, ventral view: 71, genitalia of holotype; 72,terminal abdominal segments of allotype. (sp= posteriorly directed setae on lateralprojection of ventral sclerite between vulva and anus.)