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A New Species of the Feather Mite Genus Pterotrogus Gaud (Analgoidea: Pteronyssidae) from the Ivory-billed Woodpecker Campephilus principalis L. (Aves: Piciformes)

A New Species of the Feather Mite Genus Pterotrogus Gaud
(Analgoidea: Pteronyssidae) from the Ivory-billed Woodpecker
Campephilus principalis L. (Aves: Piciformes)
Ann. Entomol. Soc. Am. 97(6): 000Ð000 (2004)
ABSTRACT A new feather mite species, Pterotrogus principalis sp. n., is described from the wing
plumage of the ivory-billed woodpecker, Campephilus principalis L. This is the Þrst record of a feather
mite species from this bird, which was distributed in southeastern North America and Cuba, but is
considered to be recently extinct.
KEY WORDS feather mites, Pterotrogus, new species, Campephilus principalis
THE IVORY-BILLED WOODPECKER,Campephilus principalis
L. (1758; Piciformes: Picidae), the largest woodpecker
of North America, once inhabited mature forests of
southeastern areas in the United States and in Cuba,
but at present is thought to be extinct. The last reliable
sightings of this species in the North American con-
tinent were in the 1950s, and in Cuba in the early 1990s
(Dennis 1979, Estrada and Alayo´n 1986, Lammertink
1992, Lammertink and Estrada 1995). Feather mites
associated with the ivory-billed woodpecker have not
previously been documented.
In the course of our current taxonomic investiga-
tions of feather mites associated with nonpasserine
birds (e.g., Dabert et al. 2002, Mironov et al. 2003), we
examined a museum skin specimen of the ivory-billed
woodpecker and recorded a new feather mite species
from this host. The new species belongs to the feather
mite genus Pterotrogus Gaud, 1981 (Analgoidea: Ptero-
nyssidae), which seems to be speciÞc to large-bodied
species of woodpeckers in North America (Faccini
and Atyeo 1981).
Among 10 feather mite families, representatives of
which have been recorded from woodpeckers (Gaud
and Atyeo 1996), the family Pteronyssidae is a feather
mite group occurring most commonly on these birds.
Pteronyssid feather mites belong to the morphotype
specialized for inhabiting primary and secondary
ßight feathers (Mironov 1999, 2001, Dabert and Mi-
ronov 1999). As most feather mite taxa specialized to
this microhabitat, pretonyssid species are commonly
monoxenous or oligoxenous in their host speciÞcity.
Six genera of this family, Conomerus Gaud, 1981,
Neopteronyssus Mironov, 2002, Pteronyssus Robin,
1877, Pterotrogus, Stenopteronyssus Faccini et Atyeo,
1981, and Zygepigynia Gaud et Corpuz-Raros, 1985,
are recently known from woodpeckers (Faccini and
Atyeo 1981, Mironov 2002). With the exception for
Conomerus, which also occurs on barbets (Piciformes:
Capitonidae), these genera are restricted in their host
associations to woodpeckers. Investigations of ptero-
nyssid species diversity and analysis of their host as-
sociations with woodpeckers of the Old World have
been carried out for mites distributed in Eurasia
(Cerny and Schumilo 1973, Mironov 1989, 2003) and
Africa (Gaud 1990, 1991). Pteronyssid mites living on
woodpeckers of the Old World are currently repre-
sented by 27 species of four genera: Conomerus (three
species), Neopteronyssus (16 species), Pteronyssus
(four species), and Zygepigynia (four species). In con-
trast to that, biodiversity of pteronyssids distributed in
the New World is almost unexplored area. Up to date,
only six species of four genera, Neopteronyssus (one
species), Pteronyssus (three species), Pterotrogus (one
species), and Stenopteronyssus (one species) have
been recorded from the New World Picidae (Mc-
Daniel and Price 1963; Faccini and Atyeo 1981).
Among them, two latter genera are apparently re-
stricted to woodpeckers of the Western hemisphere.
Materials and Methods
Mites were found on one of two museum skins of the
ivory-billed woodpecker deposited in the Uebersee
Museum (Bremen, Germany). Mites were extracted
according to the scratching method of Gaud and
Atyeo (1996) and preserved in 70% ethanol. For light
microscopy, mites were macerated in 10% lactic acid
at 60C for 4 d and mounted on slides in Faure medium
(Evans 1992).
The description of the new species follows the stan-
dard approach used for pteronyssid mites (Faccini and
Zoological Institute, Russian Academy of Sciences, Univer-
sitetskaya embankment 1, 199034 Saint Petersburg, Russia(e-mail:
Department of Animal Morphology, A. Mickiewicz University, 28
Czerwca 1956/198, Poznan´61Ð 484, Poland.
Institute of Nature Conservation and Environmental Education,
University of Vechta, Driverstrasse 22, D-49377 Vechta, Germany.
0013-8746/04/0000Ð0000$04.00/0 2004 Entomological Society of America
meadel S6 9/29/04 3:03 Art: AN-04-051 1st disk, 2nd dlw
Atyeo 1981, Mironov 2001, 2002). The nomenclature
of idiosomal chaetotaxy follows GrifÞths et al. (1990)
and the leg chaetotaxy is that of Atyeo and Gaud
(1966). All measurements are given in micrometers.
Analgoidea Trouessart and Me´gnin, 1884
Pteronyssidae Oudemans, 1941
Figs. 1–4. Pterotrogus principalis sp. n., male. (1) Dorsal view. (2) Ventral view. (3). Tarsus III, dorsal view. (4) Tarsus
IV, dorsal view. Signatures for idiosomal setae: c1, c2, c3, cpÑsegment C setae, d1, d2Ñsegment D setae, e1, e2Ñsegment E
setae, f2Ñsegment F setae, gÑgenital setae, h1–h3Ñsegment H setae, ps1–ps3Ñsetae of pseudanal segment, SE, siÑscapular
setae, viÑinternal vertical setae, 1a, 3a, 3b, 4aÑcoxal setae. Signatures for leg setae and solenidia: d, e, fÑdorsoapical setae
of tarsi I-IV, kTÑventral seta of tibiae III, IV, rÑparaxial ventral seta of tarsi III, IV, wÑventral seta of tarsi III, IV,
Ñdorsoapical solenidion of tibiae IÐIV.
meadel S6 9/29/04 3:03 Art: AN-04-051 1st disk, 2nd dlw
Pterotrogus Gaud in Faccini and Atyeo 1981
Until now, the feather mite genus Pterotrogus in-
cluded only the type species, Pterotrogus simplex
(Haller, 1882), which was described from the red-
headed woodpecker, Melanerpes erythrocephalus L.
(1758), in Canada (Haller 1882). This genus can be
distinguished from other pteronyssid genera speciÞc
to woodpeckers by the complete fusion of dorsal
shields of the body into a single uniformly punctured
shield covering almost the entire dorsal surface of the
idiosoma; only the scapular shields may be separated
or traces of their fusion with prodorsal shields are
present (Fig. 1). In a generic revision of the pteron-
yssids, Faccini and Atyeo (1981) had redescribed the
type species and noted that Þve undescribed conge-
neric species are also distributed on the New World
Picidae; however, these authors listed neither species
nor genera of woodpeckers from which they collected
Pterotrogus principalis sp. n.
(Figs. 1Ð7)
Male (Holotype). Length of idiosoma 286, greatest
width of idiosoma 132 (idiosomal size in Þve paratypes
290Ð298 by 136 Ð146). All dorsal shields of the body
fused into single idiosomal shield; traces of fusion
between prodorsal and scapular shields visible (Fig.
1), surface without ornamentation, lateral margins of
prodorsal area of the shield with small incisions ante-
rior to scapular setae, distance between setae SE 84.
Length of hysterosoma from level of setae c2 to lobar
apices 175. Setae c3 lanceolate, with acute apex, 13 in
length. Dorsal setae e1 situated at the level of lateral
Figs 5–7. Pterotrogus principalis sp. n., female. (5) Dorsal view. (6) Ventral view. (7) Head of spermatheca. SD, secondary
meadel S6 9/29/04 3:03 Art: AN-04-051 1st disk, 2nd dlw
setae e2, hysteronotal gland openings gl poorly devel-
oped, setae ps1 awl-shaped, 14 in length. Opisthosoma
wide, approximately two-thirds greatest body width,
with short and widely rounded opisthosomal lobes;
terminal cleft shallowly concave, 12 in length. Dorsal
measurements: c2–d2 88, d2– e2 47, d2– gl 32, gl–e1 25,
e2–e1 4Ð6, e2–h3 42, h2– h2 54, h3– h3 37, ps1–ps1 22,
ps2–ps2 66. Inner ends of epimerites IIIa form an
oblique T; epiandrium absent. Setae 3a anterior to 3b,
situated on sclerotized inner ends of epimerites IIIa
(Fig. 2). Genital arch 15 in length, 12 in width, ae-
deagus one-half as long as arch, genital setae g, and
coxal setae 4a at midlevel of genital arch. Adanal
apodemes, adanal shields, and adanal membranes ab-
sent. Tarsus III with apical claw, 27 in length; seta r
short, 7 in length (Fig. 3). Tarsus IV without dor-
sobasal teeth; modiÞed seta dand econical, weakly
sclerotized (Fig. 4).
Female (Paratype). Length of idiosoma 455, width
of idiosoma 164 (idiosomal size in other four paratypes
445Ð450 by 162Ð178). Structure of entire dorsal idio-
somal shield as in the male (Fig. 5), setae SE separated
by 98. Length of hysterosoma from level of setae c2 to
posterior end 322. Setae c3 lanceolate with acute apex,
20 in length. Posterior margin of opisthosoma
rounded, without lobes. Setae d1 at level of humeral
setae cp, setae e1 posterior to level of setae e2. Setae
ps1 thick setiform, 46 in length. External copulatory
tube as small weakly sclerotized terminal extension,
5 in length. Dorsal measurements: c2–d2 133, d2–e2
84, e2–f2 90, e2–e1 24 Ð30, h2–h2 69, ps1–ps1 24. Epigy-
nium a long arch, 34 in length, 25 in width; sclerotized
fold of oviporus long, extending beyond the midlevel
of epimerites IV (Fig. 6). Head of spermatheca as in
Fig. 7.
Type Material. HOLOTYPE: one male (ZISP BR
145) from the Ivory-billed Woodpecker Campephilus
principalis (Picidae), female, North America, no other
data (UBR No 6165); PARATYPES: Þve males and Þve
females, same data. Holotype male, paratype 2 males
and two femalesÑZoological Institute, Russian Acad-
emy of Sciences, Saint Petersburg, Russia (ZISP);
paratype one male and two femalesÑA. Mickiewicz
University, Poznan´, Poland (AMU); paratype one
male and one femaleÑUebersee Museum, Bremen,
Germany (UBR).
Diagnosis. The males of the new species differ from
those of the type species, Pterotrogus simplex, by hav-
ing setae e1 situated approximately at level of setae e2,
widely rounded opisthosomal lobes, legs IV not ex-
tending beyond lobar apices, and setae 3a situated on
sclerotized areas of epimerites IIIa; the females are
distinguished by having setae e1 clearly posterior to
the level of setae e2, legs IV not reaching the posterior
end of opisthosoma, and setae ps1 twice as long as the
distance between their bases. Males of P. simplex have
setae e1 situated between the levels of setae d2 and e2
and commonly equidistant from them, opisthosomal
lobes obtusely angular, legs IV extending beyond the
lobar apices by the length of an ambulacral disc, and
setae 3a are off the sclerotized areas of epimerites IIIa.
Females have setae e1 equidistant from the levels of
setae d2 and e2, legs IV extending beyond the posterior
end of opisthosoma by distal half of tarsus, setae ps1
having length approximately one-third distance be-
tween their bases.
Etymology. The speciÞc epithet derives from the
speciÞc name of the host.
The authors thank P.-R. Becker, head of the Natural His-
tory Department in the U
¨bersee Museum (Bremen, Ger-
many), for making the ornithological collection available for
our study, and H. C. Proctor (University of Alberta, Canada)
for critically reviewing the manuscript. This study was mainly
supported by the Committee of Science Promotion of the
University of Vechta (KFN), Germany, and in part, by the
Russian Foundation for Basic Researches (Project 03-04-
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Received 13 April 2004; accepted 15 July 2004.
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... This genus is one of 12 pteronyssid genera restricted in their host associations to birds of the order Piciformes . To date, the genus Pterotrogus has included 17 species; all of them are associated with woodpeckers of the New World (Mironov 2005;Mironov et al. 2005;Hernandes 2012). Phylogenetic analysis of relationships within this genus, based on morphological characters, and a key to species were provided by Mironov (2005). ...
... The mite genus Pterotrogus is widely distributed on New World woodpeckers, particularly the species-rich genera Campephilus Gray, Colaptes Vigors and Veniliornis Bonaparte in Central and South America. A few species of Pterotrogus species are also known from single host species from the genera Dryocopus Boie, Leuconotopicus, and Melanerpes Swainson in North America (Mironov 2005;Mironov et al. 2005;Hernandes 2012). We suggest that the ancestor of P. panamensis has shifted from a woodpecker species of the gen-era Colaptes or Veniliornis harboring species of the simplex group (Mironov 2005). ...
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... The diversity of the family Pteronyssidae has been most extensively studied in the Old World, particularly in Europe (erny and Schumilo 1973;Mironov 1985Mironov , 1989 and Africa (Gaud 1988(Gaud , 1989(Gaud , 1990a(Gaud , 1990b(Gaud , 1991. Representatives of this family are extremely poorly known in North America; only seven species belonging to the genera Pteronyssus (two species), Neopteronyssus Mironov, 2002 (one species), Pterotrogus Faccini and Atyeo, 1981 (two species), and Scutulanyssus (two species) have been recorded from the Nearctic region (Tyrrell 1882;Hall 1912;Banks 1915;McDaniel and Price 1963;Bruce and Johnston 1969;Faccini and Atyeo 1981;Mironov 2003;Mironov et al. 2005). ...
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... Six feather mite species were described from the museum exhibit of the Carolina parakeet Conuropsis carolinensis, the only North American parrot, 100 years after this bird became extinct through the destruction of its forest habitat (Mironov et al. 2005a). Also, the recently described new Pterotrogus (Analgoidea, Pteronyssidae) species was discovered on the museum skin of the ivory-billed woodpecker Campephilus principalis, a bird still believed by some ornithologists to exist despite the last reliable sightings of this species being in the 1950s in North America and probably in the early 1990s in Cuba (Mironov et al. 2005b). The well-known example of the extinct Diplaegidia gladiator (Analgoidea, Analgidae) from the passenger pigeon Ectopistes migratorius should not be included in this list of discovered ''museum fossils'' because it was described when the passenger pigeon was still alive (Haller 1882). ...
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A systematic review of pterolichid feather mites of the Psittophagus generic group (Pterolichidae, Pterolichinae) is presented. New feather mite taxa are described: Psittaculobius quadriglobus gen. nov., sp. nov. from the Long-tailed Parakeet Psittacula longicauda (type host) and the Red-breasted Parakeet P. alexandri (Psittacidae, Psittacinae), Psittophagus calyptorhynchi sp. nov. from the Yellow-tailed Black Cockatoo Calyptorhynchus funereus (Cacatuidae, Calyptorhynchinae). A key to all genera and species of the Psittophagus group and taxonomic comments on formerly described species are provided, and host-parasite associations of these mites with the parrots from the Old World are briefly discussed. It is hypothesised that the Psittophagus group originated on the ancestors of Psittaciformes at the same time as other pterolichine groups specific to parrots, but in the process of coevolution with these hosts it has achieved significant success on Cacatuidae only, while on Psittacidae, the representatives of this group have been retained only on few phylogenetic lineages in the Old World.
Eight species of the feather mite genus Anephippius (Avenzoariidae, Pteronyssinae) are associated with African Piciformes. Five species are parasitic on Capitonidae, 3 species on Picidae. -from English summary
The genus Pegopteronyssus seems to be restricted to Africa and includes only P. phyllurus, a parasite of the capitonid genera Gymnobucco, Lybius, Smilorhis, Stactolaema and Tricholaema. The genus Pteronyssus is ubiquitous. Four African species assigned to this genus are very closely related, forming a species complex. All of them parasitize Picidae of the genus Campethera. -from English summary
Pteronyssus centurus is described from the goldenfronted woodpecker, Centurus aurifrons, and the ladderbacked woodpecker, Dendrocopos scalaris. It is closely related to P. glossifer Gaud but has a differently shaped propodosomal shield in both sexes, and the hysterosomal plate of the female is composed of two sections rather than three as in P. glossifer. The three lots of material on which the description is based were collected in Kleberg and Hidalgo Counties in South Texas, and in Palo Pinto County in the north-central part of the State.
Within the Astigmata, setal homologies with the general chaetotaxic systems developed for acariform mites by F. Grandjean have never been convincingly established. This study deals with all body regions, exclusive of legs and gnathosoma, mostly utilizing as models astigmatid species from three different families. Six hypotheses which attempt to explain the ontogeny of segments, setae and cupules in the caudal bend region are compared using three-dimensional views obtained by scanning electron microscopy. The hypotheses are evaluated by in-group comparisons testing their consistency with presumed segmental boundaries, and by out-group comparisons with oribatid mites. The strongest, most parsimonious hypothesis suggests that larval astigmatid mites possess segment F, but not its setae, and the two pairs posterior to segment F are hl and h2, whilst the five setae added in the protonymph are h3, f2 and ps1–3. Six pairs of structures on the deutonymphal anal plate are identified as modified setae, and evidence supporting their homologies with setae of other instars is discussed for the first time. A comparison of the principal chaetotaxic systems used for the dorsal and anal regions of the Astigmata is presented. Application of Grandjean's setal signatures to these regions, as well as to the coxisternal and genital regions, is discussed and illustrated by examples.