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Feather Mites Associated with Eastern Bluebirds (Sialia sialis L.) in Georgia, Including the Description of a New Species of Trouessartia (Analgoidea: Trouessartiidae)
Reneé E. Carleton and Heather C. Proctor

Southeastern Naturalist, Volume 9, Issue 3 (2010): 605–623

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2010 SOUTHEASTERN NATURALIST 9(3):605–623 Feather Mites Associated with Eastern Bluebirds (Sialia sialis L.) in Georgia, Including the Description of a New Species of Trouessartia (Analgoidea: Trouessartiidae) Reneé E. Carleton1,* and Heather C. Proctor2 Abstract - Eastern Bluebirds inhabiting a grass-dominated agricultural environment within a northwestern Georgia land tract were examined over the course of three breeding seasons (2004 through 2006) to assess the presence of ectosymbionts. More than 90% of bluebirds examined harbored plumicolous feather mites of four species: Pterodectes sialiarum (Proctophyllodidae), Mesalgoides sp. (Psoroptoididae), Analges sp. (Analgidae), and a previously undescribed Trouessartia sp. The recovery of P. sialiarum represented the second report of this species, which had previously been recorded from Eastern Bluebirds in Guatemala. New host records for Mesalgoides sp. and Analges sp., and a description of Trouessartia sialiae sp. nov. also resulted from the study. Mite abundance did not vary among groups of birds categorized by subjective quantification, with the exception of a group of a few individuals harboring a vast number of mites. Abundance was not correlated with mean host body mass or body condition and was also independent of host sex. Feather mites were most commonly found on primary remiges, occasionally on secondary remiges, and rarely on rectrices; each mite species was located on a specific type of feather. Lice were also occasionally recovered, but were reported separately. Introduction Astigmatan feather mites are common symbionts of nearly every avian family. Members of this highly diverse assemblage (superfamilies Analgoidea, Freyanoidea, and Pterolichoidea) occur worldwide and include more than 2000 described species, with at least that many awaiting description (Dabert and Mironov 1999, Gaud and Atyeo 1996, Proctor 2003). Feather mites that dwell on the vanes of flight feathers are considered non-parasitic based on evidence that they feed on uropygial secretions, pollen, and possibly accumulations of microorganisms rather than host tissues (Blanco and Frias 2001, Blanco et al. 1997, Proctor 2003, Proctor and Owens 2000). Host specificity varies from a single host species per mite species to multiple host species from several families; however, most well-studied feather mites appear to be restricted to hosts within a single genus or family (H.C. Proctor, pers. observ.). Means of dispersal and colonization of new hosts by feather mites is unclear. Direct contact between conspecifics is the most likely method of transfer between hosts (Proctor 2003). A form of horizontal transfer involving hippoboscid flies may occur occasionally (Philips and Fain 1991), 1Department of Biology, School of Mathematical and Natural Sciences, Berry College, 2277 Martha Berry Highway NW, Mount Berry, GA 30149. 2Department of Biological Sciences, University of Alberta, Edmonton, AB, T6G 2E9 Canada. *Corresponding author - 606 Southeastern Naturalist Vol. 9, No. 3 although one recent study found no instances of phoresy by feather-dwelling mites on hippoboscids (Jovani et al. 2001). Vane-dwelling feather mites are highly adapted to aerodynamic stresses associated with specific flight feathers, although members of a given mite species do vary their locations within and among feathers depending on host activity, feather age, and ambient temperature (Dubinin 1951, Jovani and Serrano 2004, Proctor 2003). Although there is little evidence that vane-dwelling mites negatively affect their hosts, feather mite-host relationships have been frequently studied to determine effects of parasites on host condition and fitness (Behnke et al. 1995, Figuerola 2000, Pérez-Tris et al. 2002, Rózsa 1997). Some studies suggest feather mite abundance is positively correlated with the presence of viral lesions (Thompson et al. 1997) and negatively correlated with amount of pectoral muscle mass (Harper 1999). If this is the case, then even if feather mites are not directly affecting host condition, their presence and abundance on hosts may provide clues about the health status of many avian populations. We observed numerous feather mites during a banding study of a Sialia sialis L. (Eastern Bluebird) population nesting on a grass-dominated agricultural site in Georgia. Only three species of feather mites, Pterodectes sialiarum (Stoll 1893) (Proctophyllodidae), Proctophyllodes vesca Atyeo and Braasch (1966) (Proctophyllodidae), and Trouessartia sp. (Trouessartiidae) (Forrester and Spalding 2003; J. Mertins, USDA, APHIS, National Veterinary Services Laboratory, Ames, IA, pers. comm.) have been previously reported from Eastern Bluebirds. Objectives of this study were to identify mite species associated with the population, estimate mite prevalence, and make preliminary observations of ecological relationships between mite abundance and host condition. Study Area The study was conducted on a land tract in Floyd County (34.282799°N, 85.191803°W) that lies within the Ridge and Valley physiographic region of Georgia. The study area consisted of three non-contiguous sites of grass-dominated agricultural management: a Cynodon dactylon (L.) Pers. (Bermuda Grass)-dominated field managed for hay production, a cattle pasture with Festuca sp. (perennial fescue) and mixed low-growing herbaceous plants, and a low-management field with features of early succession. Methods We conducted the study during the breeding seasons (March through August) of 2004, 2005, 2006, 2008, and 2009. Adult Eastern Bluebirds (n = 228) were captured within nest boxes using a trap-door apparatus. Captures were attempted within five days after a clutch hatched and between 07:00 am–12:00 pm. Adults were banded for identification purposes with individually numbered United States Geological Survey aluminum leg bands on initial capture. Sex of each individual was recorded. Birds were weighed to the nearest 0.5 g using a spring scale and cloth bag, and the amount of visible subcutaneous fat in the depression between the furcula (SFS) was scored as no fat (0) or fat present (1) (Krementz and Pendleton 1990). A subjective 2010 R.E. Carleton and H.C. Proctor 607 measurement of pectoral muscle mass (PMS) was scored as emaciated (E), thin (T), muscled (M), well-muscled (W), or robust (R) (Graham 1993). Feather mites were detected by visual examination (see McClure 1989) aided by manual ruffling of feathers and extension of both wings to allow maximum exposure of flight feathers (remiges). Tail feathers (rectrices) were also examined for mites. Feather mite presence, specific feathers inhabited on each wing and the tail, and a subjective estimation of mite abundance were recorded for all birds. The small size of the mites and the need to minimize handling time prevented quantitative measurement of mite numbers. We subjectively defined abundance as very abundant (5), abundant (4), moderately present (3), rare (2), and very rare (1) based on the number of feathers inhabited and distribution of mites on individual wing feathers (Fig. 1). Representative feather sections harboring mites were snipped from 14 adults during 2004 and preserved in 70% ethanol (EtOH) for mite identification. Figure 1. Illustration depicting feather mites on Eastern Bluebird primary remiges corresponding to abundance scores: 1 = very rare, 2 = rare, 3 = moderate, 4 = abundant, 5 = very abundant. 608 Southeastern Naturalist Vol. 9, No. 3 Fourteen male bluebirds were sacrificed during the latter six weeks of the 2006 season for necropsy and quantification of feather mite loads. Each carcass was placed in a sealed plastic bag and refrigerated before post-mortem examination. Following the method described by Clayton and Walther (1997), a solution of water, 10% isopropyl alcohol, and household detergent was added to each bag. The contents were agitated for 5 min to dislodge ectosymbionts. After the solution was decanted into a large beaker, the carcass was removed and both the plastic bag and carcass were rinsed with clean water to remove any remaining fauna. Arthropods were recovered by filtering wash and rinse solutions. Each filter paper was examined using a dissecting microscope, and recovered specimens were placed into vials containing 70% EtOH. Mite-bearing Eastern Bluebird feathers that were snipped in 2004 were soaked in 80% EtOH to hydrate and preserve the mites. One of us (H.C. Proctor) examined feathers and the preservative under a dissecting microscope and removed the mites. Exemplars of each morphotype were cleared in lactic acid for a minimum of 12 hours, slide-mounted using commercially available poly-vinyl alcohol (PVA) (BioQuip Products, Rancho Dominguez, CA), and then cured for 4 days at ca. 40 °C on a slide-warmer. Mites were identified using a Leica DMLB compound microscope with differential interference contrast (DIC) lighting and the following literature: Gaud and Atyeo (1996), Santana (1976), and Stoll (1893). Mites from four genera were identified: Mesalgoides (Psoroptoididae), Analges (Analgidae), Pterodectes (Proctophyllodidae), and Trouessartia (Trouessartiidae). Exemplars of each taxon are deposited in the E.H. Strickland Entomological Museum, Department of Biological Sciences, University of Alberta, Edmonton, AB, Canada (UASM) Specimens of Pterodectes sialiarum (Stoll) (Proctophyllodidae) were sent to experts in Brazil for confirmation of their identity (Valim and Hernandes 2008). Additional specimens of P. sialiarum are deposited in the collection of Acari of the Departamento de Zoologia et Botânica da Universidade Estadual Paulista, São Paulo, Brazil. The Trouessartia proved to be a new species, which we describe here. Pencil drawings were made by H.C. Proctor of a male and a female mite at 400x using a 1x drawing tube attached to a Leica DMLB compound microscope with differential interference contrast (DIC) lighting. Measurements were taken at 200x and 400x using a calibrated ocular micrometer. The drawings were digitized using a flatbed scanner and then traced in Adobe Illustrator CS version 11.0.0 (Adobe Systems Incorporated, 1987–2003). A Canon Power- Shot S40 digital camera was used to photograph some aspects of morphology. The species description incorporates the formats used by Santana (1976), Mironov and Kopij (2000), and OConnor et al. (2005). Ten adult mites of each sex were measured; all measurements are in micrometers (μm). It is possible that the Mesalgoides and Analges recovered from the Eastern Bluebird samples are also species new to science; however, the taxonomy of these two genera is in too poor a state to determine this without great effort. JMP 8.0 Statistical Software (SAS Institute, Cary, NC) was used for all statistical analysis; a priori significance was set at α = 0.05. Pearson χ2 test of independence was used to evaluate relationships between subjective mite 2010 R.E. Carleton and H.C. Proctor 609 abundance categories and host sex, body condition scores, and subcutaneous fat scores. One-way ANOVA was used to test for differences in body mass among mite abundance categories. Results Feather mites were noted on 223 of 228 (97.8%) adult bluebirds, and when present, were always observed on primary remiges (100%), often on secondary remiges (56.1%), and occasionally on rectrices (10.7%). Mite abundance scores were somewhat normally distributed among the categories “abundant” (13.6%), “moderate” (38.5%), “rare” (22.9%), or “very rare” (17.9%). Abundance on eleven birds (4.8%) was scored as “very abundant” (Fig. 2). No feather damage was noted on areas harboring mites. Philopterus sialii (Osborn) (Lice) were occasionally recovered, but are reported elsewhere (Carleton et al., in press). Mites were located most commonly on the right and left eighth, ninth, and tenth primaries. Most birds (86.0%) had mites on three to five remiges, and 30.7% had mites on only three primaries of each wing. Mites located on secondary remiges were typically on the third through sixth and tenth and eleventh secondaries, and most commonly on the fourth or fifth secondaries. Although mites were found on both wings of most birds, a small number of birds (2.2%) had mites only on the right wing. These birds were included in the “very rare” category of mite loads. Mites located on primaries were attached to the ventral surface of barbs and oriented anteriorly toward the shaft, whereas mites located on secondary remiges and rectrices were only found on the dorsal surface. Mites on primaries were always located within the middle third of each feather, while those on secondaries or rectrices were always located within the proximal third of the feather and covered by the greater secondary coverts or upper tail coverts. There was no difference in mean host body mass (mean = 29.7 g, SD ± 0.95) among birds that differed in mite abundance levels (F5,227 = 0.5390, Figure 2. Histogram of feather mite abundance scores among a sample of Eastern Bluebirds (n = 228) nesting in northwestern Georgia (2004–2006, 2008–2009) with percentages of each score. Abundance scores: 0 = none, 1 = very rare, 2 = rare, 3 = moderate, 4 = abundant, 5 = very abundant. 610 Southeastern Naturalist Vol. 9, No. 3 P = 0.746; Fig. 3). Mite abundance was independent of host sex (Pearson χ2 = 8.805, 5 df, P = 0.117) and was not associated with host condition as estimated by subcutaneous fat score (Pearson χ2 = 7.848, 5 df, P = 0.165; Fig. 4). There was some evidence of an association between mite abundance and pectoral muscle score (Pearson χ2 = 31.435, 15 df, P = 0.0074; Fig. 5). Four species of feather mites were identified from samples collected by live examinations and body washes of the 14 necropsied birds (Table 1). Pterodectes sialiarum was the only species associated with the primary remiges that were collected during live examinations. A Mesalgoides sp. and a previously unidentified Trouessartia sp. (described below as T. sialiae sp. nov.) were the only species associated with collected secondary feathers. Trouessartia spp. are unusual among feather mites because they often occupy the dorsal rather than ventral surface of flight feathers (Dubinin 1951). The Trouessartia sp. was also the only species of feather mite found on rectrices. Pterodectes sialiarum, Mesalgoides sp., Trouessartia sp., and Analges sp. were recovered from wash samples. Recovery of the Analges sp. only by washing is not Figure 3. Box plots of Eastern Bluebird body mass (g) by feather mite abundance scores. Abundance scores: 0 = none, 1 = very rare, 2 = rare, 3 = moderate, 4 = abundant, 5 = very abundant. Figure 4. Histograms showing categories of subcutaneous fat (SFS) located between the furcula of Eastern Bluebirds (n = 228) by feather mite abundance scores with percentages. SFS 0: subcutaneous fat absent; SFS 1: subcutaneous fat present. Abundance scores: 0 = none, 1 = very rare, 2 = rare, 3 = moderate, 4 = abundant, 5 = very abundant. 2010 R.E. Carleton and H.C. Proctor 611 unexpected because members of this genus are typically associated with body feathers rather than flight feathers (Dubinin 1951). Taxonomy Family Trouessartiidae Gaud 1957 Genus Trouessartia Canestrini 1899 Trouessartia sialiae sp. nov. (Figs. 6–10) Holotype: male, 19 June 2004, Mount Berry, GA (34.282799oN, 85.191803oW), from Sialia sialis (Linneaus 1758) (Eastern Bluebird). Coll. Figure 5. Histograms showing categories of pectoral muscle mass scores (PMS) by feather mite abundance scores with percentages from a sample of Eastern Bluebirds (n = 228) nesting in northwestern Georgia (2004–2006, 2008–2009). Pectoral muscle mass scores: W = well muscled, M = muscled, T = thin. Abundance scores: 0 = none, 1 = very rare, 2 = rare, 3 = moderate, 4 = abundant, 5 = very abundant. Table 1. Feather mites collected from Sialia sialis (Eastern Bluebird) by body wash prior to necropsy (n = 14) and from feathers collected during live examination (n = 14) during 2004–2006 and 2008–2009. Prevalence (%) / range, Presence (X) on collected feathers Mite species by body wash Primaries Secondaries Rectrices Pterodectes sialiarum 100.0 / 5–226 X Trouessartia sialiae (sp. nov.) 85.0 / 0–19 X X Mesalgoides sp. 35.7 / 0–11 X Analges sp. 28.3 / 0–30 612 Southeastern Naturalist Vol. 9, No. 3 R. Carleton. Deposited in the Museum of Zoology, University of Michigan, Ann Arbor, Michigan (UMMZ). Paratypes: 9 males, 10 females, same location and host species, dates May– July 2004. Coll. R. Carleton. 2 male and 2 female paratypes deposited in the UMMZ, 7 male and 8 female paratypes in the University of Alberta E.H. Strickland Museum of Entomology, Edmonton, AB, Canada (UASM). Male (holotype, range given for 9 paratypes unless otherwise mentioned; Figs. 6, 7, 10). Dorsum (Fig. 6). Length of idiosoma excluding terminal Figure 6. Trouessartia sialiae sp. nov., male: dorsal view; gla = gland opening. 2010 R.E. Carleton and H.C. Proctor 613 lamella 503 (485–520), width of idiosoma as measured between bases of setae c2 210 (195–213, from 8 paratypes). Length of prodorsal shield 153 (148–160), width of shield as measured at level of setae se 170 (163–178); with narrow lateral extensions between bases of legs I, II; surface uniformly granular, without lacunae. Setae si narrowly lanceolate, 30 in length (28–30, from 7 paratypes); bases separated by 65 (65–73). Humeral shield with setae c2 narrowly lanceolate, 48 in length (45–53). Setae c3 lanceolate, 23 in length (23–25). Dorsal hysterosoma with unsclerotized gap between prohysteronotal shield and lobar shield anterior to setae e2; gap broadest at lateral margins of Figure 7. Trouessartia sialiae sp. nov., male: ventral view; TLA = translobar apodeme. 614 Southeastern Naturalist Vol. 9, No. 3 Figure 8. Trouessartia sialiae sp. nov., female: dorsal view; gla = gland opening. shields, and very narrow medially. Length of prohysteronotal shield 220 (200– 222); maximum width 185 (170–188); surface granular and without lacunae; lateral margins with heavily sclerotized notches near level of setae d2 (Fig. 6); 2010 R.E. Carleton and H.C. Proctor 615 Figure 9. Trouessartia sialiae sp. nov., female: ventral view. 616 Southeastern Naturalist Vol. 9, No. 3 d2 present as small hairlike setae, all other prohysteronotal setae represented only by alveoli. Length of lobar shield along midline, excluding lamellae, 90 (88–98). Terminal lamellae leaflike, tapered to blunt points distally, margins entire, faint longitudinal striations present. Distance from base of free terminal cleft to lamellar apices 53 (48–58). Setae ps2 and h1 setiform; bases of h1 anterior to bases of h2; ps1 represented only by alveoli. Venter (Fig. 7). Epimerites I free. Rudiments of epimerites IIa small, circular. Setae sR of trochanters III lanceolate with acute tips, length 14 (10–15). Genital apparatus massive (Fig. 10a), situated between levels of trochanters III, IV; length 80 (75–90), width at base anterior to setae g 48 (40–53); small ovoid pregenital apodeme present. Anterior and posterior genital papillae equidistant from midline. Bases of setae g clearly separated, distance between them 10 (9–12). Translobar apodeme present (TLA, Fig. 7). Setae ps3 on small triangular platelets anterolateral to adanal discs. Button-like setae d, e on tarsus IV separated by 3 (2–4), a distance approximately equal to width of one of these setae. Female (range for 10 paratypes unless otherwise mentioned). Dorsum (Figs. 8, 10b). Length of idiosoma including terminal lamellae 575–628; width of idiosoma as measured between bases of setae c2 203–225 (from 9 paratypes). Prodorsal shield as in male, 153–170 in length, 173–190 in width as measured at level of setae se; setae si narrowly lanceolate, 28–30 in length, bases separated by 73–83. Humeral shield with setae c2 narrowly lanceolate, 45–53 in length. Setae c3 lanceolate, 23–25 in length. Hysteronotal shield length to base of terminal cleft 253–270; maximum width 173–195; numerous ovoid lacunae mostly restricted to the middle third of the shield between setal bases (Figs. 8, 10b); lateral margins with heavily sclerotized notches near level of setae d2; d2 present as small hairlike setae, all other hysteronotal setae anterior to h1 represented only by alveoli. Setae Figure 10. Trouessartia sialiae sp. nov., photographs of details of morphology: (a) male genital capsule, (b) female dorsal shield sculpture. 2010 R.E. Carleton and H.C. Proctor 617 h1 robust and lanceolate, 18–28 long (depending on angle of observation), bases separated by 58–65; direct distance from bases of h1 to bases of h2 28–35; direct distance from bases of h1 to nearest lateral margin of hysteronotal shield 20–25. Width of opisthosoma at level of setae h2 115–128. Setae ps1 minute, hairlike, visible only under oil; positioned dorsally 14–19 anterior to bases of setae h3. Distance from bases of h2 to apices of lobar lamellae 110–135. Setae f2 not apparent. Supranal concavity open posteriorly. Length of terminal cleft 118–145, width of cleft at level of setae h3 38–55. Venter (Fig. 9). Epimerites I free. Setae sR on trochanter III lanceolate and 15–18 in length. Interlobar membrane very reduced, apparently restricted to area of supranal concavity rather than extending between lobes (Fig. 8b). Primary spermaduct terminating prior to interlobar region, exiting ventrally on small membranous protuberance (Fig. 10). Length of secondary spermathecal ducts 25–34 (from 8 paratypes). Etymology. The specific epithet is derived from the generic name of the host. Differential diagnosis. Trouessartia sialiae sp. nov. displays an unusual combination of characters in which males have leaf-shaped, apically pointed lamella with complete edges and females lack a tubular external spermaduct. Other described species with these features include T. bifurcata (Trouessart), T. carpi Till, T. rubecula Jabłonska, T. simillima Gaud, T. subacuta Gaud & Mouchet, T. swidwiensis Jabłonska, T. trouessarti Oudemans, and T. unicolor Berla. With the exception of T. unicolor, males of these other species have small, narrow genital apparati in which the width at the base is at most 1/3 the length and the bases of setae g are apposed. In T. sialiae sp. nov., the width of the genital apparatus is almost half the length, and setae g are clearly separated. Trouessartia sialiae sp. nov. can be differentiated from T. unicolor by the following features: setae c2 narrowly lanceolate in both sexes of T. sialiae (with filiform tips in both sexes of T. unicolor); setae h3 with long filiform tips in males (lack filiform extensions in T. unicolor); prohysteronotal and lobar shields completely separate in males (partially fused in T. unicolor); setae g in males clearly posterior to 4a (slightly anterior to 4a in T. unicolor); anterior and posterior genital papillae in males same distance from midline (anterior pair further from midline than posterior in T. unicolor); setae h1 in females robust, long, tips reaching to bases of h2 (narrow and not reaching h2 in T. unicolor) (see illustrations in Santana 1976). Trouessartia unicolor has been collected from Haplospiza unicolor Cabanis (Emberizidae) in South America (Santana 1976). Discussion There have been few reports of feather mites associated with Eastern Bluebirds despite a large volume of research on this species. The recovery of Pterodectes sialiarum represents a new host distribution record and is only the second report since the original discovery in Guatemala over a century ago (Stoll 1893). The northern-most distribution of P. sialiarum is probably not limited to the southeastern United States and Central America, given 618 Southeastern Naturalist Vol. 9, No. 3 that migration patterns of the more northern Eastern Bluebird populations are highly variable and appear to fluctuate with degree of winter severity (Gowaty and Plissner 1998). Recovery records from Michigan indicated some individuals migrating southward passed over areas with winter residents (Pinkowski 1971). Bird Banding Laboratory data reported the farthest migration occurred between southwestern Manitoba in Canada and Texas (Gowaty and Plissner 1998). Many bluebirds banded during the course of the study have been observed during winter months (R.E. Carleton, pers. observ.), suggesting that this population is non-migratory. The extent to which migratory bluebirds overwinter in the area and interact with residents is not known. The other two Sialia species, S. currucoides (Bechstein) (Mountain Bluebird) and S. mexicana Swainson (Western Bluebird), have been examined for feather mites but have yielded only Proctophyllodes spp. (Proctophyllodidae) (Atyeo and Braasch 1966; H.C. Proctor, unpubl. data for S. currucoides in Alberta, Canada). Our generic records of the unidentified Mesalgloides sp. and Analges sp. are the first for the genus Sialia. We recovered these species of feather mites in relatively low numbers and at a lower prevalence than the other two species. Recovery of Mesalgoides from collected secondary feathers represents an unusual within-host distribution because members of this genus display morphological adaptations for living in down feathers rather than flight feathers (Dabert and Mironov 1999). Analges sp. and Mesalgoides sp. are commonly found on several bird species that inhabit our study area; these include Cardinalis cardinalis (L.) (Northern Cardinal) (Wilson and Durden 2003), Sturnus vulgaris L. (European Starling) (Boyd 1951, Mitchell and Turner 1969), Turdus migratorius L. (American Robin) (Threlfall and Wheeler 1986), and Tachycineta bicolor (Vieillot) (Tree Swallow) (Lombardo and Thorpe 2000, Shutler et al. 2004). As obligate cavity-nesters, bluebirds, starlings, and swallows may occupy old nesting cavities previously used by non-conspecifics. Our nest-box design excluded starlings, but Tree Swallows occasionally usurped bluebirds from nest boxes. Although it is possible that transfer among host species occurs during use of old nesting cavities, there are no documented cases of this, and we consider mite movement between bluebirds and other species using nest boxes to be unlikely. Trouessartia sialiae sp. nov. represents a previously undescribed species of feather mite and is the first species in this genus to be recorded from the genus Sialia. Although there is a report of a Trouessartia sp. recovered from an Eastern Bluebird in Florida (J. Mertins, pers. comm.; Forrester and Spalding 2003), no formal description of this specimen was made. Trouessartia is one of the most species-rich genera of feather mites with approximately 100 named species and at least as many as yet undescribed. Trouessartia species are restricted to passeriform hosts (Mironov and Kopij 2000). Other genera of the family Turdidae that have had Trouessartia collected from them include Alethe, Brachypteryx, Catharus, Neocossyphus, Turdus, and Zoothera (Table 2). The locations of P. sialiarum and T. sialiae sp. nov. on specific types of feathers appeared to be consistent among individual bluebirds and was not influenced by relative mite abundance. Whether or not these species 2010 R.E. Carleton and H.C. Proctor 619 Table 2. Records of Trouessartia species from members of the family Turdidae. Host genus Host species Author Trouessartia sp. Author Location Reference Alethe castanea (Cassin) longifolia Gaud & Mouchet Cameroon Zumpt (1961) Brachypteryx leucophris (Temminck) Unidentified sp. Asia McClure et al. (1973) montana Horsfield Unidentified sp. Asia McClure et al. (1973) Catharus minimus (Lafresnaye) Unidentified sp. Newfoundland Wheeler and Threlfall (1986) ustulatus (Nuttall) Unidentified sp. Newfoundland Wheeler and Threlfall (1986) ustulatus (Nuttall) Unidentified sp. Alaska Wilson and Haas (1980) Neocossyphus fraseri (Strickland) stizorhinae Gaud and Mouchet Cameroon, Congo, Gabon Zumpt (1961), Santana (1976) Sialia sialis (L.) sialiae sp. nov. Proctor and Carleton Georgia This paper sialis (L.) Unidentified sp. Florida Forrester and Spalding (2003) Turdus albicollis Vieillot mangaratibensis Berla Brazil Santana (1976) albicollis Vieillot serrana Berla Argentina, Brazil, Surinam Santana (1976) chrysolaus Temminck Unidentified sp. Asia McClure et al. (1973) libonyanus (Smith) incisa Gaud Cameroon Gaud and Mouchet (1958) merula L. corvina* (Koch) no location given Santana (1976) merula L. incisa Gaud England, Morocco Santana (1976) obscurus Gmelin Unidentified sp. Asia McClure et al. (1973) pallidus Gmelin Unidentified sp. Asia McClure et al. (1973) poliocephalus Latham Unidentified sp. New Hebrides Marshall (1976) Zoothera citrina (L.) Unidentified sp. Asia McClure et al. (1973) dauma (Latham) Unidentified sp. Asia McClure et al. (1973) sibirica (Pallas) Unidentified sp. Asia McClure et al. (1973) *Santana (1976) considers this mite-host record to be questionable. 620 Southeastern Naturalist Vol. 9, No. 3 invariably remain in place or occasionally move to other feathers or body areas is unknown. Feather mites have been shown to avoid moulting feathers (Jovani and Serrano 2001, Pap et al. 2006) and to migrate from tertiary to primary and secondary feathers during seasonal and daily ambient temperature change (Wiles et al. 2000). During the present study, all examinations and collections occurred only within morning hours. We found no association between feather mite abundance scores and body mass or amount of furcular fat in the population under study. Although there was statistical evidence of an association between mite abundance and pectoral muscle mass, we suspect that this was not actually the case. Only a very small portion of the population was scored as emaciated (0.44%) or well-muscled (7.9%); no birds were scored as robust. Percentage-wise, more birds that were scored as “well-muscled” were also scored as “very abundant” in mite abundance compared to birds that were scored as “emaciated”, “thin”, or “muscled”. However, as many “well-muscled” birds were scored as “moderate” in mite abundance as those that were scored as “very abundant”. The single emaciated bird had very few mites. Previous studies of the relationships between host condition and mite load have returned contradictory findings (summarized in Proctor 2003). For example, Rózsa (1997) reported a positive relationship between mite abundance and body mass, while Poulin (1991) and Figuerola (2000) found no such relationship. Relative abundance of each mite species recovered could not be estimated by visual examination because the method only allowed recognition of mite presence per se, rather than identification of individual species (Mc- Clure 1989). Based on results obtained by body washing and examination of feather samples, P. sialiarum and T. sialiae were common throughout the population; however, estimates based on small sample sizes, as in the case of our sample obtained during necropsy (n = 14 birds, all male), may be suspect (Gregory and Blackburn 1991). Our findings yielded additional information about feather mite distribution and the relationship between mite abundance and host condition. Manipulation of mite abundances among hosts, such as by removal and addition, would allow better evaluation of relationships between host condition and mite abundance, and would also allow experimental determination of whether “misplaced” mites move to colonize the feathers on which they are typically found Authors’ note: Just before publication of this paper, we learned that Pterodectes sialiarum had been moved to Amerodectes sialiarum (Valim and Hernandes 2010). Acknowledgments This study was supported by faculty research funding provided by the School of Mathematical and Natural Sciences of Berry College to R.E. Carleton, and Natural Sciences and Engineering Research Council of Canada (NSERC) Discovery Grant funding to H.C. Proctor. We thank D.B. Conn, J. Graham, and D. Davin for access to the study area, A. Watson, B. Daniels, H. Pruett, and J. Christian for their assistance in monitoring nest boxes, P.A. 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