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Article

The High Endemism of Haemosporidian Lineages in a Southern Vietnam Avian Community

by
Gary Voelker
1,*,
Mariel Ortega
1,
McKenna Sanchez
1,
Katrina D. Keith
1,
Evgeniy A. Koblik
2,
Andrey V. Bushuev
3,4,
Anvar B. Kerimov
3,4,
Nguyễn Văn Linh
4 and
Sergei V. Drovetski
5,†
1
Department of Ecology and Conservation Biology, Texas A&M University, College Station, TX 77843, USA
2
Department of Ornithology, Zoological Museum of Lomonosov Moscow State University, 6 Bol’shaya Nikitskaya St, Moscow 103009, Russia
3
Department of Vertebrate Zoology, Faculty of Biology, Lomonosov Moscow State University, Moscow 119991, Russia
4
Southern Branch of Joint Vietnam-Russia Tropical Science and Technology Research Center, 3 Street 3/2, 10 District, Ho Chi Minh City 700000, Vietnam
5
Department of Vertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, DC 20004, USA
*
Author to whom correspondence should be addressed.
†
Current address: U.S. Geological Survey, Eastern Ecological Science Center at the Patuxent Research Refuge, Laurel, MD 20708, USA.
Diversity 2025, 17(8), 568; https://doi.org/10.3390/d17080568
Submission received: 14 May 2025 / Revised: 6 August 2025 / Accepted: 9 August 2025 / Published: 13 August 2025
(This article belongs to the Special Issue Bird Parasites—3rd Edition)
Browse Figure
Versions Notes

Abstract

Avian haemosporidians are globally distributed protozoan parasites transmitted to birds by dipteran vectors. The effects of haemosporidian infections are wide-ranging and generally manifest as negative impacts on avian survival and fitness. The study of avian haemosporidians has grown considerably over the past 20 years, such that inter-regional and global effects can be explored. However, considerable gaps in intra-regional studies remain; such studies are critical to broader assessments. In this study, we conducted a community survey of avian haemosporidians found in birds in Cát Tiên National Park, Vietnam. We quantified relative parasite abundance and host associations, and compared our results to data from nearby countries. We also assessed the broader geographic distributions of previously described haemosporidian lineages that we identified in our sample. We screened 320 birds and detected infections in just 28 (8.8%). Infections were either Haemoproteus (12, 3.8%) or Plasmodium (16, 5.0%); we detected no Leucocytozoon infections. We recovered 23 haemosporidian lineages, of which 19 are new, suggesting a high degree of parasite endemism. While our positive detection rate is considerably lower than expected, the high proportion of new versus previously described lineages highlights the need for a continued focus on localized studies to broaden our knowledge of intra- and inter-regional distributions as the field seeks the mechanisms underlying generalized patterns of parasite distribution.

1. Introduction

Avian haemosporidians are intracellular protozoan parasites that infect birds worldwide [1,2]. Found primarily in three genera (Haemoproteus, Leucocytozoon, and Plasmodium), these haemosporidians are vectored by biting midges and hippoboscid flies, black flies, and mosquitos, respectively [3,4,5]. Various abiotic factors affect the manner in which haemosporidian parasite prevalence is distributed; these factors can affect the parasites themselves or their vectors. Examples of these factors include latitude [6], elevation [7,8,9,10], season [11,12], temperature [13,14], and the availability of water [13,15,16]. Assessments of bioclimatic variables have demonstrated that abiotic factors differentially affect regional prevalences of Haemoproteus, Leucocytozoon, and Plasmodium (e.g., [2,6]), and in general, factors depressing the presence or prevalence of one haemosporidian genus are unlikely to have the same effect on the other two genera, assuming conditions that allow the parasites and vectors to reproduce. Only rarely have very low rates of infection prevalence been reported from well-sampled avian hosts, yet these analyses often do not include analyses of all three haemosporidian lineages and are often taxonomically restricted in terms of avian host diversity (e.g., [17,18,19]).
Collectively, haemosporidians have wide-ranging and often detrimental effects on their avian hosts. These effects can include changes in plumage coloration and feather growth rates [20,21], lowered fitness [22,23,24,25], and depressed immune function during migration [26]. Migratory birds are also associated with inter-regional transmission of these pathogens, thereby altering host–parasite dynamics in the avian communities in which they occur, specifically breeding and wintering grounds and possible stopover sites during migration [27,28]. Detrimental effects will become more widespread and will intensify as climate continues to change, as this allows the vectors to expand their ranges poleward and higher in elevation [29,30], and as human urbanization or other changes in land use increase [31,32].
Given the growing number of avian haemosporidian studies over the past 20 years, the field has reached a point where inter-regional and global assessment studies are possible and where unifying themes to explain distributions are being sought [2,6,33,34,35,36]. Critical to these broad-scale assessments are the foundational studies on which they depend: assessments of haemosporidian prevalence in avian host communities at localized scales. In this regard, there are many under-studied regions or countries that have seen little or no avian haemosporidian research, creating substantial gaps in regional and global assessment attempts.
Mainland southeast Asia is one such under-studied region. Using the “Hosts and Sites” table in MalAvi ([37]; http://mbio-serv2.mbioekol.lu.se/Malavi/, accessed on 10 June 2024; database version 2.5.9), we were able to determine that 95 infections have been reported in various studies of wild and captive birds in Thailand (e.g., [38,39,40]), 20 have been reported in mainland Malaysia [41], and none have been reported in Laos, Burma, or Cambodia. In this study we investigated the prevalence of haemosporidian infections in birds sampled in Vietnam, where just two samples have been previously reported, both from red junglefowl (Gallus gallus; [42,43]). Our goals were to (1) add to the (currently) scant information from mainland southeast Asia by providing a broad survey of a local avian community in southern Vietnam, (2) report identified avian host infections by each haemosporidian lineage and document the presence of both previously described and novel lineages, and (3) determine whether any of the lineages in our study are distributed outside of mainland southeast Asia, particularly in northern regions. Finding such connections would imply migratory connectivity linking birds that breed in Siberia (for example), but which broadly winter in southern Asia.

2. Materials and Methods

2.1. Study Site and Sampling

Blood samples were collected from birds (320 individuals representing 52 species) in Cát Tiên National Park (11.42133, 107.42785), Vietnam, from March to May 2015 (Table S1). The park is ca. 72,500 hectares in size, with primary, secondary, and bamboo forests as well as seasonally flooded grasslands. The eastern boundary of the park is the Đồng Nai River.
Birds were captured using mist nets arranged in a series of three interconnected nets, set primarily in bamboo and ginger thickets or heavily disturbed lowland tropical forest. Net arrays were moved every few days to avoid re-captures, but always placed within ca. 300 m of the river. Netting was conducted throughout daylight hours, and after capture birds were subsequently bled under permits from the ethics committee of Lomonosov Moscow State University (Application No.: 26(6) conforming to GOST 33215-2014 and Directive 2010/63/EU). Blood was accessed via ulnar venipuncture using a sterile syringe needle and collected via a hematocrit tube. Approximately 50–100 μL of whole blood was then preserved primarily in 96% ethanol, with 50 of the 320 samples instead being preserved in EDTA buffer. Birds were identified according to species, all were identified as adults via plumage characteristics, and where possible, using those same plumage characteristics (or in a few cases via brood patch presence), birds were designated as female or male.

2.2. DNA Extraction and PCR

DNA was extracted with the E.Z.N.A. tissue extraction kit (Omega Bio-Tek, Norcross, GA, USA) following standard protocols. The polymerase chain reaction (PCR) was used to identify haemosporidian infection through amplification of a fragment of the mtDNA cytochrome b (cytb) gene. Multiple primer pairs were used to amplify across known avian haemosporidian genetic diversity following molecular protocols previously described in [13] using primer pairs UNIVF with UNIVR1, UNIVR2, and UNIVR3 [44]. These primers pairs amplify all three genera of avian haemosporidia: Haemoproteus, Leucocytozoon, and Plasmodium. Each sample was subjected to PCR a minimum of three times, using UNIVF with each reverse primer; samples were considered to have an infection if positive for any of these first three PCRs. Each sample was subjected to a repeat PCR using the same primer pair if initially found to be infection-negative, to confirm that negative result (i.e., six negative PCRs were needed to designate a sample as negative for infection).
To verify a positive infection, 4 μL of the final PCR product was electrophoresed with 2 μL of 100 bp Promega DNA ladder (Thermo Fisher Scientific, Waltham, MA, USA) on an agarose gel. All reactions were performed with negative (same PCR mix, minus DNA) and positive controls (individuals determined positive) to evaluate the validity of the PCR results and detect possible contamination. Positive PCR fragments were sequenced in both directions, using the primer(s) that they were positive for. Successfully amplified PCR products were purified with ExoSap-It, following the manufacturer’s protocols. Sequencing was performed by Psomagen (Rockville, MD, USA).

2.3. Data Analyses

Each sequence was trimmed to 479 bp in length. In instances where a sequence contained two peaks at one or more bases, which is indicative of a co-infection, we did three things. First, we verified by eye that the two peaks at those bases were of roughly the same height, indicating a real co-infection. Where we determined that one peak was instead a weak peak, we corrected the sequence to reflect the clear, strong peak read. Second, we verified that the calls at the remaining multi-peak bases were not tri-allelic codes (i.e., not B, D, H, or V IUPAC codes). Third, we used PHASE [45,46] in DnasP v6 [47] to reconstruct single infection haplotypes. At 10,000 iterations with a burn-in of 1000 iterations, in all instances, PHASE returned two haplotypes (an “a” and a “b” lineage) only.
We then compared our sequences to published sequences housed on the MalAvi database ([37]; http://mbio-serv2.mbioekol.lu.se/Malavi/, accessed on 10 June 2024), version 2.5.9, or GenBank (NCBI, https://blast.ncbi.nlm.nih.gov/Blast.cgi, accessed on 26 August 2024). We utilized the Basic Local Alignment Search Tool (BLAST v. 2.16.0) function in both databases to compare our sequences to those in the database (of similar length), and used the “Grand Lineage Summary” table in MalAvi to determine the genus of each parasite. Sequences that were identical to a known lineage were designated as that lineage. Following [37,48], sequences that had ≥ 1 bp difference were assigned a new lineage name. In several instances, sequences we recovered were determined to be the closest match to the same named lineages, and were the same number of base pairs different from that lineage. In these instances, we used GenBank BLAST to align (compare) those sequences with one another to determine whether they were an exact match (i.e., the same novel lineage) or not (i.e., two novel lineages). We utilized the MalAvi “Parasite Summary Per Host” report tab (accessed 13 June 2024) to, for positive samples, determine which avian species represented in our sample have been previously reported to have been infected with the detected haemosporidians. To determine how common previously known lineages that we recovered were, and from where they had been reported, we used the “Hosts and Sites Table” tab in the MalAvi database (accessed 13 June 2024).
For a molecular analysis of our sequences relative to the known lineages identified in our sample, we used jModelTest 2.1.10 to select the appropriate model of evolution (TIM2+I+G) using the Akaike information criterion. Bayesian phylogenetic analyses were performed using MrBayes 3.2.7 using the CIPRES Science Gateway. This analysis consisted of 2 simultaneous runs for one million generations with four chains (one heated), sampling every 1000 generations, and a 10% burn-in. Each independent run was assessed for convergence via both PSRF (all ca. 1.0) and average ESS values (all > 200), and a 50% majority rule consensus tree was derived from the runs.

3. Results

The overall detection rate for our 320 avian host samples yielded just 28 positive individuals (8.8%) including 12 that were positive for Haemoproteus (3.8%) and 16 that were positive for Plasmodium infections (5.0%; Table 1); we recovered no Leucocytozoon lineages. We detected Haemoproteus co-infections in 2 host individuals, and Plasmodium co-infections in 5 host individuals (Table 1).
Of the 52 avian species for which we had samples, 19 (36.5%) were positive for haemosporidian infection. Neither of the 2 most heavily sampled species, the little spiderhunter (Arachnothera longirostra, n = 84) and the pin-striped tit-babbler (Mixornis gularis, n = 29), were positive for haemosporidian infections.
For Haemoproteus, 14 infections were recovered from 11 host species (Table 1) representing eight different avian families (Alcedinidae [n = 2], Strigidae [1], Phylloscopidae [1], Dicruridae [1], Aegithinidae [1], Pycnonotidae [3], Muscicapidae [1], and Cisticolidae [1]) belonging to three orders (Coraciiformes [2], Strigiformes [1], and Passeriformes [8]).
We identified three Haemoproteus sequences as known lineages (DICLEU02, GW1, and GLACUC03) from MalAvi or GenBank (100% BLAST matches). Eight Haemoproteus lineages were novel, and these differed from known lineages by 1–22 base pairs; one novel lineage (IOLPRO02, closest blast to BUL1) was recovered from two host species (Table 1, Figure 1).
Table 2. The closest BLAST matches to the new lineages described here. The species, avian family, and geographic distribution are provided for these closest matches. Our ORTATR01 and ORTATR02 blast to a large clade including widespread GW1.
Table 2. The closest BLAST matches to the new lineages described here. The species, avian family, and geographic distribution are provided for these closest matches. Our ORTATR01 and ORTATR02 blast to a large clade including widespread GW1.
Our lineageNearest BLASTSpecies found inAvian familyGeographic distribution
LARCYA01MYIFLA01Myiopagis flavivertexTyrannidaePeru
PYCCON01SHOWMAJ03Sholicola majorMuscicapidaeIndia
IOLPORO02BUL1Pycnonotus barbatusPycnonotidaeBenin
Not reported on GenBank Sweden
AEGLAF01POMSUP01Pomatostomus superciliosusTimaliidaeAustralia
DICLEU03CXPIP27Corvus coroneCorvidaeItaly, Portugal
LACPUL02CHLIND01Chloroceryle indaAlcedinidaeBrazil
GLACUC09AEFUN03Aegolius funereusStrigidaeCzech Republic
GLACUC8Butastur linventerAccipitridaeThailand
Glaucidium cuculoidesStrigidaeThailand
PITMOL02/03183Not reported on GenBank Mexico
PYCFIN01/02AEGTIP01Aegithina tiphiaAegithinidaeThailand
Dicrurus leucocephalusDicruridaeThailand
LARCYA02/03DELURB5Chrysomma sinenseTimaliidaeIndia, Myanmar
Delichon urbicumHirundinidaeSpain
Hirundo rusticaHirundinidaeSpain
Egretta garzettaArdeidaeChina
Emberiza godlewskiiFringillidaeChina
Ficedula hyperythraMuscicapidaeChina
Heterophasia melanoleucaTimaliidaeChina
Luscinia svecicaTurdidaeCzech Republic
Oriolus oriolusOriolidaeIndia
Parus monticolusParidaeChina
Prinia inornateCisticolidaeIndia
Prinia socialisCisticolidaeIndia
Saxicola rubetraTurdidaeSweden
Sylvia borinSylviidaeSpain
Zosterops palpebrosusZosteropidaeIndia
MALCIN01CERRUB01Pipra rubrocapillaPipridaeBrazil
For Plasmodium, 20 infections were recovered from nine host species (Table 1) representing five avian families (Cuculidae [n = 1], Pittidae [1], Pycnonotidae [2], Muscicapidae [4], and Pellorneidae [2]) belonging to two orders (Cuculiformes [1] and Passeriformes [10]). We identified four Plasmodium sequences as known lineages (ORW1; note that P_PAVE01 from GenBank is a known alternative name for ORW1 on MalAvi) from MalAvi (100% BLAST matches). Ten Plasmodium lineages were novel, and these differed from known lineages by 1–21 base pairs (Table 1). One recovered novel lineage (MALCIN01, closest blast to CERRUB01) was recovered from two host species, as was another novel lineage (LARCYA02, closest blast to DELURB5) (Table 1, Figure 1).
Overall, bird species of two avian families (Pycnonotidae (Iole and Pycnonotus) and Muscicapidae (Copsychus, Cyornis, Larvivora, Muscicapa)) were collectively found to be infected with both Haemoproteus and Plasmodium. At the species level, only the Siberian blue robin (Larvivora cyane) was found to be infected with both Haemoproteus and Plasmodium, although these infections were in different individuals.

4. Discussion

In this study, we present data on avian haemosporidian infections detected in an avian community in southern Vietnam; this is, to our knowledge, the first study from this country and one of just a few studies from mainland southeast Asia (see above). We sampled 320 avian host individuals collectively representing 52 species of birds. We detected positive Haemoproteus infections at a rate of 3.8%, Plasmodium at a rate of 5.0%, and detected no Leucocytozoon infections. For 13 of the avian species sampled here, these are the first reported cases of haemosporidian infection (Table 1). For the Asian brown flycatcher (Muscicapa latirostris [dauurica]), our study is the first to find a Plasmodium infection.
Our overall detection rate of 8.8% is notably low compared to most studies of which we are aware. Examples of more “typical” infection rates include birds sampled during migration along the Texas Gulf Coast (26%; [49]), breeding birds on the Texas Sky Islands (40.8%; [50]), and bird communities in the Andes (31%; [51]), Western Cape, South Africa (29%; [52]), southern Melanesia (33.8%; [53]), the northwest Caucasus and Transcaucasia (50% and 51%, respectively; [6,44]), and a bird community in Iberia (50.5%; [54]). While infection prevalence can often be even higher in some bird communities, such as those sampled in Morocco (78.5%; [54]) and Malawi (79.1%; [55]), lower detection rates in avian communities is not, as alluded to in the Introduction, typical. However, some examples of low detection rates in taxonomically restricted communities have been found, including infection prevalences of 0.009% in Argentine shorebirds [18] and 3.67% in raptors in Thailand [19]. One possible reason for the low detection rate we report here is that we missed some prevalence data because our samples consisted entirely of blood. Several studies have shown that different tissue types (e.g., pectoral muscle versus blood) can yield different infection prevalence rates of parasite genera in the same suites of avian host individuals [56,57]. Specifically, Haemoproteus infection rates tend to be higher in blood samples, while Plasmodium infection rates are higher in pectoral muscle. Our recovered prevalence rates were similar (14 versus 20, respectively), but showed a reverse pattern with respect to recovered infection rates in that we found more Plasmodium in our (exclusively) blood samples. We think it unlikely that sampling of muscle tissue would have increased prevalence to “typical” levels as found in the majority of other studies.
Other potential reasons for the low infection rate we recovered include (1) the fact that parasitemia is generally so severe that most infected individuals die before they can be sampled, (2) the fact that some birds have evolved immune mechanisms that eliminate the parasite [19], (3) the fact that innate immunity exists such that birds do not become infected, as proposed by [17] for a single songbird species and by [18] for shorebirds, or (4) the fact that birds with high levels of parasitemia are less likely to be netted due to being less active [58]. Having not assessed blood smears for microscopy analysis of parasitemia, we cannot exclude the first possibility, but would predict that the infected birds we sampled would have very low parasitemia. We think the latter two possibilities are unlikely explanations at the community level.
In Vietnam, over 70 species of black flies (which transmit Leucocytozoons) occur [59], and Leucocytozoon lineages have been reported from the region (Thailand, 42.6% of 89 recorded lineages) and the adjacent China (19.7% of 394 recorded lineages). Therefore, the fact that we did not record any Leucocytozoon infections is puzzling, as the eastern border of Cát Tiên National Park is the Đồng Nai River and the authors who participated in the collecting efforts also noted considerable water availability in the form of lakes, pools, and marshes. Because black flies require some degree of water flow for egg laying [60], this leads us to suggest that the Đồng Nai River’s flow might be too fast, and that the area overall is generally too wet for successful black fly reproduction.
Of the known Haemoproteus lineages that we recovered, DICLEU02 has been reported from a single Dicrurus leucophaeus sampled in Myanmar. As such, our finding of this lineage in a kingfisher species, which represents a different order (Coraciiformes) than that of Dicrurus (Passeriformes), is novel. We recovered GW1 from a single individual of Phylloscopus plumbeitarsus. While this is a new host–haemosporidian relationship, this haemosporidian lineage has been reported in three other Phylloscopus species (trochiloides, humei, sindianus) sampled in Turkey, Kyrgyzstan, and India. Like P. plumbeitarsus, two of these Phylloscopus species (trochiloides, humei) are migratory, have a breeding range that includes Siberia, and have wintering ranges that include various parts of southern Asia. P. sindianus is an elevational migrant that breeds in the Caucasus and Himalayas. We recovered GLACUC03 from a single individual of Glaucidium cuculoides, and this lineage has only been recorded in that host species in Thailand.
Of the known Plasmodium lineages that we recovered, ORW1 is a widespread generalist lineage, as it has been reported in the avian orders Pelecaniformes, Coraciiformes, Piciformes, Charadriiformes, Falconiforms, Strigiformes, and Passeriformes, with a distribution ranging from Australia to the United Kingdom and the Altai region of Russia. Despite the geographic proximity of Vietnam to China, and the number of lineages reported in the latter (ca. 200, see MalAvi for citations), ORW1 is the only previously known lineage that we recovered that has been found in both countries. For both haemosporidian genera, we find little evidence of connections between southeast Asia and northern Asia that would suggest that avian migratory behavior is responsible for transporting parasites between breeding and wintering areas.
We also used the “Hosts and Sites Table” (accessed on 24 October 2024) to determine where the closest BLAST matches to our new lineages occur (Figure 1); in a few cases we queried GenBank (where a lineage is not listed on MalAvi). These closest BLAST matches were largely found in a single species, with a notable exception being DELURB5, which was reported in 15 species (Table 2). Collectively, these lineages represent 20 avian families and a geographic distribution including Australia, Asia, Europe, Africa, and South America (Table 2). Overall, the majority of lineages that we identified are novel. This may suggest a high degree of endemism in the eastern portion of mainland southeast Asia. The degree and geographic extent of this endemism remains to be confirmed by additional regional sampling.
With respect to avian host infections, it was surprising to find zero infections in the two most commonly sampled bird species, the little spiderhunter (family Nectariniidae) and pin-striped tit-babbler (Timaliidae). However, a lack of haemosporidian infections in some common birds has been reported in communities with high haemosporidian prevalence, e.g., the greater spotted woodpecker (Dendrocopos major [n = 12]), the long-tailed tit (Aegithalos caudatus [n = 25]), and the common firecrest [Regulus ignicapilla [n = 23]) [54]. Nectariniidae species are readily infected by a diverse suite of haemosporidian lineages, often with high prevalence depending on the species, and can be one of the most heavily infected families at the community level, as reported in studies from Africa and southern China [13,61,62]. The high infections rates in Nectariniidae might be facilitated by host behavior, as multiple individuals of multiple species may congregate at and in the vicinity of flowering trees (GV pers. obs., SVD pers. obs.) to forage (dozens of species utilize the same types of plants [63]), which potentially increases infection chances due to proximity. Indeed, a recent study [64] found that mosquitos are attracted by birds infected with Plasmodium, so congregating behaviors might facilitate haemosporidian transfer. However, little spiderhunters forage singly or in pairs [63] and do not spend much time in a single foraging location, which might reduce infection probability due to isolation, but counter to this, the Olive Sunbird (Cyanomitra olivacea), another solitary Nectariniidae species, can be highly parasitized [62]. In contrast to the little spiderhunter, the pin-striped tit-babbler is, outside of the breeding season, found in flocks of 12 or more or in mixed species flocks [65], suggesting an increased opportunity for infection, although we found none. Timaliidae species from southern China, including the pin-striped tit-babbler, are known to be infected by both Plasmodium and Haemoproteus [61].
In conclusion, we found a remarkably low level of haemosporidian infection in a southern Vietnam bird community, and a lack of infection in heavily sampled avian host species. Most of the haemosporidian infections recovered were novel, reflecting high regional endemism; however, this result was not unexpected, given the dearth of avian malaria research in mainland southeast Asia. This study highlights the importance of regional studies that can contribute not only to our understanding of regional host–parasite interactions and distributions, but to studies at inter-regional and global scales where unifying themes to explain haemosporidian distributions are being sought.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/d17080568/s1, Table S1: Avian host samples and recovered haemosporidian lineages.

Author Contributions

Conceptualization, E.A.K., A.V.B., A.B.K., N.V.L. and S.V.D.; methodology, G.V., E.A.K., A.V.B., A.B.K. and S.V.D.; formal analysis, G.V., M.O., M.S. and K.D.K.; investigation, all authors; resources, G.V., E.A.K., A.V.B. and A.B.K.; data curation, G.V., E.A.K., A.V.B., A.B.K. and S.V.D.; writing—original draft preparation, G.V.; writing—review and editing, all authors; supervision, G.V.; project administration, G.V., E.A.K., A.V.B., A.B.K. and N.V.L.; funding acquisition, G.V., A.V.B. and A.B.K. All authors have read and agreed to the published version of the manuscript.

Funding

This research was funded by the Russian Foundation for Basic Research (grant number 15-04-07407) and the Russian Science Foundation (grant number 14-50-00029) with grants awarded to both AVB and ABK.

Institutional Review Board Statement

The study was conducted in accordance with the Declaration of Helsinki and approved by relevant review boards, as documented in the Materials and Methods section.

Data Availability Statement

New haemosporidian sequences from this study have been deposited in MalAvi and GenBank (PV941874-PV941907).

Acknowledgments

We are grateful to the Cát Tiên National Park Authority for permission to sample birds. We thank the Tropical Center for its comprehensive assistance during the material collection phase in Cát Tiên National Park. The work of AVB, ABK, and NVL took place within Task 1.1 of Topic Ecolan E-1.2 of the Tropical Center. We thank R. Weesner for running the Bayesian analysis. This is publication number 1710 of the Biodiversity Research and Teaching Collections at Texas A&M University. Any use of trade, firm, or product names is for descriptive purposes only and does not imply endorsement by the U.S. Government.

Conflicts of Interest

The authors declare no conflicts of interest. The funders had no role in the design of the study; in the collection, analysis, or interpretation of data; in the writing of the manuscript; or in the decision to publish the results.

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Diversity 17 00568 g001
Figure 1. The Bayesian analysis of all novel Haemoproteus and Plasmodium lineages detected in this study, relative to the closest BLAST matches from MalAvi. Lineage names followed by an asterisk are previously described lineages, and our samples are italicized. See Table 1 for the avian hosts associated with the new lineages, and Table 2 for the avian hosts and geographic distributions of previously known lineages.
Figure 1. The Bayesian analysis of all novel Haemoproteus and Plasmodium lineages detected in this study, relative to the closest BLAST matches from MalAvi. Lineage names followed by an asterisk are previously described lineages, and our samples are italicized. See Table 1 for the avian hosts associated with the new lineages, and Table 2 for the avian hosts and geographic distributions of previously known lineages.
Diversity 17 00568 g001
Table 1. The haemosporidian lineages recovered in our study, relative to the avian host species and malaria parasite genus; where applicable, multiple individuals of a given host species are indicated in parentheses after the taxon name. Indented taxon names indicate the same individual as on the preceding line (i.e., co-infections). Lineages were blasted to the MalAvi or GenBank databases, and where it is different from a known lineage, we provide the nearest blasted result in parentheses (column two). Lineages in the same column and without parentheses are 100% matches to a known lineage, which we provide. If different via % match and the number of base pair differences (columns three and four), in the fifth column we provide a novel lineage designation which has been registered on MalAvi. In the last column we provide the number of previously known lineages for each avian host species, for each haemosporidian genus (Haemoproteus, Plasmodium, Leucocytozoon), as reported on MalAvi (our results are not included).
Table 1. The haemosporidian lineages recovered in our study, relative to the avian host species and malaria parasite genus; where applicable, multiple individuals of a given host species are indicated in parentheses after the taxon name. Indented taxon names indicate the same individual as on the preceding line (i.e., co-infections). Lineages were blasted to the MalAvi or GenBank databases, and where it is different from a known lineage, we provide the nearest blasted result in parentheses (column two). Lineages in the same column and without parentheses are 100% matches to a known lineage, which we provide. If different via % match and the number of base pair differences (columns three and four), in the fifth column we provide a novel lineage designation which has been registered on MalAvi. In the last column we provide the number of previously known lineages for each avian host species, for each haemosporidian genus (Haemoproteus, Plasmodium, Leucocytozoon), as reported on MalAvi (our results are not included).
BLAST% MatchBP
difference		Novel Lineage
Designation		Parasite Summary Per Host (H/P/L)
Haemoproteus
ruddy kingfisher (Halcyon coromanda)DICLEU021000 0/0/0
ashy drongo (Dicrurus leucophaeus)DICLEU021000 2/2/0
two-barred warbler (Phylloscopus plumbeitarsus)GW11000 0/0/0
Asian barred owlet (Glaucidium cuculoides)GLACUC031000 3/9/0
  Asian barred owlet (Glaucidium cuculoides)(AEFUN03)99.81GLACUC09above
banded kingfisher (Lacedo pulchella)(CHLIND01)99.43LACPUL021/0/0
great iora (Aegithina lafresnayei)(POMSUP01)99.24AEGLAF010/0/0
ashy drongo (Dicrurus leucophaeus)(CXPIP27)98.57DICLEU03above
grey-eyed bulbul (Iole propinqua)(BUL1)97.910IOLPRO022/0/0
black-headed bulbul (Brachypodius melanocephalos)(BUL1)97.910IOLPRO020/0/0
Siberian blue robin (Larvivora cyane)(MYIFLA01)97.512LARCYA010/0/0
dark-necked tailorbird (Orthotomus atrogularis)(GAGLA02)9716ORTATR010/0/0
  dark-necked tailorbird (Orthotomus atrogularis)(H298)97.711ORTATR02above
streak-eared bulbul (Pycnonotus conradi)(SHOWMAJ03)95.422PYCCON010/0/0
Plasmodium
green-billed malkoha (Phaenicophaeus tristis)ORW11000 0/0/0
stripe-throated bulbul (Pycnonotus finlaysoni)ORW11000 0/0/0
Tickell’s blue flycatcher (Cyornis sumatrensis)ORW11000 0/0/0
Asian brown flycatcher (Muscicapa dauurica)(GLACUC8)99.43MUSLAT010/0/1
  Asian brown flycatcher (Muscicapa latirostris)(GLACUC8)98.95MUSLAT02above
stripe-throated bulbul (Pycnonotus finlaysoni)(AEGTIP01)98.95PYCFIN01above
  stripe-throated bulbul (Pycnonotus finlaysoni)(AEGTIP01)98.95PYCFIN02above
stripe-throated bulbul (Pycnonotus finlaysoni)ORW11000 above
stripe-throated bulbul (Pycnonotus finlaysoni)(ORW1)991PYCFIN03above
scaly-crowned babbler (Malacopteron cinereum)(CERRUB01)97.810MALCIN010/0/0
Abbott’s babbler (Malacocincla abbotti)(CERRUB01)97.810MALCIN010/0/0
white-rumped shama (Copsychus malabaricus) (4)(DELURB5)97.711LARCYA023/2/0
Siberian blue robin (Larvivora cyane) (2)(DELURB5)97.711LARCYA03above
  Siberian blue robin (Larvivora cyane)(DELURB5)97.711LARCYA02above
blue-winged pitta (Pitta moluccensis)(183)95.621PITMOL020/0/0
  blue-winged pitta (Pitta moluccensis)(183)95.820PITMOL03above
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Voelker, G.; Ortega, M.; Sanchez, M.; Keith, K.D.; Koblik, E.A.; Bushuev, A.V.; Kerimov, A.B.; Linh, N.V.; Drovetski, S.V. The High Endemism of Haemosporidian Lineages in a Southern Vietnam Avian Community. Diversity 2025, 17, 568. https://doi.org/10.3390/d17080568

AMA Style

Voelker G, Ortega M, Sanchez M, Keith KD, Koblik EA, Bushuev AV, Kerimov AB, Linh NV, Drovetski SV. The High Endemism of Haemosporidian Lineages in a Southern Vietnam Avian Community. Diversity. 2025; 17(8):568. https://doi.org/10.3390/d17080568

Chicago/Turabian Style

Voelker, Gary, Mariel Ortega, McKenna Sanchez, Katrina D. Keith, Evgeniy A. Koblik, Andrey V. Bushuev, Anvar B. Kerimov, Nguyễn Văn Linh, and Sergei V. Drovetski. 2025. "The High Endemism of Haemosporidian Lineages in a Southern Vietnam Avian Community" Diversity 17, no. 8: 568. https://doi.org/10.3390/d17080568

APA Style

Voelker, G., Ortega, M., Sanchez, M., Keith, K. D., Koblik, E. A., Bushuev, A. V., Kerimov, A. B., Linh, N. V., & Drovetski, S. V. (2025). The High Endemism of Haemosporidian Lineages in a Southern Vietnam Avian Community. Diversity, 17(8), 568. https://doi.org/10.3390/d17080568

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