Bee Buzz Box August 2026 Part II Vairimorpha and the Crithidia – Lotmaria Complex

Alan Wade

Canberra Region Beekeepers

Lotmaria passim

Source: Schwarz et al. (2015)

In Protesting Protists Part I, we surveyed amoebic, gregarine, microsporidian and trypanosomatid honey bee parasites mainly focussing on their diversity and their putative cause of disease. Here we home in on the trypanosomatids (Crithidia mellificae and Lotmaria passim) and their common co-occurrence with the amoebic Nosema (Vairimorpha) gut parasites and their widespread role in causing colony loss.

Older than beekeeping

Ruth Lotmar (1910-1989) is credited with describing the parasite Lotmaria passim (Figure 1) after whom the organism was named (Fuchs, 2019; Lotmar, 1946). The species name comes from the Latin passim, literally here and there or everywhere, an apt epithet for its wide and universal distribution across most bee families and across the Apis genus and indeed global distribution. It is a serious honey bee pathogen that likely predates beekeeping by mankind.

A little more delving signals that the microsporidians and trypanosomatids that make up the most lethal protists are themselves complex assemblages of microorganisms that play havoc with the hymenopteran bee suborder Apoidae (sphecoid wasps and bees). Individual protist species are often bee generalists. For example the Lotmaria relative, the trypanosomatid Crithidia bombi, long regarded as an exclusive parasite of bumblebees, has now been shown to infect a population of honey bees in China (Li et al., 2012).

Looking elsewhere there appear to be other species or variants of Nosema (Vairimorpha), e.g. Vairimorpha D, yet other Crithidia species, a new protist genus Tubulinosema and a variant of Nosema (Vairimorpha) ceranae that targets bumblebees. Our knowledge of the range of bee protists and their geographic distribution keeps expanding attributable in part to a lack of surveillance across many jurisdictions. Let’s keep our minds open to the emergence of such new parasites. A case in point is the inexplicable loss of colonies over the last North American winter (Oliver, 2025).

Figure 1 Crithidia mellificae

Source: Schwartz et al. (2015).

But these complexities will only muddy the waters for keepers looking for solutions not problems. Let’s stick to the simple facts: where are these nasties turning up and which amongst them are inflicting death by a thousand cuts – and what if anything we can do about them.

Where the dead men lie

Dead flies cause the ointment of the apothecary to send forth a stinking savor; so doth a little folly in him that hath a reputation for wisdom and honour.

Ecclesiastes 10:1

Crithidia, Lotmaria and Nosema are of global distribution. More intriguing is the fact that some species are globally dominant and truly serious parasites while a handful are of recent discovery and are more than likely of restricted geographical distribution and of limited importance. An oft cited example is found in Vairimorpha ceranae, syn Nosema ceranae, that evolved with the Asian honey bee Apis cerana, now a debilitating parasite of Apis mellifera having in large measure displaced the native Nosema apis. Nosema ceranae also impacts a close relative of the Asian honey bee, Apis koschevnikovi as well as the dwarf honey bee Apis florea and the giant honey bee Apis dorsata (Chaimanee et al., 2010). In a more nuanced analysis a variant of Nosema ceranae infects bumblebees.

As we saw in Part I there are a number of Nosema species (at least 81 are known) that parasitise invertebrates so the discovery of Nosema neumanni on honey bees in Uganda (see Part I) might not surprise us. We know too little of its distribution and pathogenicity to even hazard a guess as to its risk to global beekeeping or to know whether other bees are impacted. As for the two common species, Ptaszynska and coworkers found that spores of Nosema ceranae are more sculptured with deeper ornamentation than those of Nosema apis (Figure 2).

Figure 2 Nosema spp.

Source: Ptaszynska et al. (2012).

Do we have a problem? Well yes we do. We now have Nosema apis, Nosema ceranae, Crithidia mellificae and, as recently reported, Lotmaria passim in both Australia and New Zealand (Bhasi et al., 2026; Waters, 2018), as well as elsewhere in the Southern Hemisphere (Castelli et al., 2019; Quintana et al., 2021), so the global bee population has the principle protists here reported as the bee botherers. Given their almost universal presence across the bee kingdom (Figure 3) it is perhaps not too soon to say that one protist, Lotmaria passim, is on a par with Nosema ceranae as a very serious bee pathogen. The general consensus is that Lotmaria (though not widely known amongst beekeepers) and not Crithidia mellificae is problematic.

Figure 3 Global distribution of microsporidian and trypanosomatid honey bee and bumblebee parasites: N. = Nosema, C. = Crithidia, L. = Lotmaria, T. = Tubulinosema. Note: Lotmaria passim is in Australia (Bhasi, Zerna and Beddoe, 2026).

Source: Vavilova et al. (2017). Nosema D, which was determined in bumblebee populations from China (Li et al., 2012).

Grim reapers

Early studies (Fantham and Porter,1912a, 1912b; Graham-Smith et al., 1912; Kudo, 1920) pointed to the seriousness of disease caused by the microsporidian Nosema apis though their conclusion that the Isle of Wight disease was caused by this organism was clearly misplaced. They did however find Nosema widespread but of course correlation does not imply disease causation. Nevertheless one surprisingly reads:

At the commencement of the spring of I9I1, there was a great outcry among bee-keepers, who, on opening their hives, found dead bees only. Public comment in the press was a daily occurrence, and plaints of 'no bees, no fruit,' 'a fruitless year,' etc., appeared on all sides. Suggestions had been made by various investigators of the disease, under the Board of Agriculture, that in one case blood-poisoning, and in another a bacillus, was the cause of the trouble. Both cases had broken down. Having revised the accumulation of our experiments, examinations of bees, hives, combs, honey and excrement, we decided to bring forward our results. On April 4, I9I1, therefore, microscopic preparations of Nosema apis, together with infected bees and combs, were exhibited by us before the Zoological Society of London, where an outline in brief of our results was announced and published. Since that time, owing to becoming members of the Board of Agriculture's Enquiry, we have examined very many bees, but the result has been merely to confirm all our previous work, without adding any new facts regarding the parasite to our previous knowledge-a disappointing occurrence. On becoming members of the official Enquiry we delayed publishing our results in extenso, but we propose in these papers to give an account in full of our pioneer researches, confirmed, as they are, abundantly by subsequent investigations, both of our own and of our colleagues on the Enquiry under the auspices of the Board of Agriculture.

Casting aspersions aside, Farnham and Porter were eminent microbiologists and they depicted the gut region affected by Nosema apis very clearly (Figure 4).

Figure 4 Nosema apis location in honey bee gut (shaded).

Source: Fantham and Porter (1912a).

The literature records the fairly recent spread of the microsporidian Nosema (Vairimorpha) ceranaeand its having supplanted Vairimorpha apis (Nosema apis) (Klee et al., 2007), while the trypanosomatids are spread across the super family Apoidae, the bee (and sphecoid wasp) kingdom (Figure 5), the most prevalent of which are Crithidia mellificae and Lotmaria passim. The defining study of Ryan Schwartz and coworkers (2015) clarifies the confused assignment of the monoxenous (single host) trypanosomatids occasioned by classical taxonomy where species were assigned on morphological (shape and form) characteristics rather than on far more clearly defined genetic makeup. So while Crithidia mellificae and Lotmaria passim are clearly delineated as separate species (Ravoet et al., 2015), many of the earlier literature studies attributed to Crithidia mellificae clearly should have been assigned to Lotmaria passim. Two other species, Crithidia expoeki and Crithidia mexicana, are likely synonymous with Lotmaria passim (Schmid-Hempel and Tognazzo, 2010; Gallot-Lavallée et al., 2016).

Figure 5 Trypanosomatid pathogens detected across the different bee families.

Source: Adapted from Olmedo et al. (2025).

One fun fact is that not only is Lotmaria passimmore pathogenic than Crithidia mellificae, it has spread and largely replaced this crithidian. Both may co-infect our honey bees while both Vairimorpha (Nosema) ceranae (a relatively new parasite of honey bees) and Lotmaria passim may also be co-parasites. While not a microbiologist, whenever I hear the term nosemosis, we might invent a new term ‘vairi-a-lot’ to describe the complex of parasites next most common after bee viruses.

Many researchers report the sharing of honey bee pathogens with bumblebees (Gallot-Lavallée et al., 2016; Schmid-Hempel and Tognazzo, 2010) or between different honey bee species (Chaimanee et al., 2010).

The Crithidia-Lotmaria-Leptomonas-Leishmania group of trypanosomatids (Figure 6) are a complex assemblage, their similar morphology often enough masking their true identity. For example many well designed studies of the common parasite Crithidia mellificae have been shown to be actually studies of Lotmaria passim, the latter now more common, the former decreasing in prevalence and seemingly pathogenicity though findings vary (Arismendi et al., 2020, 2022; Gómez-Moracho et al., 2020).

As gut parasites they are transmitted, as are the amoebae, via the faecal oral route. They are either ingested when bees transfer food (trophallaxis) or when nurse bees feed brood. They absorb nutrients and attach, by modifying their flagella, to the hind gut wall where they multiply and are excreted to repeat the cycle (Buendía-Abad et al., 2021a). The direct effects of Lotmaria passim and Vairimorpha/Nosema ceranae on honey bees is to increase foraging and reduce in hive behaviour, i.e. to increase storage and disrupt pivotal nest behaviour (MacInnis et al., 2025; Macpherson, 2025).

Figure 6 Phylogenic reconstruction of Leishmaniinae parasites based on strains of Crithidia mellificae, Lotmaria passim n. gen., n. sp.) and from in vivo isolate sequences. Source: Schwarz et al. (2015).

Various reviews point to the ubiquity and damaging effect of the trypanosomatids: Olmedo et al. (2025) outline the biology and host resistance pointing to a general lack of evidence that Crithidia mellificae had any harmful effects on honey bee health while the effect of Crithidia bombi on Bombus terrestris – the common European bumblebee – received greater attention. The widespread detection of trypanosomatids in genetic surveys coinciding with a global increase in honey bee mortality led to the conclusion that the culprit was likely associated with a second species, Lotmaria passim (Bartolomé et al., 2018; Gómez-Moracho et al., 2020; Pascual et al., 2026; Ravoet et al., 2015; Schwarz et al., 2015; Tafi et al., 2025; Tiritelli et al., 2025). Overall, there are as many as five trypanosomatid species widely associated with honey bees: Crithidia acanthocephali, Crithidia bombi, Crithidia expoeki, Crithidia mellificae and Lotmaria passim (Tiritelli et al., 2025).

Before we leave the protists to their own devices, we might reflect on two major colony collapse events in the Northern Hemisphere in the winter of 2025 and ask ourselves whether we really know their causes. We are yet to see the full impact of Varroa establishment in Australia, but the loss of around 60% of colonies in America (Lamas et al., 2026; Nearman et al., 2025; Carreck, 2026b) over the winter of 2025 (Table 1 shows incidence of presumptive pathogens) and similar losses in the United Kingdom (Carreck, 2026a) should be a wakeup call to what may lie ahead. It would appear that Lotmaria passim may be a significant contributing cause of colony loss but that its overall importance as a truly serious parasite is only now becoming clear.

Pathogen	Number of colonies with detection (n= 113)	Prevalence of detection(%)
Acarapis woodi	0	0.0
Acute bee paralysis virus (ABPV)	81	71.7
Apis mellifera filamentous virus (AmFV)	23	20.4
Black queen cell virus (BQCV)	85	75.2
Chronic bee paralysis virus (CBPV)	5	4.4
Deformed wing virus-A (DWV-A)	88	77.9
Deformed wing virus-B (DWV-B)	19	16.8
Israeli acute bee paralysis virus (lAPV)	5	4.4
Kashmir bee virus (KBV)	32	28.3
Lake Sinai virus (LSV)	72	63.7
Lotmaria passim (Lp)	34	30.1
Nosema ceranae (NOS)	71	62.8
Sacbrood virus (SBV)	40	35.4

Table 1 US 2025 colony losses to common parasitic diseases.

Source: Lamas et al. (2026) reformatted.

Readings

Arismendi, N., Caro, S., Castro, M.P., Vargas, M., Riveros, G. and Venegas, T. (2020). Impact of mixed infections of gut parasites Lotmaria passim and Nosema ceranae on the lifespan and immune-related biomarkers in Apis mellifera. Insects 11(7):420. https://www.mdpi.com/2075-4450/11/7/420

Arismendi, N., Castro, M.P., Vargas, M., Zapata, C. and Riveros, G. (2022). The trypanosome Lotmaria passim prevails in honey bees of different ages and stages of development. Journal of Apicultural Research 61(1):63-69.https://www.researchgate.net/publication/344886987_The_trypanosome_Lotmaria_passim_prevails_in_honey_bees_of_different_ages_and_stages_of_development

Bartolomé, C., Buendía, M., Benito, M., de la Rúa, P., Ornosa, C., Martín-Hernández, R., Higes, M. and Maside, X. (2018). A new multiplex PCR protocol to detect mixed trypanosomatid infections in species of Apis and Bombus. Journal of Invertebrate Pathology 154:37-41.  https://doi.org/10.1016/j.jip.2018.03.015

Bhasi, G., Zerna, G. and Beddoe, T. (2026). From hive to lab: Molecular detection of Lotmaria passim and Crithidia mellificae in Australian honey bees using honey-derived eDNA. Journal of Invertebrate Pathology 226:108577. https://www.sciencedirect.com/science/article/pii/S0022201126000509

Buendía-Abad, M., García-Palencia, P., de Pablos-Torró, L.M., Alunda, J.M., Osuna, A., Martín-Hernández, R. and Higes, M. (2021a). First description of Lotmaria passim and Crithidia mellificae haptomonad stage in the honeybee hindgut. International Journal for Parasitology  52(1):65-75.

https://www.researchgate.net/publication/350844330_First_description_of_Lotmaria_passim_and_Crithidia_mellificae_haptomonad_stage_in_the_honeybee_hindgut

Buendía-Abad, M., Higes, M., Martín-Hernández, R., Barrios, L., Meana, A., Fernández, A.F., Osuna, A. and de Pablos, L.M. (2021b). Workflow of Lotmaria passim isolation: Experimental infection with a low-passage strain causes higher honeybee mortality rates than the PRA-403 reference strain. International Journal for Parasitology: Parasites and Wildlife 14:68-74. https://sci-hub.usualwant.com/10.1016/j.ijppaw.2020.12.003

Carreck, N. (2026a). Alarming reports of UK colony losses. The Beekeepers Quarterly 163:7.

Carreck, N. (2026b). And meanwhile in the USA. The Beekeepers Quarterly 163:8.

Castelli, L., Branchiccela, B., Invernizzi, C., Tomasco, I., Basualdo, M., Rodriguez, M., Zunino, P. and Antúnez, K. (2019). Detection of Lotmaria passim in Africanized and European honey bees from Uruguay, Argentina and Chile. Journal of Invertebrate Pathology 160:95-97. https://doi.org/10.1016/j.jip.2018.11.004

Chaimanee, V., Warrit, N. and Chantawannakul, P. (2010). Infections of Nosema ceranae in four different honeybee species. Journal of Invertebrate Pathology 105(2):207-210. https://www.sciencedirect.com/science/article/abs/pii/S0022201110001436

Fantham, H.B. and Porter, A. (1912a). Microsporidiosis, a protozoal disease of bees due to Nosema apis, and popularly known as Isle of Wight disease. Annals of Tropical Medicine and Parasitology6(2):145-161. https://doi:10.1080/00034983.1912.11687059 https://ia800607.us.archive.org/view_archive.php?archive=/8/items/crossref-pre-1923-scholarly-works/10.1080%252F00034983.1909.11685242.zip&file=10.1080%252F00034983.1912.11687059.pdf

Fantham, H.B. and Porter, A. (1912b). Note on certain protozoa found in bees. Supplement to the Journal of Board of Agriculture 19(8):138. https://books.google.com.au/books/about/Supplement_to_the_Journal_of_the_Board_o.html?id=rUAy4B0P82YC&redir_esc=y

Fuchs, W.J. (6 March 2019). Lotmaria passim: Ruth Lotmar, bee researcher and zoologist. https://www.zwitscher-maschine.org/blog/2019/3/12/hozvqc052x17tbd1x0cucu8ao5smto

Gallot-Lavallée M., Schmid-Hempel R., Vandame R., Vergara C.H. and Schmid-Hempel, P. (2016). Large scale patterns of abundance and distribution of parasites in Mexican bumblebees. Journal of Invertebrate Pathology 133(5):73-82. https://sci-hub.usualwant.com/10.1016/j.jip.2015.12.004

Gómez-Moracho, T., Buendía-Abad, M., Benito, M., García-Palencia, P., Barrios, L., Bartolomé, C., Maside, X., Meana, A., Jiménez-Antón, M.D., Olías-Molero, A.I. and Alunda, J.M. (2020). Experimental evidence of harmful effects of Crithidia mellificae and Lotmaria passim on honey bees. International Journal for Parasitology 50(13):1117-1124. https://www.sciencedirect.com/science/article/abs/pii/S0020751920302368

Graham-Smith, G.S., Fantham, H.B., Porter, A., Bullamore, G.W. and Malden, W. (May 1912). Isle of Wight bee disease. Journal of the Board of Agriculture 19(2):133-134. Summary of article of same name in Supplement to the Journal of the Board of Agriculture, Price 1s, post free.https://dn721604.ca.archive.org/0/items/dli.ernet.29495/29495-The%20Journal%20Of%20The%20Board%20Of%20Agriculture%20Vol-xix%20April%20%2C1912%20To%20March%2C1913.pdf

Klee, J., Besana, A.M., Genersch, E., Gisder, S., Nanetti, A., Tam, D.Q., Chinh, T.X., Puerta, F., Ruz, J.M., Kryger, P. and Message, D. (2007). Widespread dispersal of the microsporidian Nosema ceranae, an emergent pathogen of the western honey bee, Apis mellifera. Journal of Invertebrate Pathology 96(1):1-10.

https://sci-hub.usualwant.com/10.1016/j.jip.2007.02.014

Kudo, R. (1920). Notes on Nosema apis Zander. The Journal of Parasitology 7(2): 85–90. https://doi:10.2307/3270960 https://www.jstor.org/stable/3270960?seq=1

Lamas, Z.S., Rinkevich, F., Garavito, A., Shaulis, A., Boncristiani, D., Hill, E., Chen, Y.P. and Evans, J.D. (2026). Viruses and vectors tied to honey bee colony losses. PLoS Pathogens 22(2):e1013939. https://journals.plos.org/plospathogens/article?id=10.1371/journal.ppat.1013939

Li, J., Chen, W., Wu, J., Peng, W., An, J., Schmid-Hempel, P. and Schmid-Hempel, R. (2012). Diversity of Nosema associated with bumblebees (Bombus spp.) from China. International Journal for Parasitology 42(1):49-61. https://sci-hub.usualwant.com/10.1016/j.ijpara.2011.10.005

Lotmar, R. (1946). Über flagellaten und bakterien im Dünndarm der honigbiene (Apis mellifica). Sauerländer. Beiheft Schweizerische Bienenzeitung 2(14):49-76. Cited by Schwartz et al. (2015).

Macpherson, A. (3 March 2025). Double threat: Research shows how two gut parasites disrupt honeybee behaviour. [Adrianna Macpherson, University of Alberta.] https://www.ualberta.ca/en/folio/2025/03/double-threat-research-shows-how-two-gut-parasites-disrupt-honeybee-behaviour.html

MacInnis, C.I., Luong, L.T. and Pernal, S.F. (2025). Effects of Nosema ceranae and Lotmaria passim infections on honey bee foraging behaviour and physiology. International Journal for Parasitology 55(5):213-223. https://www.sciencedirect.com/journal/international-journal-for-parasitology/vol/55/issue/5

Maslov, D.A., Votýpka, J., Yurchenko, V. and Lukes, J. (2013). Diversity and phylogeny of insect trypanosomatids: All that is hidden shall be revealed. Trends in Parasitology 29(1):43-52. https://doi.org/10.1016/j.pt.2012.11.001

Nearman, A., Crawford, C.L., Guarna, M.M., Chakrabarti, P., Lee, K., Cook, S., Hill, S., Seshadri, A., Slater, G., Lamas, Z., Chen, Y.P., Downey, D. and Evans, J.D. (2025). Insights from U.S. beekeeper triage surveys following unusually high honey bee colony losses 2024-2025. bioRxiv 2025.08.06.668930. https://abfnet.org/wp-content/uploads/2025/08/2025.08.06-Survey-Results.pdf

Oliver, R. (2025). Is our party winding down. American Bee Journal 165(5):551-558.

Olmedo, P.G., Gómez-Moracho, T., Buendía-Abad, M., Carreira-de Paula, J., Palmer-Young, E., Martín-Hernández, R., Evans, J.D., Higes, M. and de Pablos-Torró, L.M. (2025). Trypanosomatid pathology, cell biology, host resistance and genomics in honey bee hosts: The knowns and unknowns. Parasitology 1-55. https://www.cambridge.org/core/journals/parasitology/article/trypanosomatid-pathology-cell-biology-host-resistance-and-genomics-in-honey-bee-hosts-the-knowns-and-unknowns/198B2F454982654C887B963E429CF718

Pascual, M.H., de Pablos-Torró, L.M. and Lihoreau, M. (2026). Eva Crane Trust. Effects of Lotmaria passim parasites on honeybee behaviour and cognition. https://www.evacranetrust.org/en/page/effects-of-lotmaria-passim-parasites-on-honeybee-behaviour-and-cognition

Ptaszynska, A., Borsuk, G., Mulenko, W. and Olszewski, K. (2012). Monitoring of nosemosis in the Lublin region and preliminary morphometric studies of Nosema spp. spores. Medycyna Weterynaryjna 68(10):622-625. http://www.medycynawet.edu.pl/images/stories/pdf/pdf2012/102012/201210622625.pdf

Quintana, S., Plischuk, S., Brasesco, C., Revainera, P., García, M.L.G., Bravi, M.E., Reynaldi, F., Eguaras, M. and Maggi, M. (2021). Lotmaria passim (Kinetoplastea: Trypanosomatidae) in honey bees from Argentina. Parasitology International 81:102244. https://www.sciencedirect.com/science/article/abs/pii/S138357692030194X

Ravoet, J., Schwarz, R.S., Descamps, T., Yañez, O., Tozkar, C.O., Martin-Hernandez, R., Bartolomé, C., De Smet, L., Higes, M., Wenseleers, T. and Schmid-Hempel, R. (2015). Differential diagnosis of the honey bee trypanosomatids Crithidia mellificae and Lotmaria passim. Journal of Invertebrate Pathology 130:21-27. https://sci-hub.usualwant.com/10.1016/j.jip.2015.06.007

Schmid-Hempel, R. and Tognazzo, M. (2010). Molecular divergence defines two distinct lineages of Crithidia bombi (Trypanosomatidae), parasites of bumblebees.  Journal of Eukaryotic Microbiology 57(4):337-345. https://doi.org/10.1111/j.1550-7408.2010.00480.x

Schwarz, R.S., Bauchan, G.R., Murphy, C.A., Ravoet, J., de Graaf, D.C. and Evans, J.D. (2015). Characterization of two species of trypanosomatidae from the honey bee Apis mellifera: Crithidia mellificae Langridge and McGhee, and Lotmaria passim n. gen., n. sp. Journal of Eukaryotic Microbiology 62(5):567-583. https://sci-hub.usualwant.com/10.1111/jeu.12209

Tafi, E., Capano, V., Nanetti, A. and Cilia, G. (2025). A nationwide molecular survey on Trypanosomatids occurrence in Italian honey bee (Apis mellifera L.) colonies. Veterinary Parasitology: Regional Studies and Reports 60:101253. https://www.sciencedirect.com/science/article/abs/pii/S2405939025000619

Tiritelli, R., Cilia, G. and Gómez-Moracho, T. (2025). The trypanosomatid (Kinetoplastida: Trypanosomatidae) parasites in bees: A review on their environmental circulation, impacts and implications. Current Research in Insect Science 7:100106. https://www.sciencedirect.com/science/article/pii/S2666515825000010

Vavilova, V.Y., Konopatskaia, I., Luzyanin, S.L., Woyciechowski, M. and Blinov, A.G. (2017). Parasites of the genus Nosema, Crithidia and Lotmaria in the honeybee and bumblebee populations: A case study in India.Vavilov Journal of Genetics and Breeding 21(8):943-951. https://ruj.uj.edu.pl/server/api/core/bitstreams/80cdb550-4154-4b80-b7d3-988b063fd4a6/content

Waters, T.L. (2018). The distribution and population dynamics of the honey bee pathogens Crithidia mellificae and Lotmaria passim in New Zealand. (Doctoral dissertation, Open Access Te Herenga Waka-Victoria University of Wellington). https://figshare.com/articles/thesis/The_distribution_and_population_dynamics_of_the_honey_bee_pathogens_Crithidia_mellificae_and_Lotmaria_passim_in_New_Zealand/17068349?file=31560035