Bee Buzz Box May 2025 Smite the Mite Part VI: Going Troppo

Alan Wade

Image: Travel online (2025)

Keith Garvey (1981) tells us how his Uncle Harry quickly despatched an Australian bush tick:

ssiI hurls the rum bottle and hits it [the tick] in the left eye, and slows it down enough for me to scramble up on the wagon. The only thing I can get me hands on is a packet of Cooper’s Dip. I throws it at the tick, and it busts and the yellow powder showers all over him!

That did it! He starts to wobble and shake, then like a drunken man he staggers a few steps and collapses right on the camp fire. The last thing I remember before I passed out from shock was the blue flame flyin’ out of him as his natural gases exploded.

Ticks and parasitic honey bee mites belong to the ‘Acari’ arm of the arachnids. In this they are quite separate from common a garden, 8-legged spiders, scorpions, solifuges (camel spiders) and oliopolones (daddy longlegs). Might we not use Cooper’s arsenic dip to kill Varroa and other troublesome bee parasites? Well, yes we could. The trouble is that it would also spell the death knell of the bees. What we really want are formulations that will kill mites, not take the bees with them.

The task at hand is Varroa control that is in sympathy with good beekeeping practice. However we must be equally aware of other damaging mites all present and creating carnage a short holiday flight or cruise ship trip away. Despite crashing colonies in neighbouring countries few beekeeping magazines and news reports accord these ‘extraterrestrial mites’ any airspace.

Amongst all pests, Tropilaelaps (tropi) mites – along with their viruses – are renowned as the most destructive of all bee diseases. Tropi mites are outstripping Varroa in their capacity to kill bees by a country mile. This, in large measure, is a result of their explosive breeding potential. Tropi have jumped ship from the giant honey bees onto our regular Apis mellifera buzzers as well as onto widely farmed Apis cerana honey bees and dwarf honey bees. Their arrival anywhere and everywhere will seem more like a rat-driven black plague rather than a ‘mild’ Varroa incursion.

Tropi makes Varroa look gentle!
Randy Oliver (2025).

Recent reports on advance of Tropilaelaps into Europe (Brandorf et al., 2024; Carreck, 2023, 2024; Janashia et al., 2024) were foreshadowed by Chantawannakul et al. (2018). Seemingly poor quarantine practices in eastern Europe suggests that this mite will soon become endemic across the whole the European western block. The Middle East, the Americas, the unaffected Pacific (including Australia and New Zealand) will be next in line.

In tropical climes Tropilaelaps rather than Varroa normally prevails. In colder regions, where there is a well-defined brood break, Tropilaelaps does not replicate or survive winter so its populations crash. While colonies or whole apiaries may become nearly mite free, a few mites always survive and return to breed up quickly as brood reappears.

So what happens when both mites are present and become endemic? The populations of both mite species seesaw widely so colonies under stress are prone to collapse year round. At the commencement of a normal season Varroa mite populations grow quickly and continue to develop through to the time when colonies naturally down regulate. Mite numbers remain high in autumn so the infestation rate, mites per pupa, actually increases. In this scenario winter losses of colonies, heavily parasitised by Varroa mites and weakened by viruses, can be extreme.

Tropilaelaps, when also present, behaves somewhat differently. When at the start of the season tropi mite numbers are vanishing low, it can nevertheless reestablish itself quickly and outcompete Varroa – a double jeopardy scenario. Where at all times mite control measures are less than optimal the problem morphs into potential year round colony demise.

The Asian-Pacific multi-honey bee mite scenario

The local scene

Torsten Engelhardt (March 2025), a beekeeper who has battled Varroa in Europe, and is a survivor of the present Sydney ‘mite bombings’, has made a few pertinent observations. Firstly he reports that not all the reported Sydney hive collapses can be attributed to Varroa alone. Small Hive Beetle (SHB) has formed an unholy alliance with Varroa and colonies are dying like flies with SHB slime outs. This condition replicated in tropical Hawaii (Oliver, March 2025): Varroa weakens hives but the beetle finish them off. Other hives, under mite pressure, are also being abandoned leaving full boxes of brood and stores behind. Oliver records that this is also happening in North America though, with Deformed Wing Virus, the brood left behind is invariably in poor condition.

What do these sudden losses to Varroa signal to us, beyond the imperatives of monitoring and instituting effective mite control? Firstly, as well as instituting regular mite checks, I have been installing extra beetle traps. In my experience there is nothing much worse than the prospect of cleaning up slimed out brood. I am also requeening with chalkbrood hygienic stock. Recognising that Varroa will do most of the damage, I am inclined to the view that healthy bees with good winter stores will give bees a more even chance as the mite descends.

The near Asian-Pacific scene

It has been surprising to discover that Australia, Papua New Guinea, the Solomons and most of the Pacific remain free of deformed wing virus (Roberts, 2017). Better known perhaps is that Varroa jacobsoni is widespread and that New Zealand and Tonga are blighted with both Varroa destructor and deformed wing virus (Figure 1).

Figure 1 Near Pacific honey bee-mite incursions.

So what does our near neighbour mite situation portend?Tropilaelaps (the tropi mite), is as resilient as is any bush tick and its arrival is described by USDA’s Samuel Ramsey (2024) as:

a fate worse than Varroa’.

American honey bee researcher Randy Oliver (2024) concurs:

‘…if we get Tropilaelaps it is is going to make Varroa look like nothing. It is an absolutely devastating parasitic mite far worse of a problem than the Varroa mite.’

Tropilaelaps mercedesae moved across from Apis dorsata and Apis laboriosa to Apis mellifera colonies wherever hives were collocated. The mite has now spread northwards through China into iaia westward towards Europe (Brandorf et al., 2024; Janashia et al., 2024). Namin and coworkers (2024) have tracked its genetic variations and phylogenetic patterns across its extended northern range.

Tropilaelaps has also moved eastward from the Indonesian Archipelago to Papua New Guinea, the Solomon Islands and Fiji and poses an existential risk to bees across the Pacific including Australia and New Zealand. 

The damage to Australian beekeeping is currently limited to Varroa destructor in New South Wales. It has recently spread to Victoria and Queensland and it seems inevitable that Australia and New Zealand will suffer not only from the from the Varroa destructor mite but also import three other mites, Varroa jacobsoniTropilaelaps mercedesae and Tropilaelaps clareae (Table 1). All are very proximate in the near Asia-Pacific region (Figure 1).

Destructive honey bee mitesOriginal hostsOther hosts
Tropilaelaps clareaeApis breviligula and Apis dorsata binghamiApis mellifera and Apis cerana in the Phillipines and the Sulawesi
Tropilaelaps mercedesaeApis dorsata and Apis laboriosaApis mellifera, Apis cerana and Apis florea
Varroa destructorApis ceranaKorean and Japanese halotypes only on Apis mellifera
Varroa jacobsoniApis cerana japonicaApis mellifera and Apis dorsata

Table 1 Destructive honey bee mites. Others (de Guzman et al., 2017) have reported a much greater association of parasites with other bee species, e.g. Varroa destructor and Tropilaelaps mercedesae with Apis florea.

Figure 2 Global distribution of Tropilaelaps mercedesae. Redrawn after Jordan and Rogers (2024). The type species Tropilaelaps clareae (Delfinado, 1963) is native to the Philippines (other than the Palawan Islands) and the Sulawesi in Indonesia and has been found in Thailand and India but has now spread further from Indonesia into Papua New Guinea (Figure 3a). Similar maps for Tropilaelaps mercedesae abound (Direct Science, 2025; The National Bee Unit,2017; Melissocosmos, 2025; Gill, 2025).

The Tropilaelaps mites

Of all the mites Tropilaelaps mercedesae is of greatest concern. It was originally confused withTropilaelaps clareae but reclassified as a separate species by Anderson and Morgan (2007). Widespread on the southern Asian continent and in Indonesia, and hosted by Apis dorsata and Apis laboriosa, it is spreading globally (Figure 2, Figure 3c). The record of Tropilaelaps clareae being in Burma and Pakistan (Delfinado-Baker, 1982) is incorrect and is an early record of Tropilaelaps mercedesae.

Tropilaelaps clareae, the type species, was first identified from Apis mellifera colonies and local field rats in the Philippines (Delfinado and Baker, 1961) and later found as a separate serotype in the Sulawesi. It but is now known to have had a much wider natural distribution elsewhere and across the Indonesian Archipelago (Figure 3a). It too is a very serious pest of Apis mellifera.

Tropilaelaps thaii hosted and restricted to Apis laboriosa has only been found in Vietnam (Anderson and Morgan, 2007) but these researchers suggest it is likely more widespread.

A related species Tropilaelaps koenigerum was first found in Sri Lanka (Delfinado-Baker and Baker, 1982; Koeniger, et al., 1983) but is also now recognised as being fairly widespread (Figure 3b). Neither Tropilaelaps thaii nor Tropilaelaps koenigerum have have crossed over to Apis mellifera or Apis cerana.

Chantawannakul and coworkers (2016) describe the dynamic relationship between the Varroa and Tropilaelaps affecting Apis mellifera colonies living sympatrically with Apis cerana and with the original host Apis dorsata group.

Tropilaelaps mercedesae is more prevalent in many Asian Apis mellifera colonies than Varroa: destructor; and conversely

Varroa destructor is more competitive than Tropilaelaps mercedesae in temperate climates.

Natural mite host defence mechanisms

Oldroyd and Wongsiri (2009) describe the natural defence mechanisms that Asian honey bees have mounted to control their Varroa, Euvarroa and Tropilaelaps mite populations. The propensity of most Asian bees to swarm regularly, in many instances to abscond, and in some cases to seasonally migrate (to follow flowering plant phenology), are undoubtably major factors in their achieving mite control. As significantly both the giant honey bees and the cavity dwelling Asian honey bee clades have very effective auto (self) and allo (communal) grooming behaviours (Büchler et al.,1992; Delfinado-Baker et al.,1992), limit mite development to drone brood and have well developed traits for detection and removal (or imprisonment) of parasitised larvae. Such traits are expressed more weakly in European honey bees though African honey bees races have quickly adapted to mite incursions.

So what other lessons might we learn about Tropilaelaps control from their natural giant bee host biology? Firstly we have a preemptory warning from the UK Bee Improvement and Bee Breeders Association (Gill 2025):

In areas where both Tropilaelaps and Varroa are present it has been reported that Tropilaelaps are a far more damaging pest than Varroa and cause colony losses of between 50% to 80%. This is in part due to their lifecycle and more rapid reproductive rate. Tropilaelaps can mate outside of brood cells and unmated females can reproduce via deuterotokous parthenogenesis (de Guzman et al., 2018), a form of asexual reproduction whereby unfertilised eggs can develop into either males or females.

Figure 3 Contemporary distribution of Tropilaelaps mites: a Tropilaelaps clareaeb Tropilaelaps koenigerum,c Tropilaelaps mercedesaed Tropilaelaps thaii.
Source: Chantawannakul et al. (2016).

Elsewhere de Guzman and coworkers (2017) put the threat of Tropilaelaps rather more firmly into perspective:

For much of the world, Varroa destructor single-handedly inflicts unsurmountable problems to A. mellifera beekeeping. However, A. mellifera in Asia is also faced with another genus of destructive parasitic mite, Tropilaelaps. The life history of these two parasitic mites is very similar, and both have the same food requirements (i.e., hemolymph [more correctly fat bodies] of developing brood. Hence, parasitism by Tropilaelaps spp., especially Tropilaelaps mercedesae and Tropilaelaps clareae, also results in death of immature brood or wing deformities in infested adult bees.

Tropilaelaps presents problems very different to those associated with Varroa and other parasitic mites. While all bee parasites feed on pupae and vector and amplify serious viruses their life systems differ in important ways. Tropilaelaps prevails where there is no natural brood break, a seasonal characteristic of warm-temperate and tropical climates. Tropi mites have a shorter development cycle than varroa and consequently outcompete them. The situation is more dynamic in cold regions. In the absence of brood, Tropilaelaps does not replicate and mite populations crash. And while colonies or whole apiaries may become seemingly free of this mite, some mites do survive, possibly on an alternative host, and anyway return in spring. Despite the dependence of Tropilaelaps on the presence of brood for reproduction it can reproduce on other honey bees including Apis florea (Gill, 2025) that appear to have facilitated its spread.

The dynamic is then one of Varroa mite populations continuing to develop as colonies naturally down regulate their number in autumn. This results in winter loss of colonies exacerbated by bees weakened by mites, the classic colony collapse disorder. Tropilaelaps behaves differently. Its mite numbers quickly grow from vanishingly low to outcompete Varroa by mid season. This double whammy can morphs into year round colony demise.

Tropi impact and treatment

If we look at the Tropilaelaps mite more closely we find that it has a biological profile more damaging to bees than that presented by Varroa:

  • its replication rate is almost double that of Varroa;
  • its life system is almost entirely confined to living under protective brood cell cappings; and
  • its mating and reproduction is not confined to the brood nest.

De Guzman and coworkers (2017) have shown that this mite can mate beyond the confines of the brood cell and it has a capacity to reproduce parthenogenically. While colonies or whole apiaries may become mite free in the absence of brood, the few mites that do survive allow them to bounce back. Together the presence of both Varroa and Tropilaelaps can morph into year round colony losses.

In this scenario, a common strategy to control both Varroa and Tropilaelaps will be needed. The broad-scoping review of de Guzman; Williams, Khongphinitbunjong and Chantawannakul (2017) outlines how this might be done. They survey many studies (Mahmood et al., 2012; Raffique et al., 2012; Hoppe et al.,1989) for Tropilaelaps control spanning the efficacy and limitations of use of synthetic miticides, thymol, the organic acids and various forms of brood cycle intervention. Other studies by this group (Chantawannakul et al., 2016; Khongphinitbunjong et al., 2016; Chantawannakul et al., 2018; de Guzman et al., 2018) focus further on the biology and control of Tropilaelaps. Khongphinitbunjong et al. (2015) point particularly to the loss of condition and deformities that Tropilaelaps inflicts of bees. Roberts and coworkers (2019, 2020a, 2020b) report on low cost measures, mainly inducing brood breaks, to control Varroa and Tropilaelaps in Papua New Guinea and in the Solomons.

There appears to be some overall consensus that extended brood breaks or miticides that persist for several brood cycles as well as formic acid will be the most effective ways to target Tropilaelaps. In one defining trial, Kochansky and Shimanuki (1999) employed a formulation of formic acid infused into a polyacrylamide gel to facilitate the release of the acid of a period of 2-3 weeks. It proved to be effective against parasitic honey bee mites including Tropilaelaps. Hoppe and coworkers (1989) employed cardboard plates infused with 20 g of formic acid, introduced at four days intervals placing these plates below rather than above brood frames to achieve a high kill rates of Varroa,Tropilaelaps and Acarapis mites, a scheme that avoided afflicting damage to bee brood and queens.

Other hive mites

Mercedes Delfinado appears to be the first to recognise that the triumvirate of AcarapisVarroa and Tropilaelaps would decimate keeping of western honey bees (Apis mellifera) in Asia. In later describing the behaviour of mites associated with honey bess, Delfinado-Baker, Rath, and Boecking(1992) were wont to quote Eric Binns prescient observation:

Why grow wings, said the mite, when you can thumb a ride on a passing fly [bee]? Some mites take up this specialised form of parasitism to reach an assured food supply, while others hitchhike to escape from a dying habitat.’
Eric S. Binns, 1976

Binns (1982) better known for his critical appraisal of the phenomenon of phoresy that only sometimes is associated with parasitism where he notes that:

In bee-mite relationships in the honey bee Apis mellifera L. in Germany relatively few of 18 phoretic species had any ‘permanent’ relationship to the individual or the colony.

Michael Bush (2025) lists 751 species of mites that have been found in honey bee colonies though few are phoretic and even fewer parasitic. Most of the many common mite species are saprophytic (scavenger) opportunists, unpaid house cleaners that remove fungal debris from the floor of tree hollow colonies and the bottom boards of hives. Widely commonly reported mites include Trichouropoda (Rubink et al.,1991; Delfinado-Baker et al.,1992), Pseudacarapis (Ochoa et al., 2003), Vairimorpha ceranae (Namin et al., 2024), and the widespread pollen mite Melittiphis alvearius (Barreto et al., 2004; Delfinado-Baker, 1994; Brandorf et al., 2024).

However outside the Acarapis, Euvarroa, Tropilaelaps and Varroa mite genera only Leptus ariel and possibly a few allied species (Southcott, 1989; Wilson et al.,1987; Wilson et al., 1990; Martin and Correia-Oliveira, 2016) are known to parasitise honey bees (Table 2).

There are of course other mites that have skipped species and affect honey bees other than Apis mellifera. To the Euvarroa, Varroa, Acarapis and Tropilaelaps parasite genera associated with Asian honey bees, Oldroyd and Wongsiri have compiled a list of nonpathogenic mites mainly discovered in Apis cerana honey bee colonies (Oldroyd and Wongsiri, 2009, Table 9.1, p.182; Poorna Chandra et al., 2021; Koeniger et al., 1983). For example they describe a similar association of the phoretic and non parasitic mite Neocypholaelaps indica with Apis cerana and list ten scavenger as well as Varroa mites found in Apis cerana colonies.

Mite order Mite familyParasite speciesHost species
AstigmatinaTarsonemidaeAcarapis woodiApis mellifera
Acarapis externus
Acarapis dorsalis
Not known but found on Apis dorsata, Apis cerana and Apis mellifera in Asia
MesostigmataVarroidaeVarroa destructorApis cerana
Varroa jacobsoni
Varroa rindereri
Varroa underwoodi
Likely Apis cerana
Apis koschevnikovi
Apis cerana
Euvarroa wongsirii
Euvarroa sinhai
Apis andreniformisApis florea
LaelapidaeTropilaelaps clareae
Tropilaelaps mercedesae
Tropilaelaps koenigerum
Tropilaelaps thaii
Apis breviligula and Apis dorsata binghami
Apis dorsata 
and Apis laboriosa
Apis dorsata
Apis laboriosa
ProstigmataErythraeidaeLepus arielApis mellifera scuttelata

Table 2 Phoretic honey bee parasites.

Whether we are prepared for a multi mite beekeeping future is a moot point. However it seems inevitable that Varroa control will not be the only challenge. We should be prepared for the impact of the likely establishment of bee viruses (likely deformed wing virus), Tropilaelaps mites and for marauding Asian hornets.

Readings

Anderson, D.L. and Morgan, M.J. (2007). Genetic and morphological variation of bee-parasitic Tropilaelaps mites (Acari: Laelapidae): New and re-defined species. Experimental and Applied Acarology 43(1):1-24. https://doi:10.1007/s10493-007-9103-0

Binns, E.S. (1982). Phoresy as migration: Some functional aspects of phoresy in mites. Biological Reviews 57(4):571-620. https://doi.org/10.1111/j.1469-185X.1982.tb00374.x

Barreto, M., Burbano, M.E. and Barreto, P. (2004). The bee mite Melittiphis alvearius (Berlese)(Acari: Laelapidae) in Colombia, South America. Neotropical Entomology 33(1):107-108. https://www.researchgate.net/publication/250030081_The_bee_mite_Melittiphis_alvearius_Berlese_Acari_Laelapidae_in_Colombia_South_America

Brandorf, A., Ivoilova, M.M., Yañez, O., Neumann, P. and Soroker, V. (2024). First report of established mite populations, Tropilaelaps mercedesae, in Europe. Journal of Apicultural Research 68(2):1-3. https://www.wellesu.com/10.1080/00218839.1963.11100070

Büchler, R., Drescher, W. and Tornier, I. (1992). Grooming behaviour of Apis cerana, Apis mellifera and Apis dorsataand its effect on the parasitic mites Varroa jacobsoni and Tropilaelaps clareaeExperimental and Applied Acarology 16(4):313-319. https://scispace.com/pdf/grooming-behaviour-of-apis-cerana-apis-mellifera-and-apis-4tddv4ts3o.pdf

Bush, M. (accessed 6 February 2025). Bee mites. https://bushfarms.com/beesmites.htm

Carreck, N.L. (2023). Is Tropilaelaps on the move? The Beekeepers Quarterly 152:8-9.

Carreck, N.L. (2024). Tropilaelaps in Russia and beyond. The Beekeepers Quarterly 157:22-23.

Chantawannakul, P., de Guzman, L.I., Li, J. and Williams, G.R. (2016). Parasites, pathogens, and pests of honeybees in Asia. Apidologie 47:301-324. https://doi.org/10.1007/s13592-015-0407-5

Chantawannakul, P., Ramsey, S., Khongphinitbunjong, K. and Phokasem, P. (2018). Tropilaelaps mite: An emerging threat to European honey bee. Current Opinion in Insect Science 26:69-75. https://doi.org/10.1016/j.cois.2018.01.012

de Guzman, L.I., Williams, G.R., Khongphinitbunjong, K. and Chantawannakul, P. (2017). Ecology, life history, and management of Tropilaelaps mites. Journal of Economic Entomology 110(2):319-332. https://www.wellesu.com/10.1093/jee/tow304

de Guzman, L.I., Phokasem, P., Khongphinitbunjong, K., Frake, A.M. and Chantawannakul, P. (2018). Successful reproduction of unmated Tropilaelaps mercedesae and its implication on mite population growth in Apis melliferacolonies. Journal of Invertebrate Pathology 153:35-37. https://sci-hub.sidesgame.com/10.1016/j.jip.2018.02.010

Delfinado, M.D. and Baker, E.W. (1961). Tropilaelaps, a new genus of mite from the Philippines (Laelapidae [s. lat.], Acarina). Fieldiana Zoology 44(7):53–56. https://www.biodiversitylibrary.org/partpdf/36189

Delfinado, M. (1963). Mites of the honeybee in South-east Asia. Journal of Apicultural Research 2(2):113–114. https://www.wellesu.com/10.1080/00218839.1963.11100070

Delfinado-Baker, M. (1982). New records for Tropilaelaps clareae from colonies of Apis cerana indica. American Bee Journal 125(5):382.

Delfinado-Baker, M. and Baker, E.W. (1982). A new species of Tropilaelaps parasitic on honey bees. American Bee Journal 122(6):416-417.

Delfinado-Baker, M., Rath, W. and Boecking, O. (1992). Phoretic bee mites and honeybee grooming behavior. International Journal of Acarology 18(4):315-322. https://www.wellesu.com/10.1080/01647959208683966

Delfinado-Baker, M. (1994). A harmless mite found on honeybees Melittiphis alvearius: From Italy to New Zealand. American Bee Journal 134(3):199.

Direct Science (accessed 5 February 2025). Tropilaelaps. https://tropilaelaps.info/honeybees-colony-invading-pest-180973241/

Engelhardt, T. (March 2025) pers comm.

Garvey, K. (1981). The giant tick. ABC Books, Sydney.

Gill, M. (2025). Bee Improvement and Bee Breeders Association. Tropilaelaps. https://bibba.com/tropilaelaps/

Hoppe, H., Ritter, W. and Stephen, E.W.C. (1989). The control of parasitic bee mites: Varroa jacobsoni, Acarapis woodi and Tropilaelaps clareae with formic acid. American Bee Journal 129(11):739-742. https://scholar.google.com.au/scholar?hl=en&as_sdt=0%2C5&q=+The+control+of+parasitic+bee+mites%3A+Varroa+jacobsoni%2C+Acarapis+woodi+and+Tropilaelaps+clareae+with+formic+acid.&btnG=

Janashia, I., Uzunov, A., Chen, C., Costa, C. and Cilia, G. (2024). First report on Tropilaelaps mercedesae presence in Georgia: The mite is heading westward!. Journal of Apicultural Science 68(2):183-188. https://intapi.sciendo.com/pdf/10.2478/jas-2024-0010

Jordan, M. and Rogers, S. (5 November 2024). Current distribution of Tropilaelaps mercedesae associated with Apis mellifera honey bees. Auburn University. https://www.honeybeepests.org/tropi-distribution

Khongphinitbunjong, K., Neumann, P., Chantawannakul, P. and Williams, G.R. (2015). The ectoparasitic mite Tropilaelaps mercedesae reduces western honey bee, Apis mellifera, longevity and emergence weight, and promotes deformed wing virus infections. Journal of Invertebrate Pathology 137:38-42. https://sci-hub.sidesgame.com/10.1007/s13592-015-0407-5

Kochansky, J. and Shimanuki, H. (1999). Development of a gel formulation of formic acid for control of parasitic mites of honey bees. Journal of Agricultural and Food Chemistry 47(9):3850-3853. https://www.wellesu.com/10.1021/jf9901439

Koeniger, N., Koeniger, G. and Delfinado-Baker, M. (1983). Observations on mites of the Asian honeybee species (Apis cerana, Apis dorsata, Apis florea). Apidologie 14(3):197-204. https://www.apidologie.org/articles/apido/pdf/1983/03/Apidologie_0044-8435_1983_14_3_ART0005.pdf

Mahmood, R., Wagchoure, E.S., ul Mohsin, A., Raja, S. and Sarwar, G. (2012). Control of ectoparasitic mites in honeybee (Apis mellifera L.) colonies by using thymol and oxalic acid. Pakistan Journal of Zoology 44(4):985-989. https://www.academia.edu/57831568/Control_of_Ectoparasitic_Mites_in_Honeybee_Apis_mellifera_L_Colonies_by_Using_Thymol_and_Oxalic_Acid

Martin, S.J. and Correia-Oliveira, M.E. (2016). The occurrence of ecto-parasitic Leptus sp. mites on Africanized honey bees. Journal of Apicultural Research 55(3):1–4. https//doi:10.1080/00218839.2016.1228214

Melissocosmos (accessed 25 March 2025). Tropilaelaps mite. https://melissocosmos.blogspot.com/2022/03/tropilaelaps-mite-echthros-melisswn.html

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