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International Journal for Parasitology: Parasites and Wildlife logoLink to International Journal for Parasitology: Parasites and Wildlife
. 2025 Jul 29;28:101123. doi: 10.1016/j.ijppaw.2025.101123

Unmasking Borrelia species: A comprehensive review of their presence in Iran

Parisa Soltan-Alinejad a, Mahmood Nikbakhtzadeh b, Eslam Moradi-Asl a,⁎
PMCID: PMC12345004  PMID: 40808974

Abstract

Borrelia species are the causative agents of Lyme disease (LD) and tick-borne relapsing fever (TBRF) in humans and animals. These pathogens are transmitted through hard and soft ticks. The increasing tick population, influenced by climate change, underscores the urgent need for enhanced research on tick-borne diseases.

Iran, situated in southwestern Asia, boasts a diverse climate that supports a wide range of tick species and their vertebrate hosts. While TBRF is endemic to Iran, recent reports suggest the presence of LD in the country as well. Understanding the various Borrelia species, their tick vectors, human cases, affected reservoirs, and geographical distribution is crucial for assessing the epidemiology of TBRF and LD in Iran.

This comprehensive review examines the epidemiological patterns, geographical distribution, detection methods for these pathogens, providing critical insights into their public health significance.

Keywords: Borrelia, Tick, Tick-borne disease, Lyme disease, Relapsing fever, Iran

Graphical abstract

Image 1

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Highlights

  • •

    Tick borne relapsing fever (TBRF) is endemic in Iran, and Lyme disease (LD) has recently been reported.

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    The distribution of Borrelia in Iran has increased, affecting 20 provinces.

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    Mazandaran province shows the highest diversity of Borrelia species, with six types documented.

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    Hard ticks (I. ricinus and Rh. annulatus) can also transmit TBRF in Iran, caused by B. miyamotoi and B. theileri.

1. Introduction

Borrelia is classified within the spirochaetia class, comprising organisms that belong to the Spirochaetota phylum within the eubacteria kingdom. The genus Borrelia was recognized over a century after the discovery of Borrelia recurrentis, the pathogen responsible for louse-borne relapsing fever (LBRF) (Saint Girons et al., 1994). Various Borrelia species can cause diseases, including Relapsing Fever (Jakab et al., 2022) and Lyme Disease (LD) (Stanek and Strle, 2003). Relapsing fever is classified as either tick-borne (TBRF) or louse-borne (LBRF) (Felsenfeld,1971). LBRF is an anthroponotic disease caused by B. recurrentis, whereas TBRF is a zoonotic disease resulting from various Borrelia species (Jakab et al., 2022). Borrelia spp. are transmitted through both hard and soft ticks, as well as human body lice (Koutantou et al., 2024). Relapsing fever, encompassing both LBRF and TBRF, is typically observed in the western United States, Africa, Asia, Saudi Arabia, and Spain. These diseases remain epidemic in developing countries due to poverty, war, and famine (Hoch et al., 2015). The TBRF enzootic cycles differ, encompassing various tick species and their hosts, predominantly small rodents. Furthermore, they can infect humans, who act as accidental dead-end hosts (Killilea et al., 2008; Rodino et al., 2020). TBRF has recently been associated with travel-related infections following visits to endemic regions (Rebaudet and Parola, 2006). The global reduction of lice-infested populations has led to LBRF becoming an uncommon disease, while TBRF has become a growing concern in recent years (Jakab et al., 2022) (see Table 1, Table 2).

Table 1.

Summary of published identified and unidentified Borrelia species in hosts across Iran.

Host Borrelia species Location Clinical complication Laboratory method/Target gene Accession Numbers Treatment Ref.
Identified Borrelia species responsible for TBRF Accidental dead-end host Human B. balthazardi Ardabil – Microscopic examination – – Karimi et al., 1979
Human/7-day-old neonate/Male (Congenital TBRF) B. persica Zanjan Malaise, jaundice, weakness of neonatal reflexes, and sepsis Microscopic examination – Erythromycin Mahram and Ghavami, 2009
Human/5 cases B. persica Khorasan-e Razavi Relapsing episodes of fever, chills, and headache Microscopic examination – – Shayeghi et al., 2016
Two-week-old puppy B. persica Tehran Vomiting, diarrhea, and loss of appetite Microscopic examination and PCR/IGS KR816159 Metoclopramide, Vitamin B12, and Ampicillin Shirani et al., 2016
Reservoir Meriones persicus B. duttonii-like spirochetes East Azerbaijan – Microscopic examination, qPCR, and PCR/flaB, glpQ, groEL, p66, rrs, IGS, and 16S rRNA MW767941-5 for IGS, MW795343-6 for P66, MW79533-5 for flaB, MW737426-9 for rrs, MW795340-2 for groEL, and MW79536-9 for glpQ. – Ghasemi et al., 2021
M. persicus B. duttonii East Azerbaijan – qPCR/16S rRNA – – Rezaie et al., 2025
M. persicus B. persica Hamadan – qPCR/16S rRNA – –
Identified Borrelia species responsible for LD Accidental dead-end host Human/I case B. burgdorferi Mazandaran Erythema migrans ELISA – Doxycycline Adabi et al., 2004
Dogs B. burgdorferi Gilan, Mazandaran, and Golestan No clinical signs ELISA and Western blot – – Hanifeh et al., 2012
Companion dogs B. burgdorferi Ahvaz No clinical signs Immunochromatography assay (ICA) – – Mosallanejad et al., 2015
Dogs B. burgdorferi Ahvaz – ELISA – – Rezaei et al., 2016
Companion dogs B. burgdorferi Fars No clinical signs ELISA – – Esmailnejad et al., 2017
Human/1 case (neuroborreliosis) B. burgdorferi – Seizures Chestcomputed tomography (CT) scan, MRI, and serological tests – Ceftriaxone Sayad et al., 2023
Guard dogs B. burgdorferi Isfahan No clinical signs Microscopic examination and PCR/16S rRNA – – Chaleshtory et al., 2023
Unidentified Borrelia species Human/13 cases Borrelia spp. Kazeroun – Microscopic examination – – Janbakhsh and Ardelan, 1977
Human/391 cases Borrelia spp. Ardabil Headache, fever, chills, nausea, vomiting, abdominal pain, cough, sweating, hematuria, jaundice, arthralgia, petechiae, photophobia, and eosinophilia Microscopic examination – – Arshi et al., 2002
Human/1 case Borrelia spp. Ardabil Headache, vomiting, fever, positive Kernig's sign, and neck stiffness (Meningitis) Microscopic examination – Doxycycline Majid-Pour, 2003
Borrelia meningitis
Human/97 cases Borrelia spp. Kurdistan Sweats, nervousness, headache, fever, chills, vomiting, stomach ache, myalgias, nose bleeding, cough, arthralgia, and photophobia Microscopic examination – – Rafinejad et al., 2012
Human/138 cases Borrelia spp. Kurdistan Chills, fever, sweating, headache, abdominal pain, vomiting, myalgia, arthralgia, nervousness, cough, photophobia and nose bleeding Microscopic examination – – Kassiri et al., 2014b
Human/148 cases Borrelia spp. Kurdistan Fever, chills and headache Microscopic examination – – Moemenbellah-Fard et al., 2009
Human/11 cases Borrelia spp. Kurdistan Chills, fever, sweating, headache, abdominal pain, vomiting, myalgia, arthralgia, nervousness, cough, photophobia and nose bleeding Microscopic examination – – Kassiri et al., 2014a
Human/1 case Borrelia spp. Fars Fever and tachycardia Microscopic examination – Intravenous penicillin G four times a day for 24 h and Oral penicillin for 10 days. Pouladfar et al., 2008
Human/14 cases Borrelia spp. Hormozgan Headache, fever, chills, nausea, vomiting, abdominal pain, cough, sweating, hematuria, jaundice, arthralgia, petechiae, photophobia, eosinophilia, muscle and joint pain Microscopic examination/PCR – Tetracycline Naddaf et al., 2015
One-month-old puppy Borrelia spp. Tehran Icteric mucous membranes, malaise, tenderness upon palpation, pustular lesions, loss of appetite, diarrhea, and fever Microscopic examination – Prednisolone and Doxycycline Rostami et al., 2011
Human/276 cases Borrelia spp. Hamedan – Microscopic examination – – Nazari and Najafi, 2016
Human/2 cases Borrelia spp. Jask/Hormozgan Fever, fatigue, and headache PCR Tetracycline Naddaf et al., 2017
Human/1 case Borrelia spp. Tehran Joint pain in the left elbow and left ankle, erythema migrans on foot Microscopic examination, ELISA, and Western blot Doxycycline Siadati, 2006
Sheep and goats/1018 cases Borrelia spp. West Azerbaijan – PCR/glpQ, IGS, and flaB KX683867 for IGS, KX683864 and KX683865 for glpQ, and KX683866 for flaB – Enferadi et al., 2024a
Cats and dogs Borrelia spp. West Azerbaijan No clinical signs PCR/5–23 S rRNA OR770094, OR770195, OR770198, and OR771914 – Ownagh et al., 2024
Hares and long-eared hedgehogs (Hemiechinus megalopolis) Borrelia spp. Sistan and Baluchistan – PCR/5–23S rRNA OR400934 and OR398185 – Sarani et al., 2024
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Table 2.

Summary of published identified and unidentified Borrelia species in different tick species in Iran.

Disease Tick vector Tick status in the field (Feeding or questing) Borrelia species Province Laboratory method/Target gene Accession Numbers Ref.
Identified Borrelia species related to TBRF O. tholozani – B. persica Markazi – - Moradi-Asl and Jafari, 2020
O. tholozani Questing and feeding on domestic animals B. persica Semnan Animal inoculation, microscopic examination – Nekoui et al., 1999
O. tartakovskyi Questing in rodent burrows and bird nests B. latyschewii Khorasan-e Razavi Animal inoculation, microscopic examination – Piazak et al., 2000
O. tholozani Questing in human dwelling and animal B. persica Hamedan Animal inoculation, microscopic examination – Vatandoost et al., 2003
O. tholozani Questing in human dwelling, animal shelters B. persica Kurdistan Animal inoculation, microscopic examination – Rafinejad et al., 2012
O. tholozani Questing in human dwelling and animal B. persica Qazvin Animal inoculation, microscopic examination – Aghighi et al., 2007
O. erraticus B. microti –
O. tholozani Questing in human dwelling, animal shelters, and rodent burrows B. persica Qazvin PCR-RFLP/16S rRNA EU914141 Barmaki et al., 2010
O. erraticus Feeding on rodents B. microti Alborz Animal inoculation, microscopic examination, and PCR/16S rRNA, flaB, glpQ, and IGS JF803950, JF825472, JF825473, and JQ436580 Naddaf et al., 2012
O. tholozani Questing in human dwelling, animal shelters B. persica Khorasan-e Razavi Animal inoculation, microscopic examination – Shayeghi et al., 2016
A. persicus Questing in cracks of aviary B. anserina Lorestan PCR/flaB KY438930 Chegeni et al., 2017
I. ricinus Questing and feeding (camels, goats, donkeys, hedgehogs, sheep, rodents, horses, cattle, and dogs) B. miyamotoi Mazandaran qPCR and PCR/16S rRNA, flaB, and rrs-rrlA MN958345 to MN958348 Naddaf et al., 2020
Rh. annulatus Questing and feeding (dogs, sheep, cattle, and goats) B. theileri Mazandaran qPCR and PCR/16S rRNA, flaB, and glpQ OR037296-OR037302 for flaB and OR037292- OR037295 for glpQ Milani et al., 2024
Identified Borrelia species related to LD I. ricinus Questing and feeding (camels, goats, donkeys, hedgehogs, sheep, rodents, horses, cattle, and dogs) B. bavariensis, B. valaisiana, B. afzelii Mazandaran qPCR and PCR/16S rRNA, flaB, and rrs-rrlA MN958342, MN958343, MN958344 Naddaf et al., 2020
I. ricinus B. garinii Gilan MN958341
Unidentified Borrelia species O. tholozani Questing in human dwelling, animal shelters, and rodent burrows Borrelia spp. Kurdistan Animal inoculation, microscopic examination – Moemenbellah-Fard et al., 2009
O.lahorensis, O.tholozani, and A. persicus Questing Borrelia spp. Ardabil Animal inoculation, microscopic examination – Arshi et al., 2002
O. lahorensis Questing in human dwelling, animal shelters, and rodent burrows Borrelia spp. Takistan/Qazvin PCR-RFLP/16S rRNA – Barmaki et al., 2010
O. tholozani Questing in animal shelters Q Zanjan Animal inoculation, microscopic examination – Mohammadi et al., 2013
O. tholozani/eggs Questing in animal dwelling Borrelia spp. Ardabil Animal inoculation, microscopic examination and PCR/rrs-rrlA – Aghaei et al., 2014
Rh. turanicus, and Rh. sanguineus Questing and feeding (camels, goats, donkeys, hedgehogs, sheep, rodents, horses, cattle, and dogs) Borrelia spp. Golestan qPCR and PCR/16S rRNA, flaB, and rrs-rrlA MN958349 to MN958351 Naddaf et al., 2020
Rh. sanguineus, H. asiaticumss, H.aegyptium, H. anatolicum Feeding on sheep and goats Borrelia spp. West Azerbaijan PCR/16srRNA, 5S-23SrRNA, and ospA OQ073805.1, OR342388.1, OR352151.1, and OR345451.1. Enferadi et al., 2024b
Hyalomma aegyptium. and Rh. sanguineus Feeding on rabbits and hedgehogs. Borrelia spp. Sistan and Baluchistan PCR/5–23S rRNA – Sarani et al., 2024
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LD is recognized as the most rapidly increasing tick-borne zoonotic disease worldwide, with reported cases in more than 60 countries and established endemic regions in North America, Europe, and Asia (Donohoe et al., 2015). It is transmitted through infected hard tick bites, primarily from ticks of the Ixodes genus (Giesen et al., 2024). The reservoir hosts for LD include various mammalian species, particularly rodents, as well as other animals capable of hosting hard ticks (Kazemi-Moghaddam et al., 2019). Between 1982 and 2022, the Centers for Disease Control and Prevention (CDC) documented more than 844,000 cases of LD (Mead et al., 2024). The burden of LD has increased, spreading to previously non-endemic regions (Stone et al., 2017). In this study, we assessed the status of Borrelia spp. in Iran, located in southwestern Asia.

2. Epidemiological pattern of identified Borrelia species

2.1. TBRF history, vectors, and hosts (reservoir and dead-end)

TBRF is recognized as an endemic disease in Iran. Between 1997 and 2006, more than 1400 cases were documented across 19 provinces (Asl et al., 2009). The epidemiology of the disease in various regions is influenced by the interactions between ticks, Borrelia species, and environmental conditions within their respective distribution areas (Vatandoost et al., 2003). Previously, B. persica was identified as a causative agent, with Ornithodoros tholozani reported as the main vector in Iran (Karimi et al., 1979; Naddaf et al., 2015). Additionally, B. baltazardi and B. latyschewii have been recorded in northwestern and northeastern Iran (Karimi et al., 1979). Concerning TBRF within their natural enzootic cycles, both humans and dogs serve as accidental dead-end hosts (Rodino et al., 2020). An exception to this is B. duttonii, for which humans might act as the reservoir (Jakab et al., 2022). B. persica has been documented in human TBRF cases in the provinces of Zanjan and Khorasan-e-Razavi (Mahram and Ghavami, 2009; Shayeghi et al., 2016). B. baltazardi has been detected in the blood serum of patients in Ardabil (Karimi et al., 1979). TBRF can manifest in neonates, including cases of congenital disease. A congenital TBRF case has been documented in Zanjan, where a 7-day-old neonate exhibited malaise, jaundice, diminished neonatal reflexes, and sepsis, but lacked hepatosplenomegaly, fever, and respiratory distress (Mahram and Ghavami, 2009). However, infected human cases in Khorasan-e-Razavi presented with relapsing fever episodes, chills, and headaches (Shayeghi et al., 2016). An instance of B. persica infection has also been documented in a two-week-old puppy in Tehran, which exhibited vomiting, diarrhea, and loss of appetite (Shirani et al., 2016). Wild rodents are generally the primary reservoir hosts (Rebaudet and Parola,2006). The rodent species Meriones persicus, commonly found in Iran, serves as a host for Borrelia species responsible for TBRF. In the provinces of East Azerbaijan and Hamedan, B. persica and B. duttonii, as well as B. duttonii-like spirochetes, have been successfully isolated from M. persicus (Adabi et al., 2004; Ghasemi et al., 2021; Rezaie et al., 2025).

Information on TBRF tick vectors indicates that B. persica has been identified in O. tholozani across the provinces of Markazi, Semnan, Hamedan, Kurdistan, Qazvin, and Khorasan-e- Razavi (Nekoui et al., 1999; Vatandoost et al., 2003; Aghighi et al., 2007; Barmaki et al., 2010; Rafinejad et al., 2012; Shayeghi et al., 2016; Moradi-Asl,2020). B. microti has been reported from O. erraticus in Qazvin and Alborz (Aghighi et al., 2007; Naddaf et al., 2012). Other studies have recorded B. anserine and B. latyschewii from Argas persicus and O. tartakovsky, respectively, in Lorestan and Khorasan-e-Razavi (Piazak et al., 2000; Chegeni et al., 2017).

Recent findings indicate that hard ticks can also transmit TBRF. Ixodes ricinus and Rhipicephalus annulatus have been documented as TBRF hard tick vectors for B. miyamotoi and B. theileri in Mazandaran (Naddaf et al., 2020; Milani et al., 2024).

2.2. LD history, vectors, and hosts (reservoir and dead-end)

The first report of endemic LD caused by B. burgdorferi in Iran occurred in 1997 (Chams-Davatchi, 1997), serving as an alert for medical professionals to consider LD in their diagnostic evaluations. Following this case report, several LD patients have been documented over time in the country. All identified Borrelia species in Iran are shown in Fig. 1. So far, among the various Borrelia species that cause LD, only B. burgdorferi has been reported from hosts and ticks in Iran. Humans and dogs inadvertently become part of this cycle, acting as dead-end hosts (Humair and Gern, 2000; Wagner and Erb, 2012). Conversely, small wild mammals such as rodents and possibly some migratory birds play a role as reservoirs in maintaining the Lyme borreliosis cycle (Wagner and Erb, 2012). In Iran, there have been two reported cases of human Lyme borreliosis. One case occurred in Mazandaran (Adabi et al., 2004), while the region associated with the Lyme neuroborreliosis case has not been specified (Sayad et al., 2023). In Lyme neuroborreliosis, initially, the patient exhibited low-grade fever, headache, vomiting, and malaise; however, after a week, they were hospitalized due to the onset of seizures (Sayad et al., 2023). This zoonotic disease has also been documented in dogs across various regions of Iran, including Gilan, Mazandaran, Golestan, Khuzestan, Fars, and Isfahan (Hanifeh et al., 2012; Mosallanejad et al., 2015; Rezaei and Gh, 2016; Esmailnejad et al., 2017; Chaleshtory et al., 2023). Notably, B. burgdorferi did not cause any clinical signs in dogs (Hanifeh et al., 2012; Mosallanejad et al., 2015; Esmailnejad et al., 2017; Chaleshtory et al., 2023). Findings regarding tick vectors of LD indicated that B. bavariensis, B. valaisiana, B. afzelii, and B. garinii have been successfully isolated from I. ricinus in Mazandaran and Gilan provinces (Naddaf et al., 2020).

Fig. 1.

Fig. 1

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Different types of Borrelia species by the counties.

The geographical distribution of identified Borrelia species in hosts is presented in Fig. 2. Additionally, the geographical distribution of identified Borrelia species from different tick species is shown in Fig. 3.

Fig. 2.

Fig. 2

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Identified Borrelia species in various hosts by the counties. The symbols represent humans, dogs, and rodents, indicating the Borrelia-infected specimens.

Fig. 3.

Fig. 3

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Identified Borrelia species in various tick species by the counties. The symbols represent different species of ticks infected with Borrelia.

2.3. Unidentified Borrelia species reported in vectors and hosts

Previous studies have recorded human cases of TBRF symptoms in Fars (Janbakhsh and Ardelan, 1977), Ardabil (Arshi et al., 2002; Majid-Pour, 2003), Kurdistan (Moemenbellah-Fard et al., 2009; Rafinejad et al., 2012; Kassiri et al., 2014; Kassiri et al., 2014), Fars (Pouladfar et al., 2008), Hormozgan (Naddaf et al., 2015, Naddaf et al., 2017), and Hamedan Provinces (Nazari and Najafi, 2016). However, the species of Borrelia involved in these cases have not been identified. A human case with LD clinical symptoms was reported from Tehran, while the type of Borrelia is unknown (Siadati, 2006). A one-month-old puppy was diagnosed with a Borrelia spp. infection in Tehran (Rostami et al., 2011). Blood samples from cats, dogs, sheep, and goats in West Azerbaijan (Enferadi et al., 2024; Ownagh et al., 2024), and hares and long-eared hedgehogs (Hemiechinus megalotis) in Sistan and Baluchistan revealed the presence of Borrelia spp. (Sarani et al., 2024). Unidentified Borrelia species have also been documented in soft and hard tick vectors. The identified soft ticks were infected with Borrelia spp. include O. tholozani from Kurdistan (Moemenbellah-Fard et al., 2009), Ardabil (Arshi et al., 2002; Aghaei et al., 2014), and Zanjan (Mohammadi et al., 2013); O. lahorensis from Ardabil (Arshi et al., 2002) and Qazvin (Barmaki et al., 2010); and A. persicus from Ardabil (Arshi et al., 2002). The identified hard ticks are infected with Borrelia spp. include Rh. turanicus and Rh. sanguineus from Golestan (Naddaf, Mahmoudi et al.,2020); Rh. sanguineus, Hyalomma asiaticum, H. aegyptium, and H. anatolicum from West Azerbaijan (Enferadi et al., 2023); and Hyalomma aegyptium. and Rh. sanguineus from Sistan and Baluchistan (Sarani et al., 2024). The geographical distribution of unidentified Borrelia species in hosts and ticks has been illustrated in Fig. 4.

Fig. 4.

Fig. 4

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Unidentified Borrelia species in various hosts and tick species by the counties.

3. Laboratory methods for Borrelia detection

3.1. Microscopic examination

Giemsa staining was employed for the identification of Borrelia spp. in the microscopic method. The stained slide demonstrated the presence of spirochetes within the thin blood smear obtained from an infected canine (Rostami et al., 2011; Shirani et al., 2016; Chaleshtory et al., 2023), M. persicus (Ghasemi et al., 2021), and human (Janbakhsh and Ardelan, 1977; Arshi et al., 2002; Majid-Pour, 2003, Siadati, 2006; Pouladfar et al., 2008; Mahram and Ghavami, 2009; Moemenbellah-Fard et al., 2009; Rafinejad et al., 2012; Kassiri et al., 2014; Kassiri et al., 2014; Naddaf et al., 2015; Nazari and Najafi, 2016; Shayeghi et al., 2016).

3.2. Animal inoculation

It is important to understand that the sensitivity of microscopy is defined by specific positivity thresholds, which are estimated to be 105 spirochetes per mL of blood for thin smears and 104 spirochetes per mL for thick smears (Hovette et al., 2001). As a result, more investigations use animal inoculation before direct microscopic examination. Bacteria can be introduced into the animal either by direct injection or by allowing ticks to feed on it. One or two weeks later, blood samples from the inoculated animals were tested for Borrelia using Giemsa staining, microscopic examination, or PCR. In this method, a suspension made from live ticks, including O. lahorensis, O. tholozani, and A. persicus, was injected into the peritoneum of guinea pigs and white mice in Ardabil for Borrelia spp. (Arshi et al., 2002). O. tholozani was inoculated into suckling Syrian hamsters or white mice for Borrelia spp. in Kurdistan (Moemenbellah-Fard et al., 2009). O. tholozani was also intraperitoneally inoculated into white mice with B. persica (Nekoui et al., 1999; Vatandoost et al., 2003; Mohammadi et al., 2013; Shayeghi et al., 2016), and O. erraticus was inoculated into guinea pigs with B. microti (Naddaf et al., 2012). In the tick-feeding method, unfed O. tartakovskyi infected with B. latyschewii, and O. tholozani infected with B. persica were fed on white mice in Khorasan-e-Razavi and Hamedan, respectively (Piazak et al., 2000, Vatandoost et al., 2003). In another study, infected O. tholozani were fed on guinea pigs, while O. erraticus were fed on newborn mice. As a result, B. persica and B. microti were successfully isolated from O. tholozani and O. erraticus, respectively (Aghighi et al., 2007). Unfed O. tholozani infected with Borrelia spp. fed on guinea pigs and mice in Semnan, Zanjan, and Ardabil (Nekoui et al., 1999; Arshi et al., 2002; Mohammadi et al., 2013). In a separate investigation conducted in Kurdistan, O. tholozani was examined for the presence of Borrelia spp. through a three-step process: the ticks were first crushed and inoculated into guinea pigs and white mice, followed by blood-feeding on the guinea pigs, and finally, a subcutaneous injection of the crushed ticks was administered (Rafinejad et al., 2012).

3.3. Molecular technique

The Nested-PCR technique was employed for the identification of Borrelia species after DNA extraction from tissues (tick and animal) or blood samples. This technique uses specific primers to precisely identify the species of spirochete. Multilocus sequence analysis (MLSA) used for Borrelia identification includes the intergenic spacer region (IGS) (Naddaf et al., 2012; Naddaf et al., 2015; Shirani et al., 2016; Naddaf et al., 2017; Ghasemi et al., 2021), 5–23S rRNA gene (Mosallanejad et al., 2015; Enferadi et al., 2023; Enferadi et al., 2024a; Ownagh et al., 2024; Sarani et al., 2024), non-coding rrs-rrlA spacer (Aghaei et al., 2014; Naddaf et al., 2020; Ghasemi et al., 2021), flagellin (flaB) (Chegeni et al., 2017; Naddaf et al., 2017; Naddaf et al., 2020; Ghasemi et al., 2021; Milani et al., 2024), glycerophosphodiester phosphodiesterase (glpQ) (Naddaf et al., 2012; Naddaf et al., 2017; Ghasemi et al., 2021; Milani et al., 2024), groEL (Ghasemi et al., 2021), p66 (Ghasemi et al., 2021), 16S rRNA region (Chaleshtory et al., 2023), and ospA (Enferadi et al., 2023; Enferadi et al., 2024). In several investigations, the qPCR/Real-Time PCR technique was utilized to detect Borrelia spp. using a specific probe targeting the 16S rRNA region (Naddaf et al., 2020; Ghasemi et al., 2021; Milani et al., 2024; Moradkasani et al., 2024; Rezaie et al., 2025). Furthermore, PCR-restriction fragment length polymorphism (RFLP) analysis was employed to identify B. persica in ticks (Barmaki et al., 2010).

3.4. Serological tests

The serological testing for Borrelia species, particularly in LD, generally consists of a two-phase procedure. The initial phase employs an enzyme-linked immunosorbent assay (ELISA) to identify antibodies targeting B. burgdorferi. If the outcome of this first test is either positive or uncertain, a Western blot test is administered to confirm the diagnosis. In several studies, ELISA and/or Western blot, and immunochromatography assay (ICA) techniques were utilized to detect the presence of B. burgdorferi in dogs (Adabi et al., 2004; Siadati, 2006; Hanifeh et al., 2012; Mosallanejad et al., 2015; Rezaei and Gh,2016; Esmailnejad et al., 2017; Sayad et al., 2023).

4. The negative findings regarding Borrelia presence in hosts and ticks

During the period from 2011 to 2012, blood smear examinations of febrile patients in the city of Chabahar revealed no signs of spirochetes associated with Borrelia (Metanat et al., 2014). None of the avian blood samples collected in Lorestan Province showed any signs of infection with B. anserina (Chegeni et al., 2017). In Khorasan Province, PCR testing indicated that there was no evidence of Borrelia spp. infection among 66 patients suffering from morphoea symptoms (Yazdanpanah et al., 2015). Given that small mammals and rodents are considered reservoirs of TBRF and LD disease, it is important to note that, none of the rodents and small mammals in Mazandaran, Gilan, and Golestan Provinces, including Rattus norvegicus, Microtus obscurus, Mus musculus castaneus, Rattus rattus, Apodemus hyrcanicus, Apodemus uralensis, Nesokia indica, Microtus paradoxus, Crocidura caspica, Crocidura suaveolens, and Erinaceus concolor, exhibited evidence of Borrelia spp. presence within their internal organs (Naddaf et al., 2020). In a study involving R. norvegicus gathered from Tehran, antibodies (IgG) against Borrelia spp. were not found (Azimi et al., 2024). Borrelia spp. was not detected in dromedary camels (Camelus dromedarius) assessed in Fars, Kerman, and Sistan and Baluchistan Provinces (Sazmand et al., 2019; Heidari et al., 2023). An investigation was conducted to survey the presence of Borrelia spp. in Iran, focusing on ornamental birds across four provinces: Tehran, East Azerbaijan, Yazd, and Khuzestan. The results indicated that Borrelia was absent in all the birds that were studied (Enferadi et al., 2024). Regarding ticks, no Borrelia was found in these investigations: A. persicus in Qazvin and West Azerbaijan (Barmaki et al., 2010; Enferadi et al., 2024) and I. ricinus in Gilan, Mazandaran, and Golestan (Yousefi-Behzadi et al., 2022).

5. Discussion and conclusion

5.1. Identified Borrelia species regarding TBRF and LD

Our review study found that 20 provinces in Iran are affected by Borrelia species. The geographical distribution map indicates that B. persica and B. burgdorferi are the most prevalent Borrelia species in Iran, respectively. The causative agents of TBRF are chiefly present in the northern regions of Iran. In contrast, B. burgdorferi has been reported in northern provinces, as well as in central, southern, and southwestern regions of Iran. Mazandaran exhibits the highest diversity of Borrelia, where six species of the pathogen have been documented. This notable diversity may be attributed to the province's geographical location and the presence of migratory birds, which have a remarkable ability to host various animal species, especially migratory birds (Jakab et al., 2022), likely contributing to the spread of these pathogens. Moreover, the prolonged attachment and feeding period of hard ticks enhances their ability to migrate over vast distances (Spach et al., 1993; Guberman et al., 1994), facilitating the dissemination of Borrelia species.

5.2. TBRF vectors and hosts

Humans and dogs are reported as dead-end hosts, while M. persicus has been documented as a reservoir in Iran. Considering a documented neonatal case in an endemic region, neonatal borreliosis should be considered in any newborn presenting with symptoms indicative of neonatal sepsis. Additionally, any history of febrile illnesses experienced by the mother during pregnancy, particularly in the final days of gestation, is crucial to consider. For accurate diagnosis, peripheral blood smears from the infant must be obtained at least three times (Mahram and Ghavami, 2009). The transmission of TBRF in Iran is predominantly associated with four species of soft ticks, including O. tholozani, O. erraticus, O. tartokovyskyi, and A. persicus. Surprisingly, hard ticks such as I. ricinus and Rh. annulatus can also transmit TBRF(Naddaf et al., 2020; Milani et al., 2024). Climate change can increase the risk of TBRF by expanding the activity and range of ticks and increasing the number of susceptible hosts (Bouchard et al., 2019).

5.3. LD vectors and hosts

The known dead-end hosts of LD in Iran are humans and dogs. Although B. burgdorferi has been predominantly identified in canine populations, human cases have been reported exclusively in Mazandaran. The incidence of LD in animals has increased over the past several years, specifically from 2009 to 2023. Despite reports of LD cases in six provinces across the country, the investigation of hard tick vectors responsible for the disease have been conducted only in Mazandaran.

5.4. Laboratory methods for Borrelia detection

The diagnosis of TBRF and LD is based on a combination of clinical observations and laboratory tests, none of which alone can definitively confirm infection with Borrelia species (Mead et al., 2024). For TBRF, preparing a thin blood smear followed by microscopic examination is a standard diagnostic method. However, comparative studies have demonstrated that microscopic examination may fail to detect spirochetes in some cases, whereas molecular techniques such as PCR and quantitative PCR (qPCR) can successfully identify their presence (Siadati, 2006). In fact, qPCR has been shown to possess higher sensitivity than traditional PCR, particularly for detecting spirochete DNA at low cycle threshold (Ct) values (Gil et al., 2005; Naddaf et al., 2020; Milani et al., 2024). Therefore, molecular methods could be more reliable techniques for diagnosing TBRF. The use of serological testing for diagnosing TBRF is limited because high seroprevalence does not always indicate an active infection, delayed seroconversion can hinder timely treatment decisions, and assay cross-reactivity complicates distinguishing TBRF from Lyme disease (Magnarelli et al., 1987). In contrast, the diagnosis of LD is primarily depends on serological tests, including ELISA, Western blot, and ICA (Marques,2015).

6. Literature gap and future research

Review studies often highlight gaps in the literature and suggest future research. In this study, we aim to recommend effective field studies to improve understanding and control of tick-borne diseases in Iran. In East Azerbaijan, while M. persicus has been found infected with B. duttonii, ticks in this area have yet to be tested for Borrelia species. Given that infected humans serve as reservoir for B. duttonii, studying local tick populations in this province is crucial. Similarly, an animal case report in West Azerbaijan suggests the presence of Borrelia, but comprehensive investigation into Borrelia species and tick infection are still lacking. In Zanjan Province, the identification of B. persica in O. tholozani remains inconclusive and relies primarily on earlier reports, highlighting the need for updated molecular and phylogenetic studies. Notably, molecular investigations in Hormozgan involving patients with symptoms of TBRF have revealed diverse Borrelia species that cluster into distinct clades. A focused research project on these pathogens has the potential to uncover a novel Borrelia type in Iran. Nanopore-based metagenomic studies have demonstrated higher sensitivity in detecting a wide range of tick-borne viruses and bacteria, enabling the molecular characterization of novel viral and bacterial strains in cases where traditional methods were inadequate or unsuccessful (Ravi et al., 2019; Ergunay et al., 2023; Kipp et al., 2023). Furthermore, in central, southern, and southwestern Iran—regions where LD has been identified in dogs, dedicated studies assessing tick populations are still absent, representing another important gap for future research.

CRediT authorship contribution statement

Parisa Soltan-Alinejad: Writing – review & editing, Writing – original draft, Investigation, Data curation, Conceptualization. Mahmood Nikbakhtzadeh: Writing – review & editing, Visualization, Validation. Eslam Moradi-Asl: Writing – review & editing, Writing – original draft, Software, Methodology.

Ethical approval

The study was conducted by the ethical principles and the national norms and standards for conducting Medical Research in Iran. The study was approved by the Iran National Committee for Ethics in Biomedical Research (Approval ID: IR. ARUMS.REC.1403.483).

Funding

This research was supported by Ardabil University of Medical Sciences (ARUMS), Ardabil, Iran, under Grant No.403000942.

Conflict of interest statement

The authors declare that they have no conflict of interest.

Acknowledgment

Not applicable.

Contributor Information

Parisa Soltan-Alinejad, Email: parisaalinejad1369@gmail.com.

Mahmood Nikbakhtzadeh, Email: mahmood.nikbakhtzadeh@csusb.edu.

Eslam Moradi-Asl, Email: Moradiasl83@yahoo.com, e.moradiasl@arums.ac.ir.

Data availability

All data generated this study are available upon reasonable request to the corresponding author.

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Associated Data

This section collects any data citations, data availability statements, or supplementary materials included in this article.

Data Availability Statement

All data generated this study are available upon reasonable request to the corresponding author.

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