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Symphyla
Symphyla
Symphyla
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Not to be confused with Symphyta.

Symphyla
Temporal range: 99–0 Ma Mid Cretaceous to Present
Scutigerella sp. (Scutigerellidae)
Scolopendrellid symphylan (Scolopendrellidae)
Scientific classification Edit this classification
Kingdom: Animalia
Phylum: Arthropoda
Subphylum: Myriapoda
Class: Symphyla
Ryder, 1880
Families

Scutigerellidae
Scolopendrellidae

Symphylans, also known as garden centipedes or pseudocentipedes, are soil-dwelling arthropods of the class Symphyla in the subphylum Myriapoda. Symphylans resemble centipedes but are very small, non-venomous, and may or may not form a clade with centipedes.[1][2] More than 200 species are known worldwide.[3]

Symphyla are primarily herbivores and detritus feeders living deep in the soil, under stones, in decaying wood, and in other moist places.[4] They are rapid runners,[4] can move quickly through the pores between soil particles, and are typically found from the surface down to a depth of about 50 centimetres (20 in). They consume decaying vegetation but can do considerable harm in an agricultural setting by consuming seeds, roots, and root hairs in cultivated soil.[1][2] For example, the garden symphylan, Scutigerella immaculata can be a pest of crops. A species of Hanseniella has been recorded as a pest of sugar cane and pineapples in Queensland.[5][6] A few species are found in trees[7][8] and in caves.[9] A species of Symphylella has been shown to be predominantly predatory,[10] and some species are saprophagous.

Description

[edit]
Several individuals of Scutigerella sp., showing their small size

Symphyla are small, cryptic myriapods without eyes and without pigment.[4] The body is soft and generally 2 to 10 millimetres (0.08 to 0.4 in) long, divided into two body regions: head and trunk.[4] An exceptional size is reached in Hanseniella magna, which attains lengths of 12-13 mm (0.5 in).[11]

The head has long, segmented antennae, a postantennal organ, three pairs of mouthparts: mandibles, the long first maxillae, and the second pair of maxillae which are fused to form the lower lip or labium of the mouth. The antennae serve as sense organs. Disc-like organs of Tömösváry, which probably sense vibrations, are attached to the base of the antennae, as they are in centipedes.[2]

The trunk comprises 14 segments, which are covered by microhairs on the lateral and ventral integument and by a various number of dorsal tergal plates, from 15 in Scutigerella and Hanseniella, and up till 24 in Ribautiella, increasing the flexibility of the body. Legs are found on the first 12 segments. The 13th segment, which is fused with the 12th segment, bears a pair of spinnerets that resemble cerci, and the 14th segment has a pair of long sensory hairs (trichobothria). Around the anal opening there is a small telson.[12][13][14][15] Symphylans have been reported as living up to four years, and moult throughout their life.[2] Immature individuals have six or seven pairs of legs on hatching, but they add an additional pair at each moult until the adult instar, which usually has twelve pairs of legs.[16][17] This mode of development is known as hemianamorphosis.[18] Although most adult symphylans have twelve leg pairs, the first pair is absent or vestigial in some species (e.g., those in the genus Symphylella), so adults in some species have only eleven leg pairs.[19][20] The species with 12 pairs are the only myriapods with walking legs on the first body segment, as the first pair of legs are modified into forcipules in centipedes, in pauropods this segment is a reduced collum which bears ventrally a pair of small papillae, and in millipedes this segment is a collum without any appendages at all.[21]

Symphylans have several features linking them to early insects, such as a labium (fused second maxillae), an identical number of head segments, and certain features of their legs.[1] Each pair of legs is associated with an eversible structure called a "coxal sac", which helps the animal absorb moisture, and a small stylus that may be sensory in function. Similar structures are found in the most primitive insects.

Life stages of symphylans: eggs, juvenile, and adult Scutigerella immaculata

Symphylans breathe through a pair of spiracles on the sides of their head and are the only arthropods with spiracle openings on the head.[22] These are connected to a system of tracheae that branch through the head and the first three segments of the body only.[2]

The genital openings are located on the fourth body segment, but the animals do not copulate. Instead, the male deposits 150 to 450 packages of sperm, or spermatophores, on small stalks. The female then picks these up in her mouth, which contains special pouches for storing the sperm. She then lays her eggs, and attaches them to the sides of crevices or to moss or lichen with her mouth, smearing the sperm over them as she does so. The eggs are laid in groups of eight to twelve.[2]

The spinnerets produce secretions that turn into a silk-like thread.[23] One fossil species, Symphylella patrickmuelleri, was found preserved in Burmese Amber releasing long threads of silk.[24] The silk plays a role in reproduction: the male deposits up to 450 spermatophores on stalks of silk.[25] Symphylans have also been reported releasing silk as a defense[26] and to suspend themselves in the air.[2]

Fossil record and evolution

[edit]

The symphylan fossil record is poorly known, with only five species recorded, all placed in living genera. The oldest records of both families are found in Burmese amber from the middle Cretaceous, approximately 99 million years ago. As a result, both families are thought to have diverged before the end of the Mesozoic Era.[27][28][29]

Despite their common name, morphological studies commonly place symphylans as more closely related to millipedes and pauropods than the centipedes, in the clade Progoneata.[30][31] Molecular studies have shown conflicting results, with some supporting the Progoneata clade, others aligning symphylans with centipedes or other arthropods, although some are weakly supported.[32][30] The clade is believed to be monophyletic.[33]

References

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Symphyla

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Symphyla is a class of small, soil-dwelling arthropods in the subphylum and phylum Arthropoda, characterized by their elongated, soft, white bodies measuring 2–8 mm in length, lack of eyes, prominent antennae, and 12 pairs of legs in adults. These eyeless creatures, often called garden centipedes or pseudocentipedes, inhabit moist soils worldwide and are known from approximately 200 , primarily in the families Scutigerellidae and Scolopendrellidae. Symphylans exhibit anamorphic development, hatching with six pairs of legs and adding more with each molt until reaching 12 pairs as adults, which possess 14 trunk segments, of which the first 12 bear a pair of legs each. They are omnivorous, feeding on decaying organic matter, fungi, plant roots, and small invertebrates, and can become agricultural pests by damaging seedlings and root crops in high-organic-matter soils. Despite their centipede-like appearance, symphylans lack venomous forcipules and are generally harmless to humans, playing a key role in soil ecosystems as decomposers and predators of microscopic organisms. Fossil evidence suggests symphylans diverged from other myriapods around 430–593 million years ago, making them one of the most ancient extant classes within , alongside Chilopoda (centipedes), Diplopoda (millipedes), and . Their global distribution spans diverse habitats from tropical to temperate regions, though they thrive in humid, litter-rich environments and are rarely found in arid areas. Research on symphylans remains limited due to their cryptic lifestyle, but studies highlight their ecological importance in nutrient cycling and as indicators of .

Taxonomy and classification

Etymology

The class Symphyla was established by the American zoologist in 1880, who proposed it as a new order of articulates based on specimens of the genus Scolopendrella, emphasizing their unique morphological features that distinguished them from other known . The name "Symphyla" derives from the Greek prefix syn- (together or with) and phyla (tribe or class), literally meaning "uniting the tribes" or "those that unite classes," a reference to Ryder's interpretation of these organisms as transitional forms bridging and myriapods. This underscores the historical perception of Symphyla as annectant taxa in evolution. In the broader context of 19th-century myriapod taxonomy, Ryder's introduction of Symphyla built upon earlier classifications, such as Latreille's establishment of Myriapoda in 1802 and Lubbock's description of Pauropoda in 1868, reflecting growing efforts to categorize diverse, often overlooked soil arthropods through comparative anatomy and emerging evolutionary ideas.

Systematic position

Symphyla is classified within the kingdom Animalia, phylum Arthropoda, subphylum Myriapoda, and class Symphyla. This placement positions Symphyla as one of four extant classes in Myriapoda, alongside Chilopoda (centipedes), Diplopoda (millipedes), and Pauropoda (pauropods), reflecting their shared arthropod ancestry but distinct evolutionary trajectories within the subphylum. The class Symphyla comprises two families: Scutigerellidae and Scolopendrellidae, which together encompass 14 genera and over 200 described species worldwide. The family Scutigerellidae includes genera such as Scutigerella, with representative species like Scutigerella immaculata Newport, 1845, a cosmopolitan soil-dweller often studied for its agricultural impact. In contrast, Scolopendrellidae features genera like Scolopendrella and Symphylella, accommodating a broader diversity of smaller, less economically significant forms. Recent taxonomic efforts have expanded the known diversity, particularly in underrepresented regions. For instance, in 2023, Jin and Bu described Scutigerella sinensis, the first species of its genus from , based on specimens from Hainan Province, highlighting ongoing discoveries in tropical and subtropical soils. Similarly, that year, two new Symphylella species—S. macrochaeta and S. longispina—were added from Tibetan localities, underscoring the class's cryptic distribution and the need for further surveys. In 2025, four additional species of Symphylella were described from Chongqing, southwest , further illustrating the continued expansion of symphylan taxonomy in the region. Unlike the predatory Chilopoda or the detritivorous Diplopoda, Symphyla represent a specialized lineage adapted to soil microhabitats, maintaining their status as a monophyletic class distinct in myriapod systematics.

Morphology and physiology

External anatomy

Symphyla exhibit a soft, elongated, unpigmented body adapted to subterranean habitats, typically measuring 2 to 8 mm in length, with the exception of Hanseniella magna, which attains 25 to 30 mm. The body lacks eyes and pigment, consisting of a distinct, cordiform head and a trunk of 14 segments, the first 12 of which bear legs in most species, while some exhibit 11 pairs. The trunk is covered dorsally by tergites (scuta), numbering 15 to 24 due to segmental subdivisions that vary by family and genus. The head is strongly sclerotized dorsally but weakly ventrally, featuring long, filiform, moniliform antennae that exceed the head length and bear genus- or species-specific sensory hairs for environmental perception. Tömösváry's organs, located at the base of the antennae, serve as hygroreceptors. The trunk legs are slender and thread-like, each comprising a coxa, prefemur, femur, tibia, and tarsus, with a main claw accompanied by a smaller secondary claw; the first pair is often reduced to less than half the length of subsequent pairs or modified into hairy knobs in certain genera like Symphylella. These legs facilitate navigation through soil pores, with styli present on pairs 3 through 12 and eversible vesicles positioned between the legs for sensory or adhesive functions. At the posterior end, the pre-anal segment bears a pair of large, conical spinnerets associated with sericigenous glands for production, while the anal segment features trichobothria with sensory hairs. Paired conical cerci, varying in ornamentation such as pubescence or scales across genera (e.g., longitudinal ridges in Scolopendrella), are positioned at the rear and linked to these spinning structures. The overall external morphology, studied via in diverse species, underscores adaptations for a cryptic, soil-dwelling .

Internal anatomy and respiration

The internal anatomy of symphylans is adapted to their subterranean lifestyle, featuring a soft, flexible body that facilitates movement through soil. In species that possess them, the respiratory system consists of a single pair of spiracles located anterior to the mandibles on the head, a position unique among arthropods. These spiracles connect to a tracheal system with a dorsal stem that branches to supply oxygen to the head muscles, nervous system, and anterior trunk segments, while anastomoses between opposite sides occur ventrally under the pharynx; in species lacking spiracles, gas exchange occurs primarily through the highly permeable cuticle due to the animals' small size and soft integument. The digestive system forms a simple, tubular alimentary canal suited for processing and plant material. It comprises an ectodermal with a short, rectangular and a posterior , an endodermal lined with cuboidal to columnar cells for nutrient absorption, and an ectodermal that includes a , , muscular colon, and thin-walled for waste elimination. The is open, typical of arthropods, with a tubular heart extending from the third trunk segment to the pre-anal segment and featuring lateral ostia beginning from the sixth segment to allow entry. An anterior runs alongside the into the head, where it branches into cephalic arteries that envelop the maxillary gland sacculi, and a supraneural vessel provides a ventral circulatory pathway. The includes a centralized composed of protocerebrum, deutocerebrum, and tritocerebrum, continuous with a subesophageal nerve mass and a ventral nerve cord featuring segmental ganglia. The protocerebrum innervates the postantennal organ, the deutocerebrum supplies sensory and motor fibers to the antennae, and the tritocerebrum links to the second maxillae; a stomatogastric system forms a nerve bridge around the with a recurrent nerve along the dorsal midline. Symphylans lack eyes, relying instead on their long, filiform antennae for chemosensory and in dark environments.

Life cycle and reproduction

Development

Symphyla exhibit hemianamorphic development, a post-embryonic growth pattern typical of many myriapods in which juveniles progressively add body segments and appendages through molting without undergoing a distinct metamorphosis. Newly hatched nymphs possess 6 pairs of legs and a reduced number of body segments compared to adults. With each subsequent molt, one additional pair of legs and associated segments is added, resulting in 12 pairs by maturity after approximately 6 molts, corresponding to 7 instars in total. This gradual addition continues beyond sexual maturity, as adults undergo further molts throughout their lifespan, potentially exceeding 40–50 times in total. The stage precedes this hemianamorphic phase, with females laying clutches of 4–25 , often in clusters within crevices or . These pearly white, spherical , marked by hexagonal ridges, are guarded by the female, providing limited during incubation. Hatching occurs after 25–40 days at temperatures of 50–70°F (10–21°C), though this period shortens to about 12 days above 77°F (25°C); overall development from to reproductive adult takes 3–5 months depending on environmental conditions. Juveniles differ from adults primarily in size and segment count, growing from 1–2 mm at to 8 mm or more in mature individuals, with no abrupt morphological shifts. Symphyla lifespan extends up to 4 years in favorable conditions, allowing multiple reproductive cycles supported by ongoing molting for maintenance and growth.

Reproductive behavior

Symphylans exhibit indirect transfer, a characteristic mating behavior in which males deposit stalked spermatophores in the without direct contact with females. These spermatophores, consisting of drops attached to stalks, are produced in batches, with males capable of depositing up to 450 such packages. Females locate and uptake the spermatophores by biting them off the stalks and storing the in specialized gnathal pockets within the region; the eggs are laid unfertilized and then externally fertilized by smearing the stored on them shortly after deposition. This process ensures viability in the moist environment typical of symphylan habitats. Silk plays a key role in symphylan reproduction, primarily produced by spinnerets located on projections of the next-to-last body segment to form the stalks that support spermatophores above the surface, facilitating female detection and uptake. Eggs, typically laid in groups of 4 to 25, are smeared with residual for protection. Parental care in Symphyla is provided exclusively by females, who guard clusters until , a period of approximately 1 to 3 weeks depending on (e.g., 2-3 weeks at 15-20°C). This brooding prevents and inhibits fungal overgrowth on the eggs; experimental removal of the female results in high egg mortality, with most failing to hatch due to microbial . in symphylans is attained after 4 to 6 months under conditions, allowing adults to engage in 2 to 3 reproductive cycles over their lifespan, which can extend up to 4 years in . This limited reproductive output aligns with their soil-dwelling lifestyle, where environmental constraints like and regulate breeding frequency.

Ecology and behavior

Habitat and distribution

Symphyla are primarily soil-dwelling myriapods, inhabiting the upper layers at depths typically ranging from 0 to 50 cm, though some can be found deeper than 90 cm during certain conditions. They prefer moist, organic-rich soils with high water-holding capacity, such as loamy or clay types, often occurring under stones, logs, litter, in dead wood, , or heaps. These arthropods avoid heavy, peaty, compacted, sandy, or excessively wet soils, favoring well-aggregated substrates that provide crevices and stable levels, with optimal pH ranges of 4.5 to 7.5. Their vertical distribution in the soil profile is dynamic, influenced by environmental factors; they aggregate in the top 15 cm during warm, moist periods like spring rains and migrate deeper in response to or extremes. While most Symphyla are terrestrial and confined to microhabitats, where they navigate through pores and fine channels for movement and refuge, a few species exhibit specialized habitats. Certain taxa, such as Hanseniella arborea in Amazonian blackwater inundation forests, display arboreal migrations, ascending trunks to heights of up to 3.6 m on species like Aldina heterophylla during seasonal flooding to avoid submersion. Cave-dwelling forms also exist, including Symphylella major and Scutigerella immaculata, which occupy moist, cool systems and mines, adapting to stable humidity in subterranean environments. Symphyla exhibit a , present across all continents except and absent from extreme deserts or polar regions due to their requirements. They achieve greatest abundance and diversity in temperate and tropical climates, with over 200 described worldwide; for instance, 18 taxa are recorded in alone, spanning forests, meadows, pastures, gardens, and arable lands. Their dependency directly supports respiratory needs, as the thin allows efficient but demands consistently humid conditions to prevent .

Diet and feeding

Symphyla are primarily detritivores and herbivores, feeding on decaying matter, fungi, and root exudates in the soil. They consume organic debris such as leaf litter and decomposing , which supports their role in breaking down soil organic material. Species like Scutigerella immaculata, the garden symphylan, also actively feed on live tissues, including germinating , seedlings, root hairs, and underground plant parts, often causing pitting damage to . While most symphylans exhibit detritivorous or herbivorous habits, some display predatory behavior. For instance, members of the genus Symphylella have been observed preying on soil nematodes, capturing and consuming them as part of their diet. This omnivorous flexibility allows certain symphylans to exploit a range of food sources depending on availability in the soil environment. Symphylans employ chewing mouthparts to ingest food, rasping and biting into softer materials like root tissues or fungal hyphae, and they may rapidly ingest soil particles to filter out organic components. In their foraging, they often move toward moist root zones, where adequate facilitates access to food resources. As soil decomposers, symphylans play a key trophic role in nutrient cycling by fragmenting organic matter and promoting microbial activity, thereby enhancing soil fertility in natural ecosystems. However, their feeding on crop roots can lead to occasional agricultural damage, particularly in high-organic-matter soils where populations thrive.

Evolutionary history

Fossil record

The fossil record of Symphyla is notably sparse, with only four described documented to date, reflecting the challenges posed by their small size and soft-bodied morphology that rarely fossilize outside of exceptional preservation conditions such as inclusions. These fossils span from the mid-Cretaceous to the present, providing limited insights into the group's evolutionary history despite molecular estimates suggesting an ancient divergence from other myriapods ca. 480 million years ago (latest –Early ). The oldest known symphylans are from Cretaceous Burmese amber, dated to approximately 99 million years ago, including the species Symphylella patrickmuelleri, the first fossil representative of the family Scolopendrellidae and the earliest record of the class overall. This juvenile specimen exhibits morphological features akin to modern Symphylella species, such as 17 tergites, reduced leg styli, and elongated antennae and legs, preserved alongside silk threads extruded from its spinnerets. Additional Burmese amber records include two Scutigerellidae specimens, further highlighting the mid- presence of both extant symphylan families. Subsequent fossils are younger and also amber-bound: two species from Eocene (Scutigerella baltica and Hanseniella baltica, approximately 44–54 million years old) and one from (Scutigerella dominicana, 15–20 million years old), all belonging to the Scutigerellidae. The absence of pre-Cretaceous symphylan fossils may indicate undersampling in earlier deposits rather than a late origin, as their delicate exoskeletons and soil-dwelling habits limit preservation to rare resin entrapments.

Phylogenetic relationships

Symphyla are classified within the Progoneata clade, a monophyletic group comprising Symphyla, , and Diplopoda, positioning them closer to pauropods and millipedes than to centipedes (Chilopoda). This affiliation is primarily supported by morphological synapomorphies, including a single pair of spiracles located on the head capsule and patterns of leg development that align Symphyla with Dignatha ( + Diplopoda), such as the anterior positioning of gonopores and similarities in trunk tagmosis. Molecular evidence has bolstered this placement, with analyses of 18S rRNA genes indicating a close relationship between Symphyla, , and Diplopoda, forming the Progoneata. Phylogenomic studies using transcriptomes and multiple nuclear genes further corroborate the of Progoneata, often recovering Symphyla as sister to (Edafopoda) or to Dignatha, with robust support from concatenated datasets of hundreds of loci. Phylogenetic debates have historically linked Symphyla to Chilopoda due to shared anamorphic post-embryonic growth, where segments and legs are added after hatching, contrasting with the euanamorphic development of Diplopoda. However, comprehensive phylogenomic analyses in the , incorporating quartet-based methods and extensive sampling, have resolved these conflicts in favor of Progoneata by mitigating long-branch attraction and incorporating morphological constraints. Symphyla evolved from ancestors shared with other myriapods, with their modern diversity arising from a radiation during the period, coinciding with the diversification of angiosperms and ecosystems. The record provides evidence for the ancient origins of , from which Symphyla diverged.

Relationship to humans

As agricultural pests

Symphylans, particularly species in the genera Scutigerella and Hanseniella, are recognized as significant agricultural pests due to their root-feeding habits that damage a variety of crops worldwide. The garden symphylan, Scutigerella immaculata, is a primary pest in temperate regions, targeting seedlings and young in greenhouses and field settings, including strawberries, , , cole crops, , corn, soybeans, potatoes, beets, and carrots. In tropical areas, Hanseniella ivorensis emerges as a key pest, severely affecting pineapple roots in regions like Côte d'Ivoire and , while other symphylans impact in , , and . These pests cause damage by chewing on root hairs, rootlets, and , leading to , reduced nutrient uptake, and plant death, especially in seedlings and transplants. In pineapples, H. ivorensis results in decayed roots and impaired , often exacerbating issues when combined with nematodes. Outbreaks are favored in moist s rich in , such as heavy clay types with good structure, where symphylans thrive and proliferate rapidly above 45°F temperatures. These conditions enable localized that spread gradually, up to 10-20 feet annually in affected fields. Control strategies emphasize integrated approaches to manage populations without relying solely on chemicals. Cultural methods include improving drainage through intensive , avoiding compacted or sandy soils, and periodic flooding for 2-3 weeks to reduce numbers, particularly in late spring or summer. Biological controls leverage natural predators such as centipedes, predatory mites, ground beetles, and entomopathogenic fungi, though their efficacy varies in field conditions. Recent projects as of 2025 explore biopesticides and crop rotations for , particularly in and grass seed production. Chemical options involve pre-planting applications of insecticides like or zeta-cypermethrin for spot treatments, while in tropical systems, nematicides may indirectly help but require targeted strategies for symphylids. The economic impact of symphylan pests is substantial, with untreated infestations leading to total production losses in affected areas and severe yield reductions in reduced-input farming systems across , particularly in , , Washington, and . In tropical regions, damage to and contributes to ongoing challenges in crop establishment and productivity, underscoring the need for vigilant monitoring in high-organic-matter soils.

In scientific research

Symphyla serve as important model organisms in microarthropod , particularly for investigating processes and dynamics in terrestrial ecosystems. Studies have highlighted their role in nutrient cycling through consumption of decaying , which contributes to and litter breakdown. For instance, in experiments examining community assembly at micro-spatial scales, symphylans are included among key microarthropods that influence fungal communities and turnover, with higher densities correlating to increased fungal diversity but stable rates. Post-2020 using controlled s has demonstrated how symphylan populations respond to environmental factors like and organic content, underscoring their utility in modeling responses to habitat changes. In evolutionary research, Symphyla have been instrumental in phylogenomic analyses to clarify myriapod relationships. Transcriptomic and mitogenomic studies position Symphyla as the sister group to within , resolving long-standing debates on phylogeny through multi-gene datasets. A notable application is , which facilitated the 2023 discovery of Scutigerella sinensis, the first of the genus reported from , collected from forest s in and ; its COI barcode sequence showed significant divergence (27.47%) from related taxa, aiding delimitation in understudied soil myriapods. In November 2025, four new of Symphylella were described from , southwest , using similar approaches to assess genetic divergences. Conservation efforts for Symphyla emphasize the vulnerability of undescribed species in tropical s, where poses a major to . While no Symphyla species hold formal status, reviews of myriapod conservation indicate that many tropical , including symphylans, remain undescribed and are at risk from loss, with calls for protective measures in biodiverse regions like Amazonian forests. Experimental studies on tropical show declines in diversity under land-use pressures analogous to , highlighting the need for preservation to safeguard these cryptic groups. Laboratory rearing protocols for Symphyla enable detailed studies of their development and , typically involving moist substrates enriched with decaying leaves or to mimic natural conditions. Established methods, such as those using jars with 30% moisture silt loam and bases, support and observation of life cycles, with symphylans thriving at temperatures of 15–21°C. These cultures have been used to track developmental stages and reproductive peaks, providing insights into soil-dwelling adaptations. Pest symphylan species are occasionally studied in laboratories to evaluate control efficacy, such as through baiting or chemical assays.

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  49. https://www.researchgate.net/publication/341068156_Conservation_of_terrestrial_invertebrates_a_review_of_IUCN_and_regional_Red_Lists_for_Myriapoda
  50. https://pmc.ncbi.nlm.nih.gov/articles/PMC11639662/
  51. https://www.cabidigitallibrary.org/doi/pdf/10.5555/20063069230
  52. https://academic.oup.com/jee/article/64/3/657/824857
  53. https://ir.library.oregonstate.edu/downloads/r781wg181
  54. https://ir.library.oregonstate.edu/downloads/73666829j
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