The most unusual traits exhibited by this genus, however, are found on the [[Labrum (arthropod mouthpart)|labrum]] and the telopodites of the second maxillae. The labrum features two rows of teeth, not only a posterior series of external teeth but also a dorsal series of small internal teeth.<ref name=”:02″ /> Furthermore, each telopodite of the second maxillae features an additional (fourth) article that is covered with small scales and lacks a claw at the distal end.<ref name=”Bonato-2011″ /><ref name=”:02″ /> The coxosternite of the second maxillae is not separated or divided by a longitudinal furrow down the middle.<ref name=”:02″ />
The most unusual traits exhibited by this genus, however, are found on the [[Labrum (arthropod mouthpart)|labrum]] and the telopodites of the second maxillae. The labrum features two rows of teeth, not only a posterior series of external teeth but also a dorsal series of small internal teeth.<ref name=”:02″ /> Furthermore, each telopodite of the second maxillae features an additional (fourth) article that is covered with small scales and lacks a claw at the distal end.<ref name=”Bonato-2011″ /><ref name=”:02″ /> The coxosternite of the second maxillae is not separated or divided by a longitudinal furrow down the middle.<ref name=”:02″ />
The [[Forcipule|forcipular]] coxosternite is short and lacks [[Denticle (tooth feature)|denticles]] or [[chitin]] lines, and the [[Forcipule|forcipules]] are also short and lack denticles. The ultimate article of the forcipule is flattened and relatively elongated. The [[Sternum (arthropod anatomy)|sternites]] feature pores arranged in a large field in the posterior part of each segment. The basal element of each of the [[ultimate legs]] (coxopleuron) features numerous pores that open separately and are scattered over almost the entire surface. Each of the ultimate legs features five articles with a claw at the distal end.<ref name=”:02″ /><ref name=”Bonato-2011″ /><ref name=”:7″ />
== References ==
== References ==
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Macronicophilus is a genus of soil centipedes in the family Geophilidae. This genus contains only four valid species.[1][2] These centipedes are found in the northern Andes and the Amazon basin in South America.[3]
Discovery and distribution
[edit]
This genus was created by the Italian zoologist Filippo Silvestri in 1909 to contain the newly discovered type species M. ortonedae. He based the original description of this species on a sample of specimens including both sexes. These specimens were found in Naranjito, near Guayaquil, in Ecuador.[4] In 1912, the French naturalist Henri Ribaut redescribed this species based on three male specimens found near Angelopolis in Colombia.[5] This species is known from only these two localities.[6]
In 2000, the biologists Luis Alberto Pereira, Donatella Foddai, and Alessandro Minelli described three more species in this genus. They described M. abbreviatus based on an examination of a male holotype, a female allotype, and three other specimens (two females and one male). These specimens were collected in 1986 to 1991 from three different sites in the state of Amazonas in Brazil.[6] This species is known from only these three localities.[7] The holotype and allotype are deposited in the lnstituto Nacional de Pesquisas da Amazonia in the city of Manaus in Brazil.[6]
Pereira, Foddai, and Minelli described M. unguiseta based on a male holotype found in 1982 in the state of Amazonas in Brazil. This holotype is deposited in the lnstituto Nacional de Pesquisas da Amazonia.[6] This species is known from only this holotype.[7]
Pereira, Foddai, and Minelli described M. venezolanus based on an examination of a female holotype, a male allotype, and a juvenile specimen. These specimens were collected in 1987 from three different sites in Venezuela. The allotype is deposited in the Museo de La Plata in the city of La Plata in Argentina.[6] In 2014 and 2015, nine specimens of a species of Macronicophilus were collected near Icononzo in Colombia.[8] In 2023, the biologists Lucio Bonato and Rodrigo Lopes Ferreira identified these nine specimens as M. venezolanus. This species is known from only these twelve specimens collected from the Venezuelan Coastal Range and the Colombian Andes.[7]
Puzzled by the idiosyncratic traits observed in this genus, taxonomists have struggled to the place this enigmatic genus in the appropriate family. In 1909, Silvestri originally placed this genus in the small family Gonibregmatidae.[4][6] Other authorities later placed this genus in the family Geophilidae instead.[9][6] In 2000, Pereira, Foddai, and Minelli proposed a separate monotypic family Macronicophilidae to contain this genus,[6] and authorities adopted this proposal.[3][10] In 2014, however, a phylogenetic analysis of the order Geophilomorpha using both morphological and molecular data found a representative of this genus (M. venezolanus) nested within the family Geophilidae. To avoid paraphyly of the family Geophilidae with respect to Macronicophilidae, authorities dismissed Macronicophilidae as a separate family.[11] Authorities now place the genus Macronicophilus in the family Geophilidae,[1][2] but some references continue to place this genus in the family Macronicophilidae.[10]
In 2023, a phylogenetic analysis of the family Geophilidae based on morphology confirmed the monophyly of the genus Macronicophilus, placing all four species together in a clade in a phylogenetic tree of this family. These four species also formed a sister group for the newly described monotypic genus Plutogeophilus, which emerged as the genus most closely related to Macronicophilus in this analysis.[7] In 2014, phylogenetic analysis using both morphology and molecular evidence in placed M. venezolanus in a clade with a representative of Eurygeophilus.[11] The results of these studies suggests that after the genus Plutogeophilus, the genus Eurygeophilus may be the next closest relative of the genus Macronicophilus.
Centipedes in the genus Macronicophilus feature a short head and antennae that are slightly attenuated.[3] The mandible features a pectinate lamella with a single row of less than 20 short teeth that are shaped like triangles.[6][3][7] The telopodite of the first maxillae features a single article, with not only a lappet projecting from the lateral margin but also a second lappet projecting from the dorsal surface. The lappets of the coxosternite of the first maxillae are either absent or rudimentary.[6][12]
The most unusual traits exhibited by this genus, however, are found on the labrum and the telopodites of the second maxillae. The labrum features two rows of teeth, not only a posterior series of external teeth but also a dorsal series of small internal teeth.[6] Furthermore, each telopodite of the second maxillae features an additional (fourth) article that is covered with small scales and lacks a claw at the distal end.[3][6] The coxosternite of the second maxillae is not separated or divided by a longitudinal furrow down the middle.[6]
The forcipular coxosternite is short and lacks denticles or chitin lines, and the forcipules are also short and lack denticles. The ultimate article of the forcipule is flattened and relatively elongated. The sternites feature pores arranged in a large field in the posterior part of each segment. The basal element of each of the ultimate legs (coxopleuron) features numerous pores that open separately and are scattered over almost the entire surface. Each of the ultimate legs features five articles with a claw at the distal end.[6][3][12]
- ^ a b Bonato, L.; Chagas Junior, A.; Edgecombe, G.D.; Lewis, J.G.E.; Minelli, A.; Pereira, L.A.; Shelley, R.M.; Stoev, P.; Zapparoli, M. (2016). “Alloschizotaenia Brölemann, 1909”. ChiloBase 2.0 – A World Catalogue of Centipedes (Chilopoda). Retrieved 2024-06-08.
- ^ a b “ITIS – Report: Macronicophilus”. itis.gov. Retrieved 2025-12-06.
- ^ a b c d e f Bonato, Lucio; Edgecombe, Gregory D.; Zapparoli, Marzio (2011). “Chilopoda – Taxonomic overview”. In Minelli, Alessandro (ed.). The Myriapoda. Volume 1. Leiden: Brill. pp. 363–443 [432]. ISBN 978-90-04-18826-6. OCLC 812207443.
- ^ a b Silvestri, F. (1909). “Descrizioni preliminari di vari artropodi specialmente d’America” (PDF). Rendiconti della R. Accademia dei Lincei. Classe di Scienze Fisiche Matematiche e Naturali (in Italian). 18 (1): 267-271 [267-268].
- ^ Ribaut, Henri (1912). “Contribution a l’etude des Chilopodes de Colombie”. Mémoires de la Société des Sciences Naturelles de Neuchâtel (in French). 5: 67-95 [87] – via Biodiversity Heritage Library.
- ^ a b c d e f g h i j k l m n Pereira, Luis Alberto; Foddai, Donatella; Minelli, Alessandro (2000). “New taxa of Neotropical Geophilomorpha (Chilopoda)”. Amazoniana. 16 (1–2): 1-57 [15-21]. ISSN 0065-6755.
- ^ a b c d e Bonato, Lucio; Ferreira, Rodrigo Lopes (2023-12-01). “Assessing troglomorphic and phylogenetically informative traits in troglobionts: a new cave-dwelling centipede illuminates the evolution of a soil-dwelling lineage (Chilopoda: Geophilidae)”. Organisms Diversity & Evolution. 23 (4): 833–856. doi:10.1007/s13127-023-00618-7. ISSN 1618-1077.
- ^ Prado-Sepúlveda, Camilo; Triana, Hernán Darío; Galvis Jiménez, Sebastián (2016). “Los ciempiés (Myriapoda: Chilopoda) de bosque Andino en el municipio de Icononzo (Colombia, Tolima) y clave para las familias presentes en Colombia”. Boletín de la Sociedad Entomológica Aragonesa (in Spanish). 58: 188–196 [191].
- ^ Attems, Carl (1929). Attems, Karl (ed.). Lfg. 52 Myriapoda, 1: Geophilomorpha (in German). De Gruyter. p. 276. doi:10.1515/9783111430638. ISBN 978-3-11-143063-8.
- ^ a b “Macronicophilus Silvestri, 1909”. www.gbif.org. Retrieved 2025-12-06.
- ^ a b Bonato, Lucio; Drago, Leandro; Murienne, Jerome (2014). “Phylogeny of Geophilomorpha (Chilopoda) inferred from new morphological and molecular evidence”. Cladistics. The International Journal of the Willi Hennig Society. 30 (5): 485–507. doi:10.1111/cla.12060. PMID 34794246. S2CID 86204188.
- ^ a b Bonato, Lucio; Edgecombe, Gregory; Lewis, John; Minelli, Alessandro; Pereira, Luis; Shelley, Rowland; Zapparoli, Marzio (2010-11-18). “A common terminology for the external anatomy of centipedes (Chilopoda)”. ZooKeys (69): 17–51. Bibcode:2010ZooK…69…17B. doi:10.3897/zookeys.69.737. ISSN 1313-2970. PMC 3088443. PMID 21594038.
