Research Article
Research Article
Notorhinus floresi sp. nov. gen. nov.: The first records of Siphonophorida in Chile and Siphonorhinidae in South America (Colobognatha)
expand article infoLeif Moritz§, Antonio Parra-Gómez|
‡ Zoological Research Museum Alexander Koenig, Bonn, Germany
§ University of Bonn, Bonn, Germany
| Universidad Austral de Chile, Valdivia, Chile
Open Access


The millipede family Siphonorhinidae (order Siphonophorida) shows a scattered distribution in South Africa, Madagascar, India, Southeast Asia, and North America. So far, the family is unknown from South America, while species of Siphonophoridae, the second family of the order, are relatively abundant on the continent. However, not a single Siphonophorida is known from Chile. Here we describe the monotypic genus Notorhinus gen. nov. with N. floresi sp. nov. and record a second Notorhinus (undescribed) species, as first records of the order Siphonophorida in Chile and of the family Siphonorhinidae in South America. Notorhinus gen. nov. is distinct from the remaining Siphonorhinidae by the arrangement of the sensilla basiconica on the antennae and other somatic and sexual characters. However, it shows close morphological affinities to the North American genus Illacme Cook and Loomis 1928. In the Americas Siphonorhinidae were previously only known from California (USA), where they inhabit subterranean micro-habitats. The Chilean species was found under a piece of decaying wood in a small patch of fragmented native forest. Thus, the group shows a disjunct antitropical distribution in America at ca. 37° North and 38° South. They might be the relict of a once greater distribution, which persisted in these areas due to similar climatic conditions.


antitropical, Biobio, endemic, millipede, new genus, new species

1. Introduction

Among millipedes (Diplopoda) the Colobognatha, with the four orders Platydesmida, Polyzoniida, Siphonocryptida and Siphonophorida, are extraordinary in many aspects. Colobognathan taxa show strongly modified heads and mouthparts for suctorial feeding (Moritz et al. 2021, 2022), some display maternal or paternal brood-care (Wong et al. 2020; Moritz et al. 2023), and they exhibit a fragmented and relictual distribution in widely sepa­rated areas (Cook and Loomis 1928; Shelley and Golovatch 2011). The interesting biogeography of the group is probably the relict of a once wider distribution (Marek et al. 2016) and might be related to their adaptation towards certain vegetation and climate, as the Colobognatha show greater ecological limitations than other Diplopoda (Cook and Loomis 1928) and are often restricted to very humid habitats (Golovatch et al. 2015). Prime examples of this scattered distribution can be found in the order Siphonophorida and especially in the family Siphonorhinidae (Fig. 1A). The Siphonophorida are characterized by separate tergites, pleurites and sternites, modification of male leg-pair 9 and 10 to leg-like gonopods, absence of ommatidia, a head, which is conical/pyriform or drawn out into a beak, and strongly modified mouthparts (Read and Enghoff 2009). The order comprises a total of 118 described species to date (Enghoff et al. 2015; Marek et al. 2016; Wesener 2023), which are classified in two families: Siphonophoridae Newport, 1844 and Siphonorhinidae Cook, 1895. The group has previously been referred to as a “taxonomist’s nightmare” (Read and Enghoff 2009: 543) and got the “taxonomists’ award for the least popular group among Diplopoda” (Jeekel 2001: 44). This is mainly due to the weakly modified gonopods, compared to those of other helminthomorph taxa, in which the gonopods are highly species specific (Jeekel 2001). In the Siphonophoridae the head extends into a long beak and the antennae are straight, while in the Siphonorhinidae the pyriform head lacks such a beak and carries elbowed antennae (Enghoff et al. 2015). Both groups are already known from the Cretaceous of Myanmar ca. 99 million years ago (Wesener and Moritz 2018). The Siphonorhinidae include five genera, the monotypic Kleruchus Attems, 1938 and Nematozonium Verhoeff 1939, Illacme Cook and Loomis, 1928 with two species, Siphonorhinus Pocock, 1894 with eight species (Jeekel 2001; Marek et al. 2016), and Madagascarhinus Wesener, 2023 with two species (Wesener 2023). Detailed catalogues of the group have been provided by Jeekel (2001) and Marek et al. (2016). The family Siphonorhinidae is distributed in South Africa, Madagascar, India, Southeast Asia, and North America (Fig. 1A). Not a single species of the family has been recorded from South America (Jeekel 2001; Enghoff et al. 2015; Marek et al. 2016), where the family Siphonophoridae appears to be quite diverse and widespread (Shelley 1996; Read and Enghoff 2009, 2018, 2019). However, from Chile not a single representative of the order Siphonophorida, neither the families Siphonophoridae nor Siphonorhinidae, has been reported so far (Parra-Gómez 2022).

Figure 1. 

Distribution and habitat of the Siphonorhinidae and Notorhinus floresi sp. nov. A Distribution of Siphonorhinidae in the world. Records from: Pocock (1894); Silvestri (1895); Attems (1930, 1936, 1938); Turk (1947); Shelley and Hoffman (2004); Shelley and Golovatch (2011); Marek et al. (2012, 2016); Wesener (2014, 2023). Where no coordinates were available, approximate coordinates were taken based on the locality description. Map data from NaturalEarth. B Distribution of Notorhinus sp. nov. in Chile. Map data from Olson et al. (2001). C Localities of Notorhinus floresi sp. nov. (e23) and Notorhinus sp. (e27) in the region Biobio. Map data from Maptiler and OpenStreetMap contributors. D Habitat of Notorhinus floresi sp. nov., a native forest surrounded by eucalypt monoculture. Photograph by Edgardo Flores.

With a north-south extension of more than 4,000 km, the narrow country of Chile spans a wide array of climatic conditions and biomes (Veblen et al. 2007) and contains one of the world’s major biodiversity hotspots: The Chilean Winter Rainfall–Valdivian Forest hotspot (ChV) in south-central Chile, that mainly includes the Chilean matorral and Valdivian temperate forests (Arroyo et al. 2004; Mittermeier et al. 2011; Fuentes-Castillo et al. 2020). Chile, and especially this hotspot, harbor a rich biodiversity with many endemic plants and animals (Arroyo et al. 2004), including several endemic millipede species distributed in five orders (Parra-Gómez 2022). However, these areas are already threatened by habitat loss (Arroyo et al. 2004). 65 of the 75 known Diplopoda species from Chile are considered to be endemic and most of these millipedes can be found in the south-central zone (Parra-Gómez 2022), including a relatively high diversity of the colobognathan order Polyzoniida (Mauriès and Silva 1971; Golovatch 2014; Parra-Gómez 2022). This area is characterized by evergreen broadleaved trees, evergreen coniferous vegetation and a humid temperate to Mediterranean climate (Veblen et al. 2007).

Here we describe the genus Notorhinus gen. nov. and the species Notorhinus floresi sp. nov. which is the first record of the order Siphonophorida in Chile and of the family Siphonorhinidae in South America. Furthermore, we record another representative of the genus Notorhinus gen. nov. from Chile.

2. Material and Methods

2.1. Abbreviations

br – body-ring(s); MNHNCMuseo Nacional de Historia Natural de Chile, Casilla 787, Santiago, Chile; SEM – Scanning electron microscopy; ZFMKZoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 127, Bonn, Germany; T – telson (further abbreviations in figure legends).

2.2. Specimen collecting

Specimens were collected by hand during trips throughout 2019 and 2020 to El Natri and near Caramavida in the Biobio region, Chile. Specimens were preserved in 70% ethanol and are stored at the MNHNC and ZFMK.

2.3. Distribution map

Worldwide distribution of the Siphonorhinidae (Fig. 1A) and the records of Siphonorhinidae from Chile (Fig. 1B, C) were mapped in QGIS 3.28.1 based on literature records (Pocock 1894; Silvestri 1895; Attems 1930, 1936, 1938; Turk 1947; Shelley and Hoffman 2004; Shelley and Golovatch 2011; Marek et al. 2012, 2016; Wesener 2014, 2023) and the data presented here. Where no coordinates were given, approximate coordinates were taken based on the locality descriptions. A distributional map of the species from Chile was made based on the ecoregions provided by Olson et al. (2001) and satellite data from Maptiler and OpenStreetMap contributors.

2.4. Examination and photographs

Specimens were examined with a Zeiss Discovery V12 stereo microscope. Photographs were taken at different focus planes and stacked with MicroPublisher 5.0 RTV camera (Q Imaging) mounted to a Leica Z6 imaging system with AUTO-MONTAGE PRO version 5.03.0061 (Synoptics Ltd). The number of body-rings was counted (including the collum, excluding the telson), and the length of the animals were measured from the photographs, and body-width was measured based on SEM images (see below) in IMAGEJ 1.53c (Schneider et al. 2012). Additional images are deposited on Zenodo (

2.5. Scanning electron microscopy (SEM)

For SEM the head, body-rings, telson and gonopods of the male holotype of N. floresi sp. nov. (MNHNC 8387), and the head, body-rings and telson of the single female Notorhinus sp. (MNHNC 8390) were dissected under a Zeiss Discovery V12 stereo microscope. The body parts were transferred via an ascending ethanol series from 70% ethanol to 100% ethanol, air dried and placed with conductive tape on SEM stubs. Due to the small size of the dissected specimens and the mesh size of the available sample container critical point drying was not possible. The samples were sputtered with gold (35 nm) with a Cressington Sputter Coater 108auto and examined with a Zeiss Sigma 300 VP scanning electron microscope. Brightness and contrast of the images were adjusted in GIMP 2.10.20.

3. Results

3.1. Taxonomy

Class Diplopoda de Blainville in Gervais, 1844

Subclass Chilognatha Latreille, 1802/1803

Infraclass Helminthomorpha Pocock, 1887

Subterclass Colobognatha Brandt, 1834

Order Siphonophorida Newport, 1844

Siphonorhinidae Cook, 1895


The specimens (Notorhinus floresi sp. nov. and Notorhinus sp. (MNHNC 8390)) can be placed in the family Siphonorhinidae based on the following characters: Head pyriform, not extended into beak (rostrum) (Figs 3A, 8A); antennae elbowed; antennomere 2 longer than wide (Figs 3B, 8C); gnathochilarium consisting of separate plates (Figs 3G, 8B); anterior margin of collum straight (compare Enghoff et al. 2015; Marek et al. 2016). According to a key provided by Shelley (1996: 23) members of the family Siphonorhinidae lack sensory pits on antennomeres 5 and 6 (see also Marek et al. 2012), as is the case in the specimens studied here (Figs 3E, 8C). However, lack of these sensory pits seems to apply only to the North American genus Illacme (Marek et al. 2012, 2016), the Malagasy genus Madagascarhinus (Wesener 2023) and the South African genus Nematozonium (Shelley and Hoffman 2004), as these sensory pits have been reported for Siphonorhinus (Attems 1930, 1938) and Kleruchus (Attems, 1938).

Notorhinus gen. nov.

Type species

Notorhinus floresi sp. nov.


Pale, thin and elongated Siphonorhinidae with pyriform heads. In Notorhinus gen. nov. (Figs 2, 7) two backwards projecting spines are situated behind the elevated ozopores (Figs 4E, 8G), and the posterior margin of the metazonite (limbus) carries sculptures in shape of a fluke (tail-fin) (Figs 4D, 8H). Notorhinus gen. nov. differs from all other Siphonorhinidae genera by the absence of sensilla basiconica on antennomere 5 and the arrangement of numerous sensilla basiconica on antennomere 6 in a field not sunken into a sensory pit (Figs 3E, 8C). In Siphonorhinus and Kleruchus (see Attems 1930 for S. pellita; Attems 1938 for Teratognathus (syn. of Siphonorhinus) and Kleruchus) numerous sensilla basiconica are located in sensory pits (Sinnesgruben sensu Attems 1930) on antennomeres 5 and 6. In Illacme few sensilla basiconica are arranged in rows along the apical margin of antennomeres 5 and 6 (Marek et al. 2012, 2016) and in Madagascarhinus sensilla basiconica are arranged in 2 – 3 rows on antennomeres 5 and 6 (Wesener 2023). In Nematozonium no sensory structures are evident on the antennae according to Shelley and Hoffman (2004). Notorhinus gen. nov. differs from Illacme by the fusion of the first legs’ coxae to the sternite (Fig. 3G), forming a coxosternite (see Marek et al. 2012, 2016 for Illacme), as is the case in Siphonorhinus (see Enghoff et al. 2015). Furthermore, the specimens differ from Kleruchus by the presence of an anal scale/hypoproct (Figs 4F, 8I) (Analschuppe sensu Attems 1938).


Several characters are shared with the genus Illacme Cook and Loomis, 1928 (see Marek et al. 2012, 2016 for detailed documentation): antennomeres 5 and 6 longer and wider; antennae elbowed between antennomeres 3 and 4 (Fig. 3B); presence of cluster of spiniform sensilla basiconica (spiniform basiconic sensilla sensu Marek et al. 2016) next to apical sensory cones (Fig. 3F); labrum with incision, lined by teeths; pores of salivary glands arranged in a single circular field above the labrum (Fig. 3C); gonopods with 7 podomeres (Fig. 5E); posterior gonopods apical podomere with three branches: two apically laminate/flattened branches and a single posterior spine-like branch (Fig. 5F). However, some of these characters are not well studied in the remaining genera of the Siphonorhinidae.


Noto is derived from the ancient greek νότος (nótos) meaning south and refers to the distribution of the genus in South America and the fact that it is the most southern record of the family Siphonorhinidae. Rhinus is derived from the ancient Greek ῥῑ́ς (rhī́s; genitive: ῥῑνός (rhīnós)), meaning nose, and refers to the acuminate head shape. Rhinus is often part of taxonomic names in the group (e.g. Siphonorhinidae Cook, 1895, Siphonorhinus Pocock, 1894, Madagascarhinus Wese­ner, 2023).

Notorhinus floresi sp. nov.

Figs 2, 3, 4, 5


Small (< 13 mm) elongated Siphonorhinidae with arched metazonites. Body pale, covered by setae, creating a velvety appearance (Fig. 2A). Anterior and posterior gonopods with 7 podomeres (Fig. 5E). Posterior gonopod apically with 3 branches, 2 laminate and 1 spinous (Fig. 5F). Pseudopenis prominent, cone-shaped (Fig. 5B). Posterior margin of metazonite (limbus) with small fluke-shaped sculpture (Fig. 4D). Antennomere 7 with spiniform sensilla basiconica in 2 rows (Fig. 3F) (1 row in Notorhinus sp.). Metazonite posterior margin straight (Fig. 4A) (sinuate in Notorhinus sp.). Differs from Notorhinus sp. (MNHNC 8390) in coloration, number of body-rings and its length relative to the number of body-rings (Fig. 2A): Notorhinus sp. has 80 body-rings but is only 11.5 mm long, while Notorhinus floresi sp. nov. has up to 54 body-rings, but a length of up to 13.8 mm.

Figure 2. 

Notorhinus floresi sp. nov., photographs. A, B Living specimen in its natural habitat. Photographs by Edgardo Flores. C Male paratype (ZFMK-MYR 12267), habitus, lateral view. D Female paratype (ZFMK-MYR 12269), head, lateral view.

Figure 3. 

Notorhinus floresi sp. nov., male holotype (MNHNC 8387), head, SEM. A Head, frontal view. B Head, lateral view. C Labrum, frontal view. D Labrum and gnathochilarium, ventral view. E Apical antennomeres, lateral view. F Apical antennomere, apical view. G Gnathochilarium and leg-pair 1, ventral view. Abbreviations: I–VII = antennomeres, ac = apical cones, cl = claw, co = collum, cx = coxa, ep = epicranium, fe = femur, fh = forehead, ge = genae, in = inscision of the labrum, la = labrum, LL = lamella lingualis, lp = spinous protuberences of labrum, lt = teeth lining labral incision, me = mentum, pa = palp-like structure, po = pores of the salivary glands, pof = postfemur, prf = prefemur, sb = sensilla basiconica, ssb = spiniform sensilla basiconica, sti = gnathochilarial stipes, ta = tarsus, ti = tibia.

Figure 4. 

Notorhinus floresi sp. nov., male holotype (MNHNC 8387), body-rings and legs, SEM. A Mid-body-rings, dorsal view. B Mid-body-rings, lateral view. C Mid-body-rings, ventral view, legs removed. D Metazonite posterior margin (limbus), arrow indicates fluke-shaped sculpture. E Ozopore, lateral view, arrows indicate spines behind ozopore. F Telson, posterior view. G Mid-body leg, anterior view. H Leg 2, tarsal claw, arrow indicates division of tarsal claw. Abbreviations: cl = claw, cs = coxal sac, cx = coxa, fe = femur, hp = hypoproct (subanal scale), mz = metazonite, oz = ozopore, pl = pleurite, pof = postfemur, pp = paraproct (anal valve), pr = preanal ring, prf = prefemur, pz = prozonite, sp = spiracle, st = sternite, ta = tarsus, ti = tibia, tr = trochanter.

Figure 5. 

Notorhinus floresi sp. nov., male holotype (MNHNC 8387), sexual characters, SEM. A Leg 2, ventral view. B Male pseudopenis bearing the gonopore, ventral view. C Male pseudopenis bearing the gonopore, posterior view. D Anterior and posterior gonopods, ventral view. E Anterior and posterior gonopods, ventro-lateral view. F Right anterior and posterior gonopod, detail of the apical podomeres. Abbreviations: a1–a7 = podomeres of the anterior gonopod, cx = coxa, f1 = pointed/spinous branch of the posterior gonopod’s apical podomere, f2 and f3 = laminate branches of the posterior gonopod’s apical podomere, go = pseudopenis carrying the gonopore, p1–p7 = podomeres of the posterior gonopod, st = sternite associated with anterior gonopods.


The species epithet floresi refers to Edgardo Flores, who collected the examined specimens, and honors his continuous engagement in nature conservation and his persistence on the protection of Nahuelbuta National Park and adjacent areas. Noun in genitive.

Material examined

Holotype: ♂ (MNHNC 8387); CHILE: Biobío, near Caramavida; 37.72698°S 73.19339°W; 21.iix.2019; Edgardo Flores leg. (e23); hand collected under a decaying piece of wood, in a relictual forest surrounded by eucalypt plantations (Fig. 1C, D). — Paratypes: 2 ♂ (ZFMK-MYR 12267, ZFMK-MYR 12268); 3 ♀ (MNHNC 8388, MNHNC 8389, ZFMK-MYR 12269); same data as holotype.


Measurements : Male holotype (MNHNC 8387): br47+T, 12 mm long; male paratype (ZFMK-MYR 12267): br37+T, 8.9 mm; male paratype (ZFMK-MYR 12268): br54+T, 13.8 mm; female paratype (MNHNC 8389: br42+T, 10.7 mm; female paratype (MNHNC 8388): br49+T, 12.8 mm; female paratype (ZFMK-MYR 12269): br47+T, 10.9 mm. Body thread-like, elongate thin, pale white (Fig. 2A–C), in ethanol head lighter in coloration than body (Fig. 2C, D). — Head: Head pyriform, anteriorly tapering, widest behind antennae, covered by long setae (Fig. 3A), posterior and lateral of antennae covered by scale-like sculpture (Fig. 3B). Labrum incised, two lateral spinous protuberances on each side of incision, incision lined by. by up to 5 teeth on each side. Salivary glands opening via > 50 small pores, arranged in a circular field above labrum (Fig. 3C, D). Lateral margin (genae) and anterior margin (labrum) of head capsule straight, tightly appressed to gnathochilarium (Fig. 3B). Antennae inserting laterally in posterior half of head capsule (Fig. 3A), with 7 antennomeres + apical disc, elbowed between antennomeres 3 and 4. Antennomere 6 longest (6 > 2 > 5 > 1 = 3 = 4 > 7), 2 and 6 longer than wide, 1, 3–5 and 7 wider than long, 5 and 6 widest. Each antennomere with several rows of setae (Fig. 3B). Antennomere 5 without sensilla basiconica. Antennomere 6 with ca. 25 sensilla basiconica arranged in a circular field (not sunken into a sensory pit) located laterally in some distance from antennomere’s apical margin (Fig. 3E). Antennomere 7 on apical margin with a lateral group of 8 spiniform sensilla basiconica arranged in two rows, and a single spiniform sensilla basiconica each at anterior and posterior margin, projecting above apical disc between apical cones. Apical disc sunken into antennomere 7 with 4 apical sensory cones, base of apical cones elevated (Fig. 3F). Gnathochilarium triangular, tightly appressed to head capsule consisting of stipites, mentum and lamellae linguales (proximal part not visible). Stipites each with 5 marginal setae, apically with palp-like extension, each with 5 sensilla (Fig. 3D). Mentum with 6 (3+3) setae arranged in 2 longitudinal rows. Lamellae linguales each with 2 short setae, mesal margins apically with hair-like structures (Fig. 3D, G). Mandibles internalized, not visible externally (Fig. 3A, B). — Body-rings: Collum longer than following tergite, covering posterior part of head, anterior margin almost straight. Collum covered by long setae and laterally with scale-like sculpturing (Fig. 3A, B). Each body-ring consisting of free tergite (1), pleurites (2) and sternites (2) (Fig. 4A–C). Body-rings 2–4 shorter than following body-rings. Tergites clearly divided into prozonite and metazonite (Fig. 4A, B). Prozonite narrower than metazonite, covered by circular cylinder/pillar-shaped sculpturing, lateral sculpturing becoming more acuminate, anterior margin with scale-like sculpturing, with few setae along anterior and posterior margin (Fig. 4A, B). Metazonite arched, circular/cylindrical sculpture restricted to anterior and lateral areas, resulting in a crown-shaped smooth dorsal area. Metazonite covered by setae (Fig. 4A, B). Posterior margin of metazonite (limbus) with 2 irregular rows of fluke-shaped sculptures (Fig. 4D). Ozopores from body-ring 5 onwards, positioned laterally in posterior half of metazonite, slightly elevated, surrounded by a ring of setae, with 2 posterior spines and 3 setae between spines (Fig. 4E). Pleurites almost rectangular, anteriorly and posteriorly rounded, laterally slightly curved towards legs. Pleurites anteriorly covered by setae and circular/cylindrical sculptures, posterior lateral sculpturing acuminate, anterior part of pleurite covered by flat scale-like sculptures (Fig. 4C). Sternites rectangular with a transverse elevation projecting between the legs’ coxae. Margins of this elevation with scale-like sculpturing, spiracles located laterally of legs (Fig. 4C). — Telson: Preanal ring ca. as long as preceding body-ring, posteriorly rounded in dorsal view, covered by setae. Cylindrical sculpture restricted to lateral and ventral part and the anterior margin, dorsal surface without sculpture (Fig. 4F). Anal valves (paraprocts) covered by setae, proximally with scale-like sculpturing. Subanal scale (hypoproct) present, with few setae and flat scale-like sculpture (Fig. 4F). — Legs: Leg pairs 1 and 2 with 6 podomeres: coxa, prefemur, femur, postfemur, tibia and tarsus. Coxae of leg-pair 1 fused to sternite (coxosternite) (Fig. 3G). From leg-pair 3 onwards with coxal sacs and short additional podomere (trochanter) between coxa and prefemur: Prefemur = tarsus > femur = coxa > postfemur = tibia > trochanter (Fig. 4G). Coxae almost touching mesally. Podomeres with few long setae (Fig. 4G). Tarsal claw bifurcated, ventral branch ca. 2/3 of length of dorsal branch on anterior legs (Fig. 4H), ventral branch decreasing in size on posterior legs. Last body-ring anterior of telson apodous. — Male sexual characters: Male gonopores on pseudopenes situated posteriorly on coxae of leg-pair 2 (Fig. 5A), pseudopenis knob-like/cone-shaped in ventral view (Fig. 5B), latero-posterior surface excavated with membranous surfaces (Fig. 5C). Leg-pair 9 and 10 modified to leg-like gonopods (Fig. 5D). Anterior gonopods with 7 podomeres, tapering distally. Each podomere with few setae arranged in a row. Podomere 1 with 2 anterior setae, podomere 2 with 1 anterior seta, podomere 3 with 3 anterior setae, podomere 4 with 1 anterior and 1 posterior setae, podomere 5 with 1 lateral and 4 posterior setae, podomere 6 with 2 lateral and 3 posterior setae, podomere 7 with 4 posterior setae (Fig. 5D, E). Apical podomere (7) flap-like, forming a sheath engulfing the tip of the posterior gonopod (Fig. 5F). Posterior gonopod with 7 podomeres, tapering distally. Podomeres 1–4 without setae, podomere 5 with 1 lateral seta, podomere 6 with 2 lateral setae, podomere 7 proximally with 1 posterior seta (Fig. 5D, E). Apical podomere (7) elongated, forming 3 branches, 1 short and pointed (spinous) posterior branch, and 2 long anterior branches, both proximally thin cylindrical, distally flattened (laminate) and paddle-shaped, almost triangular with sinuate margin. The 2 long branches rest in the sheath formed by anterior gonopod (Fig. 5F). — Female sexual characters: Female gonopores inconspicuous, behind 2nd coxa.


A fungus in amphoromorph state was located on the 4th antennomere (left) of the male holotype (Fig. 6).

Figure 6. 

Fungus in amphoromorph stage on antennomere 4 of the male holotype (MNHNC 8387) of Notorhinus floresi sp. nov.

Notorhinus sp.

Figs 7, 8

Material examined

1♀ (MNHNC 8390); CHILE: Biobío, El Natri, Lanalhue, Contulmo; 37.9°S 73.27°W; 23.iv.2020; Edgardo Flores leg. (e27).


Body elongate and thread-like, pale white (Fig. 7), 11.5 mm long, 0.37 mm wide, br80+T. Head pyriform (Fig. 8A) with incised labrum (Fig. 8B) and antennae elbowed between antennomere 4 and 5 (Fig. 8C). Sensilla basiconica on antennomere 5 absent, sensilla basiconica on antennomere 6 arranged in oval field (Fig. 8C), 6 spiniform sensilla basiconica on antennomere 7 arranged in single row (Fig. 8D) (2 rows in N. floresi sp. nov.). Lateral metazonal sculpturing present (Fig. 8F). Two backwards projecting spines behind the ozopores and 3 setae between spines (Fig. 8G) as present in N. floresi sp. nov. Fluke-shaped projections on the posterior margin of metazonites (limbus) (Fig. 8H) as in N. floresi sp. nov. Because only a single female specimen of Notorhinus sp. (MNHNC 8390) is known, we refrain from describing it as a new species until male specimens are available.

Figure 7. 

Notorhinus sp., female (MNHNC 8390). A Habitus, lateral view. B Head and anterior body, lateral view.

Figure 8. 

Notorhinus sp., female (MNHNC 8390), SEM. A Head and anterior body-rings, frontal view. B Apical parts of gnathochilarium and labrum, ventral view. C Left antenna, lateral view. D Antennomere 7 with apical sensory structures. E Tarsal claw of mid-body leg. F Mid-body tergite, dorsal view. G Ozopore, lateral view, arrows indicate spines behind ozopore. H Posterior tergal margin (limbus), dorsal view, arrow indicates fluke-shaped sculpture. I Posterior body-rings and telson, ventral view. Abbreviations: I–VII = antennomeres, ac = apical cone, cl = tarsal claw, co = collum, ep = epicranium, fh = forehead, gn = gnathochilarium, hp = hypoproct, L1 = leg-pair 1, L2 = leg-pair 2, la = labrum, LL = lamella lingualis, md = mandible gnathal lobe, me = mentum, mz = metazonite, oz = ozopore, pp = paraproct, pr = preanal ring, pz = prozonite, sb = sensilla basiconica, ssb = spiniform sensilla basiconica, sti = gnathochilarial stipes, ta = tarsus.

4. Discussion

4.1. Affinities of Notorhinus gen. nov. to Illacme Cook and Loomis, 1928

Notorhinus gen. nov. is distinct from the remaining Siphonorhinidae genera in its morphology as pointed out in the genus diagnosis. However, it shows morphological similarities to the North American genus Illacme Cook and Loomis, 1928 or at least one of its members (­Table 1). Notorhinus gen. nov. shares with Illacme the structure of the gonopods with 7 podomeres in both pairs, the absence of sensory pits on the antennae and the presence of a lateral group of spiniform sensilla basiconica on antennomere 7, reaching between the apical cones. Furthermore, the single isolated spiniform sensilla basiconica on the anterior and posterior margin of antennomere 7 might also be present in Illacme plenipes Cook and Loomis, 1928 (compare Marek et al. 2012, fig. 13).

Table 1.

Comparison of morphological characters in Notorhinus gen. nov. and Illacme Cook and Loomis, 1928 based on Marek et al. (2016: table 1). n indicates the number of specimens measured. Asterisk (*) indicates characters not included in Marek et al. (2016: table 1).

Notorhinus floresi sp. nov. Notorhinus sp. (e27) Illacme plenipes Cook and Loomis, 1928 Illacme tobini Marek, Shear and Krejca, 2016
Number of body-rings* (T = telson) 37–54 + T [n = 6] 80 + T [n = 1] 84–192 + T(Marek et al. 2012: table 1) [n = 8] 108 + T (Marek et al. 2016: table 2) [n = 1]
Length* 8.9–13.8 mm [n = 6] 11.5 mm [n = 1] 13.4–40.4 mm (Marek et al. 2016: table 3) [n = 8] 19.7 mm (Marek et al. 2016: table 2) [n = 1]
Antennae, Antennomere 5 sensilla basiconica* Absent Absent Present, row along apical margin (Marek et al. 2012: fig. 14; Marek et al. 2016: fig. 3B) Present, row along apical margin (Marek et al. 2016: fig. 3A)
Antennae, Antennomere 6 sensilla basiconica* Present, in field in some distance from apical margin (Fig. 3E) Present, in two rows close to apical margin (Fig. 8C) Present, row along apical margin (Marek et al. 2012: fig. 14; Marek et al. 2016: fig. 3B) Present, row along apical margin (Marek et al. 2016: fig. 3A)
Antennae, Antennomere 7, spiniform sensilla* In cluster of 8 (Fig. 3F) In row of 6 (Fig. 8D) In cluster of 5 (Marek et al. 2012: fig. 13) In cluster of 4 (Marek et al. 2016: 18)
Metazonite/prozonite Metazonite wider than prozonite (Fig. 4A) Almost equal in width (Fig. 8F) Almost equal in width (Marek et al. 2016: fig. 10B) Metazonite wider than prozonite (Marek et al. 2016: fig. 10A)
Metazonite sculpture (except margin)* Extending laterally and dorsally (Fig. 4A, B) Restricted to lateral potion (Fig. 8F) Absent (Marek et al. 2016: fig. 10B) Absent (Marek et al. 2016: fig. 10A)
Spines behind ozopore (on paraproct) 2 short backwards projecting spines (Fig. 4E) 2 short backwards projecting spines (Fig. 8G) 2 large backwards projecting spines (Marek et al. 2016: fig. 10D) Absent (Marek et al. 2016: fig. 10C)
Sculpture on metazonite posterior margin (limbus) Fluke-shaped spines (Fig. 4D) Fluke-shaped spines (Fig. 8H) Anchor-shaped spines (Marek et al. 2016: fig. 10F) Quadrate spines (Marek et al. 2016: fig. 10E)
Metazonite posterior margin shape Straight (Fig. 4A) Sinuate (Fig. 8F) Straight (Marek et al. 2016: fig. 10B) Sinuate (Marek et al. 2016: fig. 10A)
Telson sculpture/spines Lateral and ventral surfaces (Fig. 4F) Lateral surface only (Fig. 8I) All surfaces (Marek et al. 2016: fig. 11B) Lateral surface only (Marek et al. 2016: fig. 11A)
Hypoproct setation > 2 setae (Fig. 4F) > 2 setae (Fig. 8I) > 2 setae (Marek et al. 2016: fig. 11B) 2 setae (Marek et al. 2016: fig. 11A)
Tarsal claw bifurcation, length of smaller claw* Long (2/3 of length of claw) (Fig. 4H) Long (2/3 of length of claw) (Fig. 8E) Long (2/3 of length of claw) (Marek et al. 2012: fig. 27) Long (2/3 of length of claw) (Marek et al. 2016: fig. 17C)
Anterior gonopod, podomere 3, setation 3 setae (Fig. 5E) ? 6 setae (Marek et al. 2016: fig. 9B, D, F) 2 setae (Marek et al. 2016: fig. 9A, C, E)
Anterior gonopod apex, setae/spines 5 setae (Fig. 5F) ? 3 setae (Marek et al. 2016: fig. 9D, F) 9 setae (Marek et al. 2016: fig. 9C, E)
Posterior gonopod apex, branches (articles) Bundle of 3 styliform branches (Fig. 5F) ? Bundle of 3 styliform branches (Marek et al. 2016: fig. 11D) Bundle of 4 styliform branches (Marek et al. 2016: fig. 11C)
Posterior gonopod apex, margin of apically flattened branches* Sinuate (Fig. 5F) ? Serrated (Marek et al. 2012: fig. 12) Serrated (Marek et al. 2016: fig. 11C)

Notorhinus gen. nov. shares with Illacme plenipes the two backwards projecting spines behind the ozopores with three setae in between, but the spines are shorter and stouter than in I. plenipes. Notorhinus gen. nov. also shares with I. plenipes the projections on the posterior margin of the metazonites (limbus), but in Notorhinus gen. nov. these projections are rather fluke-shaped and not as well developed as in I. plenipes, in which these have been described as anchor-shaped (Marek et al. 2012, 2016). Moreover, Notorhinus floresi sp. nov. shares with I. plenipes the presence of 3 branches (articles) on the posterior gonopod’s apical podomeres, two laminate flattened ones and one spinous. N. floresi sp. nov. also shares with I. plenipes the straight posterior margin of the metazonite, while the margin is sinuate in Illacme tobini Marek, Shear and Krejca, 2016 and Notorhinus sp. (MNHNC 8390). Therefore, we suggest that the South American Notorhinus gen. nov. is more closely related to the North American Illacme, than to the Asian and African genera. However, most of these characters are not well studied in Siphonorhinidae, except for Illacme thanks to Marek et al. (2012, 2016).

4.2. Biogeography and habitat preferences of Siphonorhinidae in the Americas

Notorhinus floresi sp. nov. and the undescribed Notorhinus species (MNHNC 8390) occur in Chile at ca. 38° South, while Illacme plenipes and Illacme tobini occur in California (USA) at ca. 37° North (Marek et al. 2012, 2016). All three species occur close to the Pacific coast mostly in wet subterranean/poorly lighted microhabitats within temperate to Mediterranean humid native woodlands (Veblen et al. 2007; Marek et al. 2012). Thus, the known distribution of Siphonorhinidae in America is antitropical (amphitropical), meaning that the taxon shows a disjunct distribution with representatives occurring at similar latitudes in the northern and southern hemisphere, but not in the tropical region in between (Hubbs 1952). Examples of antitropical distributions include many marine taxa, like fishes (Randall 1981), whales (Barnes 1985), echinoderms (Naughton et al. 2014), mollusks (Koufopanou 1999; Hilbish et al. 2000), and terrestrial plants from North and South America (Raven 1963; Simpson et al. 2017), but also terrestrial insects like some wingless darkling beetles from Africa and Europe (Kamiński et al. 2021) and bees from North and South America (Wilson et al. 2014; Freitas et al. 2022). Observing such a distribution in millipedes is remarkable as these soil-organisms show a very limited dispersal ability and are often adapted to certain micro-habitats (Sierwald and Bond 2007; Golovatch and Kime 2009). The exchange between the North and South American flora and fauna, referred to as Great American Biotic Interchange (GABI), is often associated with the rise of the Isthmus of Panama in the late Pliocene about 3 million years ago (Leigh et al. 2014). However, for some taxa it has been shown that this exchange probably already started earlier in the Miocene (Cody 2010; Pinto-Sánchez et al. 2011; Wilson et al. 2014). The antitropical distribution of the American Siphonorhinidae might be explained by a once wider distribution in North and South America followed by extinction events in the tropics due to changes in climate and environmental conditions. However, the observed pattern might also be due to incomplete sampling, as these enigmatic animals, typically restricted to small soil habitats, are often overlooked, with I. plenipes collected for the first time since its description (Cook and Loomis 1928) in 2006 (Marek and Bond 2006), a second Illacme species only described recently (Marek et al. 2016), and the specimens described here only discovered in 2019 and 2020. Due to their restriction to very small habitats, Shelley (1996: 22) concluded that to find Siphonophorida “one must not only investigate the right habitat but also fortuitously dislodge the one or few rocks or logs that shelter them”. Thus, “the probability of discovering siphonophorids is low and apparently chiefly a matter of chance” (Shelley 1996: 22). Furthermore, the group received only little attention by taxonomists (Jeekel 2001). Antitropical distributions have been assumed for other taxa, which had to be corrected later, when species were discovered in the tropics (e.g. Harris and Armitage 1997; Holzenthal and Harris 2002). Currently, we exclude an anthropogenic introduction of the species to Chile, as there is no species of the order known to have been expanded through human activities. This is further supported by the rarity of these millipedes across the globe (Marek et al. 2012), and the habitat in a small relict of native forest in which they were found.

Specimens of N. floresi sp. nov. were encountered in a small, fragmented patch of native forest between eucalypt plantations near Camaravida, which lays within the Chilean Winter Rainfall–Valdivian Forest hotspot. The temperature and precipitation in this biodiversity hotspot is affected by anthropogenic climate change, and especially smaller protected areas appear to be vulnerable in this region (Fuentes-Castillo et al. 2020). The siphonorhinid millipedes might have evolved in mild consistent habitats and climate for hundred millions of years and thus they are dependent on certain microhabitats and climatic conditions (Marek et al. 2012). Therefore, climate change and habitat loss pose a serious risk to them, thus protection and preservation of these habitats is essential.

4.3. The Diplopoda fauna of Chile

The Chilean Diplopoda fauna shows predominantly Gondwanan affinities, with closely related taxa occurring in South Africa, Australia, and other Gondwanan landmasses, rather than in other South American regions (Shelley and Golovatch 2011; Golovatch 2014). With the addition of Siphonophorida a total of six native orders can be found in Chile (Chordeumatida, Polydesmida, Polyxenida, Polyzoniida, Siphonophorida and Spirostreptida) (Parra-Gómez 2022). This is the first record of a new millipede order for the country in 65 years, the last addition was the order Polyzoniida in 1957 (Chamberlin 1957). Furthermore, the order Julida has been introduced to Chile, with 5 species native to Europe, and can even be found in natural habitats inside national parks (Golovatch 2014; Parra-Gómez 2022). The native millipede species usually have a narrow distribution, and are often endemic, with the highest diversity in the temperate regions at mid-latitudes (Parra-Gómez and Fernández 2022). However, the knowledge on their distribution remains obscure, with several latitudinal gaps without any records and fragmentary distributions for various taxa (Parra-Gómez 2022).

5. Conclusion

The discovery and description of Notorhinus gen. nov. is a significant contribution to our knowledge of the Colobognatha, as this is the first record of the order Siphonophorida in Chile and of the family Siphonorhinidae in South America. Notorhinus gen. nov. represents the most southern occurrence of the family Siphonorhinidae. While Notorhinus gen. nov. is morphologically distinct from the remaining Siphonorhinidae genera it shares several somatic and sexual characters with the North American genus Illacme Cook and Loomis, 1928.

6. Acknowledgements

We would like to express our deepest gratitude to Edgardo Flores for his collection efforts and for providing the specimens studied here, as well as photographs of the specimens and their habitat. Furthermore, we would like to thank Pooja Avinipully Anilkumar (ZFMK) for taking some additional SEM images of Notorhinus floresi sp. nov., and Jorge Pérez Schultheiss (MNHN, Chile) for logistic support. LM thanks Thomas Wesener (ZFMK) for discussions and encouragement. We thank Paul Marek for discussions. We are grateful to Andy Sombke and Klaus-Dieter Klass, and the two reviewers Henrik Enghoff and Zoltan Korsós for valuable comments on the manuscript.

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