Giant Millipede Research Articles

A new distinct, disjunct giant millipede of the genus Spirostreptus Brandt, 1833, from Tanzania, and a solution for orphaned Spirostreptus species (Diplopoda, Spirostreptida, Spirostreptidae).

Spirostreptus digitus sp. nov. is described based on specimens from Tanzania, and the genus Spirostreptus Brandt, 1833 is discussed, as is the higher classification of the family Spirostreptidae. Gonopod illustrations of S. heros Porat, 1872, and S. tripartitus Cook & Collins, 1893, are provided for comparison with the new, morphologically very distinct species. Analocostreptus Silvestri, 1910, is suggested as the valid genus name for the species currently being placed in Spirostreptus sensu auctorum, nec Brandt, 1833, and 32 nominal species of Spirostreptus are placed in new combinations with Analocostreptus, viz., Analocostreptus amandus (Attems, 1914), comb. nov., Analocostreptus biconus (Attems, 1934), comb. nov., Analocostreptus bonifatius (Attems, 1914), comb. nov., Analocostreptus castaneus (Attems, 1934), comb. nov., Analocostreptus cornutus (Attems, 1934), comb. nov., Analocostreptus damasus (Attems, 1953), comb. nov., Analocostreptus dartevellei (Attems, 1953), comb. nov., Analocostreptus dentiger (Attems, 1953), comb. nov., Analocostreptus garambanus (Chamberlin, 1927), comb. nov., Analocostreptus gregorius (Attems, 1914), comb. nov., Analocostreptus ineptus (Kraus, 1958), comb. nov., Analocostreptus informis (Attems, 1938), comb. nov., Analocostreptus makarius (Attems, 1914), comb. nov., Analocostreptus manyemanus (Attems, 1927), comb. nov., Analocostreptus manyemanus biserialis (Attems, 1938), comb. nov., Analocostreptus medjensis (Chamberlin, 1927), comb. nov., Analocostreptus micromelas (Saussure & Zehntner, 1902), comb. nov., Analocostreptus missionarius (Attems, 1953), comb. nov., Analocostreptus montivagus (Karsch, 1881), comb. nov., Analocostreptus multisulcatus (Demange, 1957), comb. nov., Analocostreptus pancratius (Attems, 1914), comb. nov., Analocostreptus pavani (Demange, 1981), comb. nov., Analocostreptus phthisicus (Saussure & Zehntner, 1902), comb. nov., Analocostreptus pictus (Saussure & Zehntner, 1902), comb. nov., Analocostreptus rolini (Silvestri, 1897), comb. nov., Analocostreptus sculptus (Saussure & Zehntner, 1902), comb. nov., Analocostreptus semilunaris (Peters, 1855), comb. nov., Analocostreptus servatius (Attems, 1914), comb. nov., Analocostreptus tetricus (Attems, 1934), comb. nov., Analocostreptus tiburtius (Attems, 1953), comb. nov., Analocostreptus triangulicollis (Attems, 1934), comb. nov., Analocostreptus versicolor (Saussure & Zehntner, 1902), comb. nov. and Analocostreptus yambatanus (Attems, 1934), comb. nov. Illustrations of the gonopods of A. ibanda (Silvestri, 1907), type-species of Analocostreptus, are provided, and the species is recorded as new for Kenya.

Histologic lesions of giant African millipedes (Archispirostreptus gigas) from a zoological institution.

Histopathologic data of millipedes are scarce. Little is known about health and disease of these invertebrates despite their exhibition at zoological institutions and use in ecotoxicological studies. In a retrospective study of 69 zoo-housed giant African millipedes (Archispirostreptus gigas) submitted between 2018 and 2021, most deaths occurred during midwinter and in 2021. The most common lesion was inflammation (n = 55; 80%). Necrosis was seen concurrently in 31 (45%) millipedes and of these, bacteria (20; 29%) and fungi (7; 10%) were detected in lesions. Inflammation was seen in the head/collum (20; 29%), hemocoel (16; 23%), and appendages (9; 13%), specifically in perivisceral fat body (42; 61%), gut (16; 23%), tracheae (26; 38%), skeletal muscle (24; 35%), and ventral nerve (17; 25%). Inflammatory cell types and patterns included agranular hemocytes (61; 88%), granular hemocytes (39; 57%), and nodulation/encapsulation (47; 68%) often accompanied by melanization. The oral cavity or gut (ingestion), spiracles (inhalation), or cuticular defects were considered plausible routes of bacterial entry. Metazoan parasites (adult nematodes: 2, 3%; trematode ova: 2, 3%; and arthropods: 1, 1%) were associated with gut necrosis and inflammation in 5 millipedes. In addition, adult nematodes were noted in the gut of 4 millipedes without lesions. Neoplasia was not detected in any millipedes. Speculatively, environmental factors may have predisposed to disease, as most deaths occurred during winter months. Disease surveillance of millipedes is critical to optimize husbandry practices in zoo populations and investigate potential impacts of environmental degradation and climate change on wild millipedes.

Some Aspects of the Physiology of the Nyctotherus velox, a Commensal Ciliated Protozoon Taken from the Hindgut of the Tropical Millipede Archispirostreptus gigas.

In this paper, the growth requirements, fermentation pattern, and hydrolytic enzymatic activities of anaerobic ciliates collected from the hindgut of the African tropical millipede Archispirostreptus gigas are described. Single-cell molecular analysis showed that ciliates from the millipede hindgut could be assigned to the Nyctotherus velox and a new species named N. archispirostreptae n. sp. The ciliate N. velox can grow in vitro with unspecified prokaryotic populations and various plant polysaccharides (rice starch-RS, xylan, crystalline cellulose20-CC, carboxymethylcellulose-CMC, and inulin) or without polysaccharides (NoPOS) in complex reduced medium with soluble supplements (peptone, glucose, and vitamins). Specific catalytic activity (nkat/g of protein) of α amylase of 300, xylanase of 290, carboxymethylcellulase of 190, and inulinase of 170 was present in the crude protein extract of N. velox. The highest in vitro dry matter digestibility was observed in RS and inulin after 96 h of fermentation. The highest methane concentration was observed in xylan and inulin substrates. The highest short-chain fatty acid concentration was observed in RS, inulin, and xylan. In contrast, the highest ammonia concentration was observed in NoPOS, CMC, and CC. The results indicate that starch is the preferred substrate of the N. velox. Hydrolytic enzyme activities of N. velox showed that the ciliates contribute to the fermentation of plant polysaccharides in the gut of millipedes.

Plant Growth-promoting Ability and Pathogen Inhibitory Effect of Actinomycetes Isolated from Fecal Pellets of the Giant Millipede Thyropygus resimus (Diplopoda)

The microbial properties of millipede fecal pellets have been studied mainly in Glomerida (pill millipedes), and much less in the significant majority of other millipede groups. Therefore, the present study examined actinomycetes isolated from the fecal pellets of the non-glomerid giant millipede Thyropygus resimus Attems, 1938 (Spirostreptida) to (1) test their plant growth-promoting ability, and (2) evaluate their potential to control and inhibit plant pathogenic microorganisms. Millipedes were collected from Phu Kum Khao, Kalasin Province, Thailand. A total of 59 actinomycete isolates were obtained and identified as belonging to the genus Streptomyces using 16S rRNA sequencing. The plant growth-promoting properties of the isolates were tested by screening four characteristics: nitrogen fixation, phosphate solubility, siderophore production, and indole-3-acetic acid (IAA) production. A nitrogen-fixation test on nitrogen-free solid malate media (NFM) showed that 54 isolates were capable of fixing nitrogen. Phosphate solubility was tested on double-layered glucose yeast extract agar (GYA) medium containing tricalcium phosphate. This showed that 42 isolates formed a clear zone around the colonies due to phosphate dissolution. Siderophore production was tested on chrome azurol sulfate (CAS) agar. This showed that 55 isolates could grow on this medium and form clear yellow to orange zones around their colonies. IAA production tests revealed that 41 isolates could produce IAA. Based on the combined results of these four tests, eight of the 59 isolates were the most effective in promoting plant growth: KLS-AC04, KLD-AC01, KLD-AC02-1, KLD-AC08, KLD-AC09, KLD-AC16, KLD-AC29-1, and KLD-AC30. Seventeen isolates inhibited the growth of Xanthomonas oryzae pv. oryzae, the causal agent of bacterial leaf blight disease in rice, more effectively than rifampicin (100 ppm), with isolate KLS-AC02-1-1 being the most effective (inhibition zone, 58.25 mm in diameter). Therefore, these isolates can be used for growth promotion and rice disease control in the future.

Geographic distribution, diversity and conservation status of giant millipedes in southern Cameroon rainforest

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A strikingly coloured new giant millipede from Vietnam has copycat in Borneo (Diplopoda, Spirostreptida, Harpagophoridae)

A new giant species of the subfamily Rhynchoproctinae with strikingly bi-coloured, red-yellow legs contrasting with a black body is described: Alienostreptus bicoloripes sp. nov. from Vietnam. The new species is assigned to the genus Alienostreptus Pimvichai, Enghoff & Panha, 2010 hitherto comprising one species, A. alienus (Attems, 1936), and differing from other rhynchoproctine genera by having the femoral spine duplicated. Species of this genus share three synapomorphic characters of the subfamily, viz (1) anterior coxal fold forms deep concavity in posterior view, (2) posterior coxal fold very low, and (3) posterior coxal fold with mesal flap. A superficially very similar colourful species from Borneo, also with bi-coloured legs but clearly not belonging to Alienostreptus due to the position of ventral soft pads on male legs, is documented based on photographs.

Autophagy and Apoptosis in the Midgut Epithelium of Millipedes.

The process of autophagy has been detected in the midgut epithelium of four millipede species: Julus scandinavius, Polyxenus lagurus, Archispirostreptus gigas, and Telodeinopus aoutii. It has been examined using transmission electron microscopy (TEM), which enabled differentiation of cells in the midgut epithelium, and some histochemical methods (light microscope and fluorescence microscope). While autophagy appeared in the cytoplasm of digestive, secretory, and regenerative cells in J. scandinavius and A. gigas, in the two other species, T. aoutii and P. lagurus, it was only detected in the digestive cells. Both types of macroautophagy, the selective and nonselective processes, are described using TEM. Phagophore formation appeared as the first step of autophagy. After its blind ends fusion, the autophagosomes were formed. The autophagosomes fused with lysosomes and were transformed into autolysosomes. As the final step of autophagy, the residual bodies were detected. Autophagic structures can be removed from the midgut epithelium via, e.g., atypical exocytosis. Additionally, in P. lagurus and J. scandinavius, it was observed as the neutralization of pathogens such as Rickettsia-like microorganisms. Autophagy and apoptosis ca be analyzed using TEM, while specific histochemical methods may confirm it.

A giant African millipede re-discovered, re-located and re-classified after120 years (Diplopoda, Spirobolida, Pachybolidae).

The giant millipede Pachybolus tectus Cook, 1897, described from Zanzibar, East Africa, and never re-collected till now, is re-described based on newly collected specimens from Cameroon, West Africa. The species is transferred to the genus Pelmatojulus Saussure, 1860, and compared with the most similar congener, P. togoensis (Cook, 1897). The type specimen of P. tectus, which no longer exists, probably was mis-labelled. An overview of the distribution of Pelmatojulus species and records of the genus from Cameroon are given, including P. brachysternus (Cook, 1897) as new to the fauna of Cameroon. Pelmatojulus insignis (Saussure, 1859) and P. togoensis are recorded as new to the fauna of Ghana.

Mating trackways of a fossil giant millipede

Three closely associated arthropleurid trackways, Diplichnites cuithensis , from the Lower Carboniferous of Fife, Scotland, exhibit signs of interaction between track-makers. An extra file of footprints is found on the downslope side of two trackways (A and C), the upslope side of another (B). These additional files of footprints suggest that either: each trackway resulted from two arthropleurids of different sizes walking in tandem, matching their footprints exactly on one side for some distance; or that one arthropleurid was partially mounted on the back of another producing the three parallel files. It is here argued that the latter is correct and that this represents evidence of mating behaviour in arthropleurids.

The ultrastructure of the hepatic cells in millipedes (Myriapoda, Diplopoda)

The hepatic cells are characteristic cells that occur in millipede body where they form a continuous layer which surrounds the midgut epithelium. The following species, which represent five millipede orders were selected: Julus scandinavius (Julida), Polyxenus lagurus (Polyxenida), Polydesmus angustus (Polydesmida, Polydesmidae), Strongylosoma stigmatosum (Polydesmida, Paradoxosomatidae), Epibolus pulchripes (Spirobolida, Pachybolidae), and two species of the order Spirostreptida, Archispirostreptus gigas and Telodeinopus aoutii. The hepatic cells are absent in P. lagurus in which the midgut epithelium is surrounded by the visceral muscles. In the other species they were arranged around the midgut as coherent hepatic cells of mesenchymal character. Each hepatic cell possessed its own basal lamina and formed the cellular processes which protrude into the basal lamina of the midgut to make contact with midgut digestive cells. Accumulation of reserve material in different millipede taxa has been described with the special emphasis on the process of autophagy in the cytoplasm of the hepatic cells. The lack of hepatic cells may represent an ancestral condition within millipedes.

Three-dimensional chitin rings from body segments of a pet diplopod species: Characterization and protein interaction studies

Physicochemical characterization of new chitin isolates can provide valuable insights into designing of biomimetic materials. Chitin isolates with a definite three-dimensional (3D) structure can exhibit characteristics that distinguish them from other chitin specimens that are in form of powder or flakes without a definite and uniform shape. Herein, 3D chitin rings were produced from body segments of a diplopod (Archispirostreptus gigas) inhabiting tropical regions. This organism is cultured easily and can reach 38cm in length, which makes it a suitable source for isolation of chitin. The chitin rings were characterized via TGA, FT-IR, SEM and XRD analyses. Enzymatic digestion test with chitinase demonstrated that chitin isolates had high purity (digestion rate: 97.4%). The source organism had high chitin content; 21.02±2.23% on dry weight. Interactions of the chitin rings with bovine serum albumin (BSA) protein were studied under different conditions (pH: 4.0–8.0, chitin amount: 6–14mg, contact time: 30–360min, protein concentration: 0.2–1mg/mL). The highest BSA adsorption was observed at pH5.0 at 20°C. The adsorption equilibrium data exhibited a better fit to Langmuir adsorption and the pseudo-first order kinetic models. The findings presented here can be useful for further studies aiming to develop biocompatible and nontoxic biomaterials.

The ultrastructure of the midgut epithelium in millipedes (Myriapoda, Diplopoda)

The midgut epithelia of the millipedes Polyxenus lagurus, Archispirostreptus gigas and Julus scandinavius were analyzed under light and transmission electron microscopies. In order to detect the proliferation of regenerative cells, labeling with BrdU and antibodies against phosphohistone H3 were employed. A tube-shaped midgut of three millipedes examined spreads along the entire length of the middle region of the body. The epithelium is composed of digestive, secretory and regenerative cells. The digestive cells are responsible for the accumulation of metals and the reserve material as well as the synthesis of substances, which are then secreted into the midgut lumen. The secretions are of three types – merocrine, apocrine and microapocrine. The oval or pear-like shaped secretory cells do not come into contact with the midgut lumen and represent the closed type of secretory cells. They possess many electron-dense granules (J. scandinavius) or electron-dense granules and electron-lucent vesicles (A. gigas, P. lagurus), which are accompanied by cisterns of the rough endoplasmic reticulum. The regenerative cells are distributed individually among the basal regions of the digestive cells. The proliferation and differentiation of regenerative cells into the digestive cells occurred in J. scandinavius and A. gigas, while these processes were not observed in P. lagurus. As a result of the mitotic division of regenerative cells, one of the newly formed cells fulfills the role of a regenerative cell, while the second one differentiates into a digestive cell. We concluded that regenerative cells play the role of unipotent midgut stem cells.

Molecular phylogeny of the Thyropygus allevatus group of giant millipedes and some closely related groups

Giant cylindrical millipedes of the family Harpagophoridae, especially species of the genus Thyropygus, are broadly distributed in Thailand and nearby countries. They show a great deal of variation in body size, color patterns and gonopodal characters. Phylogenetic analyses of 26 nominate species from six genera in the subfamilies Harpagophorinae and Rhynchoproctinae, as well as nine new morphotypes (regarded as new species), were performed with the DNA sequences from two mitochondrial gene fragments (16S rRNA and COI). The genus Thyropygus (Harpagophorinae) was recovered as monophyletic under all analyses, whilst the representatives of Rhynchoproctinae also formed a monophyletic group. However, the analyses suggested that the tribe Gonoplectini should be elevated to a separate subfamily, Gonoplectinae. The molecular analyses were largely (but not totally) congruent with, and so supported the usefulness of, gonopodal characters for the classification and identification of harpagophorid millipedes, and additionally supported previous studies on the delimitation of species and subgroups. This is the first molecular study inside the family Harpagophoridae and provides the basis for further studies of the evolutionary processes and biogeographic patterns of millipedes in Southeast Asia.

Population dynamics of a threatened giant millipede: implications for restoration

AbstractThe population dynamics of threatened invertebrates have important implications for their conservation and restoration. The Seychelles giant millipede (SGM), Sechelleptus seychellarum, is a threatened and functionally important macro‐detritivore endemic to the Seychelles granitic islands. Here, we studied the population dynamics of the SGM from 1998 to 2009 on Cousine Island, Seychelles, to make practical restoration recommendations. Large fluctuations in millipede densities were found between 1998 and 2009. In 2002, 2003, 2005, and 2007 millipede densities were low, while densities were high in 1998 and 2009. Although the SGM is active all year round, millipede surface activity was positively correlated with rainfall, with millipede density high during the wet NW monsoon period (i.e., October to April) and low during the SE trade wind period (i.e., May to September). Female:male:juvenile ratios were ∼3:1:1. The implications of these results for restoration are that translocations should preferably be done in years of high millipede densities and during the wet season. Furthermore, chemical control of the invasive ant Pheidole megacephala, which is currently being carried out on Cousine Island and in future could be conducted on other Seychelles islands, should preferably be done during the low rainfall months, as the SGM readily consumes the hydramethylnon‐based bait.

Molecular and functional characterization of hemocyanin of the giant African millipede,Archispirostreptus gigas

In contrast to other terrestrial arthropods, where gaseous O2 that fuels aerobic metabolism diffuses to the tissues in tracheal tubes, and most other metazoans, where O2 is transported to tissues by circulating respiratory proteins, the myriapods (millipedes and centipedes) strikingly have tracheal systems as well as circulating hemocyanin (Hc). In order to elucidate the evolutionary origin and biological significance of millipede Hc, we report the molecular structure (subunit composition and amino acid sequence) of multimeric (36-mer) Hc from the forest floor-dwelling giant African millipede Archispirostreptus gigas and its allosteric oxygen-binding properties under various physico-chemical conditions. Archispirostreptus gigas Hc consists of only a single subunit type with differential glycosylation. Phylogenic analysis revealed that millipede Hc is a sister group to centipede HcA, which supports an early divergence of distinct Hc subunits in myriapods and an ancient origin of multimeric Hcs. Archispirostreptus gigas Hc binds O2 with a high affinity and shows a strong Bohr effect. O2 binding is, moreover, modulated by Ca(2+) ions, which increase the O2 affinity of the Hc in the tense (T; deoxygenated) as well as the relaxed (R; oxygenated) states, and by (l)-lactate, which modulates Hc-O2 affinity by changing the allosteric equilibrium constant, L. Cooperativity in O2 binding at half O2 saturation (n50) is pH dependent and maximal at ~pH 7.4, and the number of interacting O2-binding sites (q) is markedly increased by binding Ca(2+). The data are discussed in the light of the mutually supplementary roles of Hc and the tracheal system for tissue O2 supply.

Beyond vegetation-based habitat restoration for a threatened giant Spirostreptid millipede

Vegetation-based restoration efforts often fail to provide suitable habitat for many invertebrates. Restoring habitat for target invertebrates requires an understanding of their resource and condition requirements. The Seychelles giant millipede (SGM), Sechelleptus seychellarum, is a functionally important millipede which is Red Listed as ‘Vulnerable’. Here, we studied the habitat requirements of this macro-detritivore in relation to the ongoing forest restoration programme on Cousine Island, Seychelles. SGM spatial density did not significantly correlate with edaphic and litter properties, but did positively correlate with the toposcape (i.e. elevation and granitic rock cover). Granite rock crevices in forest covered areas were important diurnal refuges for the SGM, as microclimate conditions in non-forest covered rock refuges were unsuitable. SGM physical condition was significantly lower in non-shaded crevices compared to those shaded by forest. Low granite rock cover in the restored forest limited the SGM colonisation of this area in large numbers, despite canopy cover in the restored forest being comparable with that in the reference natural forest. As most restoration practices are primarily vegetation-based, this study demonstrates that such an approach can be inadequate for restoring habitat for target invertebrates, as many species habitat requirements extend beyond that of vegetation. For the SGM, selecting restoration sites that already have abundant rock cover would be the most practical way to increase SGM habitat through forest restoration practices. Taking into consideration the habitat requirements of target invertebrates can help in setting or redirecting restoration goals and thus enhance the conservation value of such practices.

A Behavioural Ecology Approach to Assessing the Effect of Alien Vegetation on a Threatened Giant Millipede

The importance of taking into account behavioural ecology when assessing the impacts of habitat deterioration on threatened species is evaluated. The Seychelles giant millipede (SGM), Sechelleptus seychellarum, is IUCN Red-Listed as “Vulnerable”. Here, we studied the effect of alien bamboo and coconut on the behavioural ecology of this species on Cousine Island, Seychelles. Coconut did not affect SGM density, but negatively affected its foraging behaviour, whereas bamboo negatively affected both its density and foraging behaviour. The SGM showed feeding preferences for Pisonia grandis and Ficus sp. leaf litter types. Alien bamboo and coconut pose a varied threat to the SGM, and their removal and replacement by indigenous forest species (e.g. P. grandis and Ficus sp.) should form part of an island’s restoration programme.

Response of a threatened giant millipede to forest restoration

Understanding the response of threatened and functionally important island invertebrate species to plant community restoration is essential for the successful conservation of these invertebrate species. The Seychelles giant millipede (SGM), Sechelleptus seychellarum, is a threatened and functionally important macro-detritivore endemic to the Seychelles granitic islands. Here, we studied the response of this species to the ongoing forest restoration programme on Cousine Island, Seychelles. This study was conducted over an 11 year period, representing the floral succession of a formerly degraded open scrubland area, dominated by alien plants, to a closed canopy forest, dominated by planted indigenous trees. While the time span of this study was insufficient for vegetation height in the restored area to equal that in the reference natural forest, canopy closure was nevertheless comparable. We found SGM density to be an order of magnitude lower in the restored site compared to the natural forest. In contrast, SGM physical condition improved significantly in the restored site, as vegetation structure increased. Furthermore, SGM behaviour in the restored site switched from a predominantly walking to a dominantly feeding behaviour over the study period, resulting in the forest restoration programme on Cousine increasing the foraging area of the SGM by 43 %. Competition for key resources, e.g. food and day-time refuges, are suggested as possible factors limiting SGM numbers in the restored forest.

Effect of an invasive ant and its chemical control on a threatened endemic Seychelles millipede

The impact of invasive species on island faunas can be of major local consequence, while their control is an important part of island ecosystem restoration. Among these invasive species are ants, of which some have a disruptive impact on indigenous arthropod populations. Here, we study the impact of the invasive African big-headed ant, Pheidole megacephala, on a small Seychelles island, Cousine, and assess the impact of this ant, and its chemical control, using the commercially available hydramethylnon-based bait, Siege, on the endemic keystone Seychelles giant millipede species, Sechelleptus seychellarum. We found no significant correlations in landscape-scale spatial overlap and abundance between the ant and the millipede. Furthermore, the ant did not attack healthy millipedes, but fed only on dying and dead individuals. The chemical defences of the millipede protected it from ant predation. Ingestion of the bait at standard concentration had no obvious impact on the millipede. The most significant threat to the Seychelles giant millipede in terms of P. megacephala invasion is from possible catastrophic shifts in ecosystem function through ant hemipteran mutualisms which can lead to tree mortality, resulting in alteration of the millipede's habitat.

The Rhynchoproctinae, a south-east Asiatic subfamily of giant millipedes: cladistic analysis, classification, four new genera and a deviating new species from north-west Thailand (Diplopoda:Spirostreptida:Harpagophoridae)

The subfamily Rhynchoproctinae is revised at the generic level. Four new genera and a remarkable new species are described: Alienostreptus, gen. nov. from Vietnam, Armatostreptus, gen. nov., Heptischius, gen. nov., and Heptischius lactuca, sp. nov. from Thailand, and Prominulostreptus, gen. nov. from China. Agaricogonopus Zhang & Zhang, 1997, is removed from synonymy with Junceustreptus Demange, 1961. All 14 genera of the subfamily share three diagnostic gonopodal characters: the posterior surface of the anterior coxal fold forms a deep concavity, there is a mesal flap on the gonopod coxa, and the posterior coxal fold is very low. A cladistic analysis based on morphological data supports monophyly of Rhynchoproctinae and the distincness of the newly described genera. A dichotomous key to the genera is included.