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Research Article
New records of Lophoproctus coecus Pocock, 1894 (Diplopoda, Polyxenida, Lophoproctidae) extend the range of the genus Lophoproctus
expand article infoMegan Short
‡ Deakin University, Burwood, Australia
Open Access

Abstract

The geographic distribution of the genus Lophoproctus Pocock, 1894 has greatly expanded with new records of the species Lophoproctus coecus Pocock, 1894, together with the reassignment of a number of millipedes formerly identified as Lophoproctus lucidus (Chalande, 1888). L. coecus was found to be the sole representative of the family Lophoproctidae in collections examined from Crimea and the Caucasian region. The species was also identified from Iran and Kyrgyzstan. Lophoproctus specimens collected in Italy by Verhoeff were reassigned as L. coecus with the exception of one specimen of L. jeanneli (Brölemann, 1910) from Capri. These data were combined with all available information from the literature to look at the pattern of distribution of the four species in the genus. The range of the genus Lophoproctus extends from Portugal to Central Asia. Lophoproctus coecus is widespread from Italy eastward, while the morphologically very similar species L. lucidus is confined to France and northern Africa. The two species have a narrow overlap in the Alpes Maritimes region of France. L. jeanneli has a scattered coastal distribution around the Mediterranean Sea. The troglobitic species L. pagesi (Condé, 1982) has only been recorded from a cave on Majorca, Spain.

Keywords

Penicillata , millipedes, geographic distribution, Caucasus, Crimea, Italy

Introduction

Genera and species in the family Lophoproctidae Silvestri, 1897 have very similar morphology, with species adapted to an endogenous mode of life, being found in soil, caves and under stones. All Lophoproctidae lack ocelli, their integument lacks pigmentation and the 8th antennal article is elongate. Species in the family also share the same arrangement of caudal trichomes and similar organisation of tergal trichomes (Fig. 1).

Figure 1. 

Habitus drawing of Lophoproctus coecus Pocock, 1894 showing typical morphology of the family Lophoproctidae.

Characters used to determine genus and species of the family can be difficult to observe and include number and arrangement of antennal sensilla, number and arrangement of linguiform processes along the anterior margin of the labrum, structure of the telotarsus, leg setae, and tarsal spine. There are currently 5 genera: Lophoproctus Pocock, 1894, Lophoturus Brolemann, 1931, Ancistroxenus Schubart, 1947, Lophoproctinus Silvestri, 1948, and Alloproctoides Marquet & Condé, 1950. Two further genera Barroxenus Chamberlin, 1940 and Trichoproctus Silvestri, 1899, known only from single collections, are of uncertain status as they are inadequately described.

Pocock (1894) established the genus Lophoproctus for a species collected from soil at Nervi in Liguria, Italy. As the species lacked ocelli he called it coecus (‘blind’ in latin). Previously in 1888, Chalande had described the species Pollyxenus (sic) lucidus from Palalda, Eastern Pyrenees, France, which he initially described as having ocelli. In 1894 Silvestri identified Polyxenus lucidus from Italy, then later the same year recognising that the specimens had no ocelli he moved the species into the genus Lophoproctus. He further suggested that L. coecus and L. lucidus were synonymous (Silvestri 1894b) as did Verhoeff some years later (Verhoeff 1921). Both Silvestri and Verhoeff collected widely throughout Italy (Silvestri 1894a, 1894b, Verhoeff 1921, 1952) and identified all lophoproctids they found as L. lucidus with the exception of those from Capri that Verhoeff mistakenly described as a new species Lophoproctus litoralis (Verhoeff, 1952). L. litoralis was later determined to be Lophoproctus jeanneli (Brölemann, 1910) (Condé 1969). Condé (1978) re-examined material from Verhoeff’s collection from Isernia and Teramo, Zannone (Pontine Islands) and Sardinia, Italy, and noted that they differed from L. lucidus in that they had a different arrangement of sensilla on antennal article VI which was also more elongate. On the basis of Condé’s description, Nguyen Duy-Jacquemin (1993) confirmed that these specimens were L. coecus.

In the Caucasian and Crimean regions the previous records of Lophoproctus are by Lignau. He collected Lophoproctus in Krasnodar Polyanna (Lignau 1903), Crimea (Lignau 1905) and in Gagri (Lignau 1914) and although in his earlier papers he had identified the specimens incorrectly, in his 1911 paper he identified all as Polyxenus (Lophoproctus) lucidus (Lignau 1911). Subsequent species lists published all include Lophoproctus lucidus (Lohmander 1936, Kobakhidze 1965, Talikadze 1984) presumably based on Lignau’s early collections.

In 1993, Nguyen Duy-Jacquemin redescribed L. coecus from syntypes from Nervi, Italy that together with specimens collected in Zannone, Italy by Verhoeff and from Rome, Italy by Silvestri, confirmed that L. lucidus and L. coecus were not synonymous. Her paper clearly illustrates that the two species differ in arrangement of sensilla on antennal article VI (Fig. 2), structure of the median lobe on the anterior edge of the labrum, the number of ridges on the leg setae and the ratio of the length of the tarsal spine to length of the claw (Fig. 4). Initially described as a subspecies of L. lucidus, L. jeanneli is also found in the Mediterranean region of Europe but is easily distinguishable from other species of Lophoproctus by the presence of a denticle on the claw of the telotarsus.

Figure 2. 

Typical pattern of basiconic sensilla on the right 6th antennal article. A Lophoproctus coecus B Lophoproctus lucidus. The coeloconic sensilla are not visible due to angle of view. Scale bar: 20 µm (A, B).

Figure 3. 

Lophoproctus coecus (Pocock, 1888), Krasnodar Province, Russia. Illustration of diagnostic features. A labrum showing triangular median linguiform process B typical setae on coxa C right antenna showing articles VI–VIII with sensilla D right antennal article VI showing arrangement of sensilla E tarsus 2 and telotarsus. Scale bars: 10 µm (A, B); 50 µm (C); 20 µm (D, E).

Figure 4. 

Lophoproctus coecus (Pocock, 1888) Krasnodar Province, Russia. A Lateral view showing antenna B Dorso-lateral view showing tergal trichomes. Scale bars: 1 mm.

The most recently described species Lophoproctus pagesi Condé, 1982, is a troglobitic species collected in caves on the island of Majorca, Spain. Specimens similar to L. pagesi have also been collected from caves in Portugal. L. pagesi differs from other species of Lophoproctus in details of the labrum, antennal sensilla and telotarsus. As well it has elongate antennae and legs typical of troglobitic species (Condé 1982, Nguyen Duy-Jacquemin 1993).

In this study I re-assessed specimens from The Zoological State Collection, Munich collected by Verhoeff in Italy and identified as L. lucidus. I also identified Lophoproctidae from Crimea, Caucasus, Iran and Kyrgyzstan in the collection of the Zoological Museum of Moscow. These data were then combined with details obtained from the published literature on Lophoproctus species to determine distribution of species in the genus.

Methods

The material examined for this study is lodged in the Zoological Museum of Moscow and the Zoological State Collection in Munich, Germany.

Specimens from the Zoological Museum of Moscow were examined and identified. All specimens were preserved in ethanol. These specimens were examined by light microscopy and scanning electron microscopy. For light microscopy, specimens were mounted on slides in Hoyer’s medium, dried at 60 °C and examined with an Olympus CX 41 compound microscope. Scanning electron micrographs were obtained of selected whole specimens that were dehydrated in a graded series of ethanol, 80%, 90% and 100%, then air-dried. Specimens were then mounted on stubs using adhesive tabs, sputter-coated with gold and examined with a Philips XL20 scanning electron microscope. Photographs of whole specimens were taken with a Leica Integrated Stereo-microscope System comprising a Leica 205C microscope with a DFC425 camera and 5000HDI dome illuminator. Images were stacked using Leica Application Suite and enhanced using Adobe Photoshop CS6.

Specimens from the Zoological State Collection, Munich, are slide mounts in Canada Balsam made by KW. Verhoeff. The slides lack both date of collection and site habitat details. The slides were examined by light microscopy using an Olympus CX41 compound microscope. Due to the thickness of the slide mounts, they could not be examined at magnifications higher than 400×.

As no coordinates were available for most of the material examined, Google Earth was used to provide an estimate of geographical position for mapping purposes (a table of localities with coordinates is available in supplementary material). A map of the distribution of all species in the genus Lophoproctus was generated using SimpleMappr (Shorthouse 2010). New records determined in this study were included together with all known published records. Many records in the literature, especially those by Verhoeff, Silvestri and Tabacaru are questionable and these have been treated separately.

Results

Sixty collections of Polyxenida in the Zoological Museum of Moscow were examined and L. coecus identified in 15. In most cases less than 5 specimens were collected at a site. No other species of Lophoproctidae were found.

Twenty slides from the Zoological State Collection, Munich (ZSM/Myr. 20031594–612, 615) all labelled as L. lucidus were examined and 19 found to be L. coecus. Slide ZSM/Myr. 20031615 contained a whole mount of L. jeanneli.

The geographic distribution of all known localities of the genus Lophoproctus was plotted using the data listed below (Fig. 5).

Figure 5. 

Map indicating geographic distribution of Lophoproctus species. Legend: red star = L. coecus; yellow circle = L. lucidus; white star = L. coecus/L. lucidus? (many in similar localities as L. coecus and hence hidden); aqua triangle = L. jeanneli; purple square = L. pagesi; purple hexagon = L. cf. pagesi; white circle = Lophoproctus sp. indet. Map created using SimpleMappr, http://www.simplemappr.net, (Shorthouse 2010).

Systematics

Order Polyxenida Lucas, 1840
Family Lophoproctidae Silvestri, 1897
Lophoproctus Pocock, 1894

1. Lophoproctus coecus Pocock, 1894

Records from literature. Nervi, suburb of Genova, Italy, on the open hill-side behind the town, beneath stones, alt. 400–500 ft. (Pocock 1894); Grotta di San Antonino, Finale Ligure, Italy, leg. Ascenso 1950 (Nguyen Duy-Jacquemin 1993); Grotta di San Antonio =Antonino, Finale Ligure, Italy, leg. Comotti Baldan 13 Aug 1986 (Nguyen Duy-Jacquemin 1993); Grotta di Arma do Rian, Finalborgo, Italy, leg. Franciscolo 16 Mar 1952 (Nguyen Duy-Jacquemin 1993); Zannone, Ponziane Islands, Italy, leg. Condé 28–29 Jan 1966, 26–27 Feb 1966, 17 Jan 1967 (Nguyen Duy-Jacquemin 1993); Grotta di Nettuno, Porto Conte, Alghero, Sardinia, Italy, near the pier, 4 Oct 1955, leg. Condé (Nguyen Duy-Jacquemin 1993); Isernia, Italy, leg. Verhoeff (Condé 1982), ? same specimen as listed below; Teramo, Italy, leg. Verhoeff (Condé 1982), ? same specimen as listed below; Villa Pamphyli, Rome, Italy, leg. Silvestri Nov 1893 (Silvestri 1894a, reclassified Nguyen Duy-Jacquemin 1993).

Unpublished record. Nice (pers. comm. Nguyen Duy-Jacquemin 2014).

Re-examined material collected in Italy by Verhoeff (date of collection known for only 2 specimens). ZSM/Myr-20031594, Tivoli, Lazio; ZSM/Myr 20031595, Isernia, Molis; ZSM/Myr-20031596,Teramo, Abruzzo; ZSM/Myr-20031597, Portofino, Genova, Liguria; ZSM/Myr-20031598, Portofino, Genova, Liguria, molt; ZSM/Myr-20031599, Italy, molt; ZSM/Myr-20031600, Elba, Livorno, Toscana; ZSM/Myr-0031601, Chiesetal, Vestone, Brescia, Lombardia; ZSM/Myr-20031602, Isernia, Molise; ZSM/Myr-20031603, Ferrania, Liguria, 01.07.1933; ZSM/Myr-20031604, Mele, Genova, Liguria; ZSM/Myr-20031605, Monte Cimino, Soriano, Viterbo, Lazio; ZSM/Myr-20031606, Porto Santo Stefano, Grosseto, Toscana; ZSM/Myr-20031607, Veneto, Vicenza; ZSM/Myr-20031608, Ferrania, Liguria; ZSM/Myr-20031609, Santuario, Savona, Liguria; ZSM/Myr-20031610, Frigido, Toscana, from under stones at a mill ruin, April 1907; ZSM/Myr-20031611, illegible labelling; ZSM/Myr-20031612 Capri (No 27), Napoli, Campania.

New material from Zoological Museum of Moscow. Nikita Botanical Garden, Cape Martyan, near Yalta, Crimea, 4 Nov 1947, leg. M.S. Ghilarov; Gurzuf, Yalta, Crimea, Jun–Sep 1947, leg. M.S. Ghilarov (2 vials); Utrish Nature Reserve, Krasnodar province, Russia, oak hornbeam forest, 15 Jun 2013, leg. I. Tuf; Utrish Nature Reserve, Krasnodar province, Russia, hornbeam forest, 14 Jun 2013, leg. I. Tuf; Goryachy Klyuch, Mtn ridge, Markotkh plateau, Krasnodar province, Russia, 3 Jul 1956, leg. M.S. Ghilarov; Dagomys, Sochi, Krasnodar province, Russia, Quercus shrub, Carpinus, Fagus etc., 18 May 1983, leg. S. Golovatch; on road 2 km N of Dagomys, Krasnodar province, Russia, 2 Jun 2014, leg. M. Potapov; Cave “Our Lady”, ca 8 km from Khosta Sochi, Krasnodar province, Russia, Buxus, Fagus, Acer etc., forest near entrance, litter and under stones, 16 May 1985, leg. S. Golovatch; Khosta, Sochi region, Krasnodar province, Russia, Sambucus, 26 Jun 1956, leg. M.S. Ghilarov; Ris Forest, Bobcai east, Gumista River, Abkhazia, Russia, litter, 5 Jun 1982, leg. J. Bohàc; environs of Keda, Adjaria, Georgia, Picea and deciduous forest, 1 Oct 1975, A. Druk; Nedzura River valley 8km SE of Akhaldaba, Borzhomi district, Georgia, Picea, Carpinus and Fagus forest, litter, logs, 12 May 1983, leg. S. Golovatch; Arslanbob, Fergana mountain range, environs of Yarodar, Kyrgyzstan, dry limestone slopes with grass, under stones, 28 Sep 1983, leg. K. Eskov; Sari, Mazanderan province, Iran, Quercus and Carpinus forest, 11 Apr 2013, leg. M. Mehrafrooz.

Distribution. South-East France, Italy, Russia, Georgia, Iran and Kyrgyzstan.

2. Lophoproctus lucidus (Chalande, 1888)

Records from literature. Palalda (now Amélie-les-Bains-Palalda), Pyrénées-Orientales, France, in soil under litter layer in oak woods, leg. Chalande (Chalande 1888); cave Gourgue,Canton Aspet, No. 229, Haute-Garonne, France, (Nguyen Duy-Jacquemin 1993); Baumo de las Fadas, Canton du Barjac, Dept du Gard, France, 26 Aug 1909, leg. Brölemann (Brölemann 1910); Albères, France, 1926, leg. Brolemann (Condé 1950); Banyuls sur Mer, France, leg. Brölemann (Nguyen Duy-Jacquemin 1993); Hyères, France leg, A Dollfus (Nguyen Duy-Jacquemin 1993); Ariège, France, leg. Nguyen Duy-Jacquemin (Nguyen Duy-Jacquemin 2000); Souk el Arba, Jendouba district, Tunisia, 30 Mar 1896, leg. Silvestri (Silvestri 1896); La Pérouse (now known as Tamentfoust), Dar El Beïda district of Algiers, Algeria, in wave washed and dry plant material (Seurat 1930); Marrakech, Morocco, 10 Dec 1950, (Condé 1954); Marrakech, Morocco, Jardin de l’ Aguedal, near the Mechouar (Condé 1954); Marrakech, Morocco, Jardin de la Bahia (Condé 1954); Marrakech, Morocco, Parc de la Villa Majorelle, under flower pots and stones, 10 Dec 1950 (Condé 1954).

Unpublished records. El Ghazalaf Ariana, nr. Tunis, Tunisia, garden of private house in earth, beneath Cydonia tree (pers. comm. N. Akkari, 2014).

New material. La Parc Phoenix, Nice, France, in cold greenhouse, Jan 2014, leg. JM Lemaire.

Distribution. France, Tunisia, Algeria, Morocco.

3. Lophoproctus jeanneli (Brölemann, 1010)

Re-examined material collected by Verhoeff (date not given). ZSM/Myr-20031615: Corsica, France.

Records from literature. Baume (grotto) du Colombier, Alpes-Maritimes, commune de Roquefort-les-Pins, canton de Bar-sur-Loup, France, 17 Sep 1905, 27 Sep 1908 (Brölemann 1910); Grotte de la Chèvre d’Or, canton de Bar-sur-le-Loup, France 25 Nov 1987, leg. V. Aellen, (Condé 1989); City park, Barcelona, Spain, Sep 1950 leg. Condé (Condé 1954); Lower Gravona River, left bank of western arm of river, opposite Canapajolo, Corsica, France (Condé 1953); Pointe de Porticcio, sur la côte S. du golfe d’Ajaccio, near houses, 1 km to south west of Fallaccioli, Corsica, France (Condé 1953); Togna, commune de Sari-di-Porto Vecchio, at the edge of a ravine and a garden well, Corsica, France, (Condé 1953); Cueva de la Moriguilla (Vilacarrilo), Andalucia, Spain (Golovatch and Mauries 2013); Esporlas near Palma, Majorca, Balaeric Islands, Spain, at irrigation canal overgrown with dry compact rootlets 17 Aug–23 Sept 1954, leg. J. Pagès (Condé 1955); near Bagno di Tiberio, beach, Capri, Italy, leg. Verhoeff (Condé 1969); Malta leg. Silvestri (Nguyen Duy-Jacquemin 1993, with reservation); near Dékouané, 7 Km to east of Beirut, Lebanon April 1952, leg. PJ. Corset, (Condé 1954, Nguyen Duy-Jacquemin 1993 with reservation); Tel Dan, Israel, 26 Dec 1963, leg. G. Levy (Condé and Nguyen Duy-Jacquemin 1970).

Distribution. France (mainland and Corsica), Spain (mainland and Majorca), Italy (Capri), Malta, Lebanon, Israel.

4. Lophoproctus lucidus/coecus

Specimens identified as L. lucidus but likely to be L. coecus as Silvestri and Verhoeff thought the two species were synonymous.

Records from literature. Bevagna, Umbria, Italy, in meadow and forest, late October 1893, leg. Silvestri (Silvestri 1894a); Woods at Madama and Acquacetosa, near Rome, Italy, on ground in plant debris, in forest and open places, leg. Silvestri, November 1893 (Silvestri 1894a); at Colle Pezzo, Mt. Martano, Umbria, Italy 15 Oct 1893, leg. Silvestri (Silvestri 1894a); Medol Casello, Lombardy, Italy, on floor of cave 1935 –1940, leg. various unnamed (Manfredi 1940): Sicily, Italy, leg. Silvestri (Silvestri 1898); Mt. Schignano, Italy, in soil among plant debris, not only in forests, but also in open places, 15 Oct 1893, leg. Silvestri (Silvestri 1894a); Syracuse, Sicily, Italy, 1962–1968, leg. Institute of Zoology of Catania (Strasser 1970); Ciminà, Aspromonte, Calabria, Italy 25 Oct 1966, leg. G. Osella (Strasser 1970); St Remo, Italy under stones in an olive terrace 7–21 April 1907, leg. Verhoeff (Verhoeff 1921); Pegli, Italy in creek valley 7–21 April 1907, leg. Verhoeff (Verhoeff 1921); Massa, Carrara, Italy, in sandstone gorge 7–21 April 1907, leg. Verhoeff (Verhoeff 1921); St. Margherita, Italy, in chestnut wood 7–21 April 1907, leg. Verhoeff (Verhoeff 1921).

5. Lophoproctus sp. indet., reported as Lophoproctus lucidus

Records from literature never formally identified (pers. comm. Nguyen Duy-Jacquemin, 2014): Pădurea Comorova, Romania (Tabacaru 1966); Mangalia, Romania, 20 Nov 1963 (Tabacaru 1966); Pădurea Hagieni, Romania, 17 May 1963, (Tabacaru 1966); Canaraua de pe Graniţă - comuna Băneasa, Romania 2 Aug 1962, leg. Dumitrescu et al. (Tabacaru 1966); Cave Gura Dobrogii litoclazic, Romania 17 Jun 1963 leg. Dumitrescu et al. (Tabacaru 1966); Cave Gura Dobrogii, Romania, at entrance 17 Sep 1963 leg. Dumitrescu et al. (Tabacaru 1966); Casian, Romania 27 Jul 1962, 7 May 1963, 30 Aug 1964 leg. Dumitrescu et al. (Tabacaru 1966).

Further record from literature, identification uncertain. Marine de Sisco, Corsica, France, 3 Sept 1942, leg. P. Remy (Verhoeff 1943).

6. Lophoproctus sp. indet

Record from literature. Kalimantsi, South Pirin Mountains,Bulgaria, in ant nests 1 Mar 2003, leg. Lapeva-Gjonova (Stoev and Lapeva-Gjonova 2006)

7. Lophoproctus pagesi (Condé, 1982)

Records from literature. Cueva de Genova near to Palma, Majorca, Balearic Islands, Spain (Condé 1982); Cueva de Bellver, Palma, Majorca, Balearic Islands, Spain, unpublished, Condé det. (Nguyen Duy-Jacquemin 1993);

Lophoproctus cf. pagesi : “Gruta do Fumo”, Parque Natural da Arrábida, Portugal (Cardoso et al. 2008, Nguyen Duy-Jacquemin 2014).

Distribution. Caves on Majorca (L. cf. pagesi – cave in Portugal).

Discussion

Lophoproctus coecus has previously been considered to occupy a scattered range within the Central Mediterranean region, but the results of this study indicate that the species is widespread throughout Europe particularly in Eastern Europe with its distribution extending into Central Asia. L. lucidus in comparison seems limited to the Southern France, as well as Morocco, Algeria and Tunesia in Northern Africa. The identification of L. jeanneli from Capri, Italy reinforces the Mediterranean coastal distribution previously noted by Kime and Enghoff (2011). It is of interest that species within the genus may overlap in their geographic distribution with both L. jeanneli and L. coecus being found in Capri and in the Alpes Maritimes region of France. Lophoproctus pagesi and L. jeanneli both occur on the island of Majorca, with L. pagesi restricted to caves while L. jeanneli was found in humid, sunny hilly areas.

A number of identifications were unable to be checked. In the case of the Lophoproctus identified by Lignau as L. lucidus, specimens identified as L. coecus in this study were found at all 3 of Lignau’s collection areas, indicating that it is most probable that the specimens collected by Lignau were in fact L. coecus. Prior to publication of Nguyen Duy-Jacquemin (1993), the difference between L. coecus and L. lucidus was not understood with Silvestri (1894b) considering the two species synonymous. Hence, until it can be confirmed, the identification of specimens of Lophoproctus as L. lucidus, from Romania (Tabacaru 1966) and Sicily (Silvestri 1903, Strasser 1970) must remain questionable. Lophoproctus has also been collected from Bulgaria but has not yet been identified to species (Stoev and Lapeva-Gjonova 2006).

The distribution map (Fig. 5) indicates very clearly that there is a big gap in our knowledge of Lophoproctus in Greece, the Balkans and Turkey. It is predicted that L. coecus does occur in these three locations, and that L. jeanneli may also occur in coastal regions. All species except the troglobitic L. pagesi have the ability to live in many of the habitats in these areas of Europe and Asia. Unfortunately there is limited information available to guide collection of these tiny millipedes. In most cases millipedes in this study were collected by hand collecting in the field, or by sieving of litter and/or soil followed by direct collection from a tray of sieved material (Fig. 6). Recently in Dagomys, Russia, L. coecus was collected from forest litter by funnel extraction (M. Potapov pers comm. 2014). Habitats from which Lophoproctus has been collected vary from maquis and forest litter, top layer of soil, cave floors, to under stones and logs, and in ants nests (Stoev and Lapeva-Gjonova 2006). In the case of L. coecus from Kyrgyzstan, the single specimen was collected from under stones on a dry grassy limestone slope, a similar habitat to that of the type collected by Pocock (1894), ‘open hillside beneath stones’. As well as the above mentioned habitats, L. lucidus has been found in North African cities in city parks, in gardens and under pot plants and stones (N Akkari pers comm. 2014, Condé 1954), and near Algiers on the beach in accretions of both dry and damp marine plant material (Condé 1954). Lophoproctus jeanneli has been collected in abundance from cracks in bricks and under dry stones on the ground in heavy shade in the Barcelona City Park (Condé 1954) as well as on the sea shore (Verhoeff 1952).

Figure 6. 

Ivan H. Tuf and Daria Kuznetsova collecting invertebrates including Lophoproctus coecus from sieved soil and litter, Utrish Nature Reserve, Krasnodar province, Russia.

Methods of dispersal have not been studied, but it is probable that the presence of L. lucidus in North African cities is due to anthropogenic activities dating back to the French colonial period as millipedes confirmed to be L. lucidus appear limited to Southern France and centres of French colonial activity in North Africa. Polyxenida are thought to passively disperse via the wind and incidental attachment to the feathers of birds. It is likely in the case of soil and litter dwelling lophoproctids that dispersal via wind is less common as they do not appear to be living in elevated situations in trees and bushes in contrast to Polyxenida from the families Synxenidae and Polyxenidae. However, limited methods of dispersal do not seem to have restricted the geographic ranges of species of the genus Lophoproctus, especially L. coecus.

Acknowledgements

I am grateful to Sergei Golovatch for providing me with the opportunity to study the Crimean and Caucasian Polyxenida from the Zoological Musum of Moscow and to Mikhail Potapov and Ivan Tuf for some very recent specimens from the same region. Thanks also to Cuong Huynh and Stefan Friedrich for access to Verhoeff’s Lophoproctus slides, to Khanh Tran and Nick Porch for assistance with SEM and photomontage imaging. I greatly appreciated the improvements to the manuscript suggested by Jean-Francois David and an anonymous reviewer. Finally many thanks to Monique Nguyen Duy-Jacquemin for her continuing advice and support. This study was funded by the author.

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Supplementary material

Supplementary material 1 

Record details of all Lophoproctus collections.

Author: Megan Short

Data type: Comma-separated values files (CSV)

Explanation note: A record of all specimens of Lophoproctus examined, or mentioned in the literature, is available.

This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.
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