Research Article |
Corresponding author: Megan Short ( mshort@deakin.edu.au ) Academic editor: Ivan H. Tuf
© 2015 Megan Short.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Short M (2015) New records of Lophoproctus coecus Pocock, 1894 (Diplopoda, Polyxenida, Lophoproctidae) extend the range of the genus Lophoproctus. In: Tuf IH, Tajovský K (Eds) Proceedings of the 16th International Congress of Myriapodology, Olomouc, Czech Republic. ZooKeys 510: 209-222. https://doi.org/10.3897/zookeys.510.8668
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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.
Penicillata , millipedes, geographic distribution, Caucasus, Crimea, Italy
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.
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.
In the Caucasian and Crimean regions the previous records of Lophoproctus are by Lignau. He collected Lophoproctus in Krasnodar Polyanna (
In
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).
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 (
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.
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 (
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.
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, (
Records from literature. Nervi, suburb of Genova, Italy, on the open hill-side behind the town, beneath stones, alt. 400–500 ft. (
Unpublished record. Nice (pers. comm.
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.
Records from literature. Palalda (now Amélie-les-Bains-Palalda), Pyrénées-Orientales, France, in soil under litter layer in oak woods, leg. Chalande (
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.
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 (
Distribution. France (mainland and Corsica), Spain (mainland and Majorca), Italy (Capri), Malta, Lebanon, Israel.
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 (
Records from literature never formally identified (pers. comm. Nguyen Duy-Jacquemin, 2014): Pădurea Comorova, Romania (
Further record from literature, identification uncertain. Marine de Sisco, Corsica, France, 3 Sept 1942, leg. P. Remy (
Record from literature. Kalimantsi, South Pirin Mountains,Bulgaria, in ant nests 1 Mar 2003, leg. Lapeva-Gjonova (
Records from literature. Cueva de Genova near to Palma, Majorca, Balearic Islands, Spain (
Lophoproctus cf. pagesi : “Gruta do Fumo”, Parque Natural da Arrábida, Portugal (
Distribution. Caves on Majorca (L. cf. pagesi – cave in Portugal).
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
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
The distribution map (Fig.
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.
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.
Record details of all Lophoproctus collections.
Data type: Comma-separated values files (CSV)
Explanation note: A record of all specimens of Lophoproctus examined, or mentioned in the literature, is available.