European Journal of Taxonomy 15: 1-12

http://dx.doi.org/10.5852/ejt.2012.15

BY

This work is licensed under a Creative Commons Attribution 3.0 License.

ISSN 2118-9773

www. europeanj ournaloftaxonomy. eu

2012 • Henrik Enghoff

Research article

hree new species of Dolichoiulus millipedes from the underground

of Gran Canaria, with notes on the circumscription of the genus

(Diplopoda, Julida, Julidae)

Henrik ENGHOFF

Natural History Museum of Denmark, University of Copenhagen,

Universitetsparken 15, DK-2100 Copenhagen OE, Denmark.

Email: henghoff@snm ku.dk

Abstract. Dolichoiulus typhlocanaria sp. nov., D. oromii sp. nov. and D. longunguis sp. nov. are

described from caves and the mesovoid shallow stratum (MSS) on Gran Canaria. The genus Anagaiulus

Enghoff, 1992 is synonymized under Dolichoiulus Verhoeff, 1900, resulting in Dolichoiulus hlancatypa

(Enghoff, 1992) comb nov.

Key words. Millipede, Canary Islands, new species, cave. Gran Canaria.

Enghoff H. 2012. Three new species of Dolichoiulus millipedes from the underground of Gran Canaria, with notes

on the circumscription of the genus (Diplopoda, Julida, Julidae). European Journal of Taxonomy 15: 1-12. http://

dx.doi.org/10.5852/eit.2012.15

Introduction

The millipede genus Dolichoiulus Verhoeff, 1900 includes one of the largest among the many radiations

of plants and animals on the Canary Islands (Fernandez-Palacios 2011). Of the 56 species hitherto

included in the genus, 46 are Canarian endemics (Enghoff 1992, 2011; Enghoff & Baez 1993). Enghoff

(1992) revised the genus and described 39 new species. Despite considerable collecting activity on the

Canary Islands since 1992, no additional new species have been discovered until now.

Five Canarian Dolichoiulus species have been found in caves, three endemic species on Tenerife, one

endemic species on El Hierro and the introduced D. typhlops Ceuca, 1973 on La Palma and El Hierro

(Enghoff 1992, 2002). The three Teneriffan cave species are blind, like the introduced D. typhlops. In

recent years, blind Dolichoiulus millipedes have been collected in several caves and MSS (mesovoid

shallow stratum, Romero 2009) on Gran Canaria. This material includes three new species and is the

subject of the present paper. One of the new species shows a character that necessitates a re-evaluation

of the genus Anagaiulus Enghoff, 1992, a monotypic genus from Tenerife.

Material and methods

Specimens were collected in MSS with special pitfall traps designed for collecting in this habitat

(see Lopez & Oromi 2010), whereas those captured in caves and mines which were found by visual

exploration. All were preserved in 70% ethanol and were examined with a stereomicroscope. A few

1

European Journal of Taxonomy 15: 1-12 (2012)

specimens were ‘analysed’, i.e., gnathochilarium, an antenna, and some midbody legs were temporarily

mounted in glycerol and examined at high magnification in a compound microscope. Gonopods/vulvae

of some specimens were s im ilarly placed in temporary mounts. Specimens for scanning electron

microscopy were transferred to 96% ethanol and then to acetone, air-dried, mounted on aluminium

stubs, coated with platinum/palladium and studied in a JEOL JSM-6335F scanning electron microscope

(ZMUC).

Terminology follows Enghoff (1992) except that the term ‘ring’ or ‘body ring’ is used instead of

‘segment’, and ‘mesomerital process’ instead of‘mesomerite’.

Abbreviations used in the text

DZUL

GE

H

L

MCNT

MELANSIS

MSS

ZMUC

GIET

Department of Animal Biology (Zoology), University of La Laguna, Tenerife,

Canary Islands

Grupo de Espeleologia

height (vertical diameter) of body

length of body

Museum of Natural Sciences of Tenerife, Canary Islands

Sociedad Entomologica Canaria Melansis, Gran Canaria, Canary Islands

mesocavernous shallow stratum, or milieu souterrain superficiel

Natural History Museum of Denmark (Zoological Museum), University of Copenhagen

Grupo de Investigaciones Espeleologico de Tenerife

Results

Class Diplopoda Blainville-Gervais, 1844

Order Julida Leach, 1814

Family Julidae Leach, 1814

Genus Dolichoiulus Verhoeff, 1900

Dolichoiulus Verhoeff, 1900: 85.

Nesopachyiulus Attems, 1902: 599, 607.

Trichopachyiulus Verhoeff, 1910: 194.

Anagaiulus Enghoff, 1992: 144 syn. nov.

See Enghoff (1992) for an in-depth discussion of Dolichoiulus. Anagaiulus was erected by Enghoff

(1992) for a species which differs from the Dolichoiulus species in having a whorl of metazonital setae.

Enghoff (1992) realized that this is a poor diagnostic character but preferred not to ‘dilute’ the already

rather vague concept of Dolichoiulus by including a species with metazonital setae. Since then, two lines

of evidence have emerged that contradict separate status for Anagaiulus'.

1. One of the new species described in the present paper has metazonital setae; the other doesn’t, but

apart from this, and a size difference, the two species are as good as identical.

2. A preliminary molecular phylogenetic analysis by I. Sanmartin (unpublished) suggests that

Anagaiulus is nested deeply inside Dolichoiulus.

Diagnosis

(from Enghoff 1992, emended): Pachyiulini. Eyes present or absent. No frontal setae. Mostly without

metazonital setae (exceptions: D. blancatypa (Enghoff, 1992) comb. nov. and D. oromii sp. nov.).

Metazonital striae mostly distinct also dorsally. No grooves in suture. No (or almost no) preanal

projection. Preanal pilosity marginal or extensive. Anal valve pilosity extensive. Male mandibles mostly

without lobes. First and second male legs typically julid. Male legs mostly with ventral pads. Anterior

2

ENGHOFF H., Three new Dolichoiulus millipedes

gonopod spoon-shaped, with a mesal ridge, a lateral prominence, and two apical denticles (which may

be fused or reduced). Posterior gonopod without a sperm fovea, with a mesomerital process and with

two opisthomerital processes, posterior process varying from short-triangular to flagelliform; one or

both opisthomerital processes sometimes more or less reduced.

Included species

Dolichoiulus hlancatypa (Enghoff, 1992), comb. nov. ex Anagaiulus, D. typhlocanaria sp. nov.,

D. oromii sp. nov., D. longunguis sp. nov., plus the 56 species included in Enghoff (1992).

Dolichoiulus typhlocanaria sp. nov.

Figs 1, 2A, 3A, B, 4A, B

Diagnosis

A blind species of Dolichoiulus. Differs from other blind Dolichoiulus species as shown in Table 1.

Differs from the other blind Dolichoiulus on Gran Canaria in size (Figs 1, 5). Differs from D. oromii

sp. nov. in lacking metazonital setae, in having only marginal preanal pilosity, and in having a terminal

projection on the mesal ridge of the anterior gonopod. Differs from D. longunguis sp. nov. and the

widespread D. typhlops in having clearly slenderer anterior gonopods (Fig. 3A, B, cf. Fig. 3D).

Etymology

The name (noun in apposition) refers to the blindness of this species from Gran Canaria.

Type locality

GRAN CANARIA (Canary Islands), Barranco del Draguillo.

Material examined (all from Canary Islands: Gran Canaria)

Holotype

3 , Barranco del Draguillo, UTM 28R 4561/30913 (27°56 , 45 ,, N, 15°26 , 45 ,, W), MSS, 26 Dec. 2006, H.

Fopez leg. (DZUF).

Paratypes

3 33 , 2 $ ?, same locality, 16 Aug. 2007, H. Fopez leg. (1 3 , 1 ? DZUF; 2 3 , 1 $ ZMUC); 3 33 , 2

55, same locality, 26 Dec. 2010, H. Fopez leg. (ZMUC); 9 33 , 6 5 5> same locality, 30 Dec. 2011, H.

Fopez leg. (DZUF).

Referred (non-type) material

2 S3, 1 5 , Cuevade laFuna, UTM 28R 4478/30873 (27°54 , 36 ,, N, 15°31’48”W), 23 Dec. 2006, GIET

leg. (ZMUC); 2 33 Cueva de los Arrepentidos, UTM 28R 4482/30878 (27°54 , 51 ,, N, 15 0 3r35”W),

26 Dec. 2006, GIET leg. (DZUF); 1 & 2 55, Eos Majateles, UTM 28R 4505/30915 (27°56 , 08 ,, N,

15°30’10”W), MSS, 26 Dec. 2010, H. Fopez leg. (DZUF).

Total material

21 33 , 13 55 (1 3 , 1 5 analyzed).

Description

S' L 14-20 mm, H 1.03-1.26 mm, F/H 13-16, 33-52 podous rings + 1-2(3?) apodous rings in front of

telson.

5: F 15-20 mm, H 1.17-1.41 mm, F/H 12-16, 36-52 podous rings + 2 apodous rings in front of telson.

3

European Journal of Taxonomy 15: 1-12 (2012)

Fig. 1. Size diagram for Dolichoiulus typhlocanaria sp. nov. The diagram shows number of podous

(leg-bearing) body rings (p.r., x axis) and vertical body diameter in mm (y axis). For a given number of

podous rings D. typhlocanaria sp. nov. is thicker than the two other species (see Fig. 5), and females are

thicker than males.

Fig. 2. A D. typhlocanaria sp. nov., male from Cueva de la Luna, midbody rings. Scale 0.1 mm. B. D.

oromii sp. nov., detail of posterior margin of midbody ring, showing diagnostic setae. Scale 0.05 mm.

4

ENGHOFF H., Three new Dolichoiulus millipedes

Colour. The preserved specimens are uniform whitish yellow. Defence glands in part visible by

transparency as dark spots.

Head. No eyes. 4 supralabral setae. Setae on gnathochilarial stipes: 3 apical, 2-3 (c?) / 0 (5) nonapical.

Fength of ante nn ae 156% of H in S, 124% of H in §.

Body rings. (Fig. 2A) Prozonites posteriorly furrowed, at least in specimens from Cueva de la Funa.

Metazonites unvaulted, with c. 12 striae per dorsal quarter. Ozopore c. !4 metazonite length behind

suture. Fimbus cell length c. 4x width; their free margin with a few tiny denticles (as in Enghoff, 1992:

figs 28-29).

Fegs. Fength 94% of H in S, 78% of H in §. Claw: length 6-8% of leg, length/height 3.4-5.2. Acces¬

sory claw 0.4x shorter than claw.

Preanal ring. Without a projection, pilosity marginal.

Male

Mandibular stipites. Without lobes.

Fegs. Second pair with ventral pads on postfemur and tibia. Postfemoral pads tiny, tibial pads small on

midbody legs. Second pair with coxal pores.

Anterior gonopods. (Fig. 3 A, B) Slender, basal shaft more than half as broad as distal bowl. Ridge with

a terminal projection. Fateral prominence distinct. Bowl slender, rounded-triangular. Apical denticles

distinct.

Posterior gonopods. (Fig. 4A, B) Mesomerital process almost as long as opisthomerite, straight.

Opisthomerite slender, straight. Anterior process short, triangular, pointed. Posterior process longer,

slender.

Female

Receptaculum seminis. A stalked sphere (as in Enghoff 1992: fig. 48).

Distribution and habitat

Endemic to Gran Canaria. Known from two caves and three MSS in the east-central part of the island.

Dolichoiulus oromii sp. nov.

Figs 2B, 3C, 4C, 5

Diagnosis

A blind species of Dolichoiulus. Differs from other blind Dolichoiulus species as shown in Table 1.

Differs from all other Dolichoiulus species, except D. blancatypa (Enghoff, 1992) comb, nov., in having

metazonital setae. Differs further from the other blind Dolichoiulus on Gran Canaria in size (Figs 1,

5), in having extensive preanal pilosity, and in lacking a terminal projection on the mesal ridge of

the anterior gonopod. Differs from D. longunguis sp. nov. and the widespread D. typhlops in having

slenderer anterior gonopods (Fig. 3C, cf. Fig. 3D). Differs from D. blancatypa (from Tenerife) in having

the mesomerital process clearly separated from the opisthomerite.

Etymology

The species in named in honour of Pedro Oromi, Fa Faguna, Tenerife, in recognition of his vast effort

to document and analyse the Canarian fauna.

5

Table 1. Characteristics of blind Dolichoiulus species. Data in part from Enghoff (1992, 2002). Relative lengths of antennae and legs are given in

percentage of body diameter. Relative length of claw is given in percentage of total leg length. Diagnostic or near-diagnostic characters shown in

underlined boldface . There are further subtle gonopod differences between the species.

European Journal of Taxonomy 15: 1-12 (2012)

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6

ENGHOFF H., Three new Dolichoiulus millipedes

Type locality

GRAN CANARIA (Canary Islands), El Sao.

Material studied

all from Canary Islands: Gran Canaria.

Fig. 3. Anterior gonopods, posterior views. A. D. typhlocanaria sp. nov., specimen from Barranco

Draguillo. B. D. typhlocanaria sp. nov., specimen from Cueva de la Funa. C. D. oromii sp. nov.

D. D. typhlops Ceuca, 1973, specimen from Fa Palma, right anterior gonopod. ad: apical denticles, lp:

lateral prominence, mr: mesal ridge, p: projection. Scales 0.1 mm (A-C), 0.01 mm (D).

7

European Journal of Taxonomy 15: 1-12 (2012)

Holotype

6 El Sao (Agaete), UTM 28R 4354/31049 (28°04 , 06 ,, N, 15°39’27”W), MSS, 26 Dec. 2005, H. Lopez

leg. (DZUL).

Paratypes

3 SS, 2 5?, same data. (2 SS, 2 55 DZUL, 1 S ZMUC); 1 $, same loc.. May 2005, H. Lopez leg.

(DZUL); 8 SS, 3 ??, 1 juv., same loc., 28 Dec. 2006, H. Lopez leg. (1 S 1 ? DZUL, 6 <$<$ 1 ? 1 juv.

ZMUC, 1 $ 1 $ MCNT); 2 $ $, same loc., 11 Aug. 2007, H. Lopez leg. (DZUL), 5 SS, 1 ?, same loc.,

7 Jan. 2009, H. Lopez leg. (DZUL); 1 3, 1 5 , same loc., 3 Jan. 2010, H. Lopez leg. (DZUL); 2 <$<$,

same loc., 4 Dec. 2010, H. Lopez leg. (DZUL).

Referred (non-type) material

1 5, Caldera de los Marteles, UTM 28R 4481/30920 (27°57’01”N, 15 o 31’40”W), MSS, 19 Aug. 2006,

H. Lopez leg. (MCNT); 1 §, same loc., 7 Nov. 2008, H. Lopez leg. (DZUL); 2 5 55> same

loc., 27 Dec. 2008, H. Lopez leg. (DZUL); 1 5 , Los Majateles, UTM 28R 4505/30915 (27°56 , 08 ,, N,

15°30 , 10”W), MSS, 26 Dec. 2010, H. Lopez leg. (DZUL).

Total material

23 SS, 17 55, 1 juv. (1 (?, 1 5 analyzed).

Description

S'- L ll- 21 1 / 2 mm, H 0.82-1.00 mm, L/H 13-21, 31-53 podous rings + 1-4 apodous rings in front oftelson.

5: L 14-19 mm, H 0.95-1.08 mm, L/H 15-18, 42-47 podous rings + 1-2 apodous rings in front oftelson.

Colour. The preserved specimens are uniform whitish yellow. Defence glands in part visible by

transparency as dark spots.

Fig. 4. Posterior gonopods, mesal views. A. D. typhlocanaria sp. nov., specimen from Barranco

Draguillo. B. D. typhlocanaria sp. nov., specimen from Cueva de la Luna. C. D. oromii sp. nov. ap:

anterior process, pp: posterior process, mp: mesomerital process. Scales 0.1 mm.

8

ENGHOFF H., Three new Dolichoiulus millipedes

Head. No eyes. 4 supralabral setae. Setae on gnathochilarial stipes: 3 apical, 5 (c?) / 0 ($) nonapical.

Fength of antennae 162% of H in 123% of H in §.

Body rings. Prozonites not furrowed. Metazonites unvaulted, with ca. 12 striae per dorsal quarter and a

whorl of short setae on the posterior margin (Fig. 2B). Ozopore % - % metazonite length behind suture.

Fimbus cells length slightly more than 2x width; their free margin with a few tiny denticles (as in

Enghoff 1992: figs 28-29).

Fegs. Fength 84% of H in S, 24% ofH in §. Claw: length 7-8% of leg, length/height 4.4-5.4. Accessory

claw 0.4-0.5x shorter than claw.

Preanal ring. Without a projection, pilosity extensive.

Male

Mandibular stipites. Without lobes.

Fegs. Second pair with ventral pads on postfemur and tibia. Postfemoral pads tiny, tibial pads small on

midbody legs. Second pair with coxal pores.

Gonopods. (Figs 3C, 4C) As in D. typhlocanaria sp. nov., except for the lack of a terminal projection on

the mesal ridge of the anterior gonopod.

Female

Receptaculum seminis. A simple sack (as in Enghoff 1992: fig. 47).

Distribution and habitat

Endemic to Gran Canaria. Known only from the type locality in the NW part of the island, and from two

further MSS sites in the East-Central part.

Dolichoiulus longunguis sp. nov.

Fig. 5

Diagnosis

Ablind species of Dolichoiulus. Differs from other blind Dolichoiulus species as shown in Table 1. Differs

from the other blind Dolichoiulus on Gran Canaria in size (Figs 1, 5). Differs from D. typhlocanaria

sp. nov. and D. oromii sp. nov. in having broader anterior gonopods, further from D. oromii sp. nov. in

lacking metazonital setae and in having only marginal preanal pilosity, and further from D. typhlocanaria

sp. nov. in lacking a terminal projection on the mesal ridge of the anterior gonopod. Differs from the

widespread D. typhlops by being slenderer (Fig. 1, cf. Enghoff 2002: fig. 1).

Etymology

The name (noun in apposition) meaning “long claw” and refers to a distinguishing character of this

species.

Material studied

Holotype

c?, GRAN CANARIA, Mina Eos Roques, UTM 28R 4476/30943 (28°0 , 0”N, 15°3F56”W), 18 Mar.

2012, M. Naranjo leg. (DZUF).

9

European Journal of Taxonomy 15: 1-12 (2012)

Paratype

1 5, same loc., 23 Jan. 2012, M. Naranjo leg. (DZUL).

Total material

1 (?, 1 ? (analyzed).

Description

f \ L 13 mm, H 0.71 mm, L/H 19, 51 podous rings + 2 apodous rings in front of telson.

L 13 mm, H 0.72 mm, L/H 18, 49 podous rings + 1 apodous ring in front of telson.

Colour. The preserved specimens are uniform pale brown.

Head. No eyes. 4 supralabral setae. Setae on gnathochilarial stipes: 3 apical, 4 (c?) / 0 ($) nonapical.

Length of antennae 138% of H in <$, 126% of H in $.

Body rings. Prozonites not furrowed. Metazonites unvaulted, striae quite indistinct dorsally (not studied

with SEM). Ozopore c. 0.4 metazonite length behind suture. Limbus not studied (requires SEM).

Legs. Length 65% of H in S, 61% of H in §. Claw: length 11% of leg, length/height 4.5-5.4. Accessory

claw 0.3-0.4* shorter than claw.

Preanal ring. Without a projection, pilosity marginal.

Male

Mandibular stipites. Without lobes.

Number of podous body rings

Fig. 5. Size diagram for Dolichoiulus oromii sp. nov. and D. longunguis sp. nov. The diagram shows

number of podous (leg-bearing) body rings (p.r., x axis) and vertical body diameter in mm (y axis). For

a given number of podous rings D. oromii sp. nov. is thicker than D. longunguis sp. nov. but thinner than

D. typhlocanaria sp. nov. (see Fig. 1); specimens of D. oromii sp. nov. from East-Central Gran Canaria

are thicker than those from the Northwest (El Sao), and females are thicker than males.

10

ENGHOFF H., Three new Dolichoiulus millipedes

Fegs. Second pair with ventral pads on tibia only. Postfemoral pads absent, tibial pads tiny on midbody

legs. Second pair with coxal pores.

Anterior gonopods. Indistinguishable from the one shown in Fig. 3D, broader than in D. typhlocanaria

sp. nov. and D. oromii sp. nov. Mesal ridge without a terminal projection. Fateral prominence distinct.

Bowl broad, lateral margin regularly convex. Apical denticles distinct.

Posterior gonopods. Indistinguishable from those of D. typhlocanaria sp. nov. and D. oromii sp. nov.

Female

Receptaculum seminis. A stalked sphere (as in Enghoff 1992: fig. 48).

Distribution and habitat

Endemic to Gran Canaria. Known only from the type locality in the east-central part of the island.

Collected in an artificial gallery. The short legs and long claws in this species may suggest that it is a

soil-digger species rather than a true cave species ( cf. Enghoff 1982). Among other Dolichoiulus species

claws as long as those of D. longunguis sp. nov. are found only in D. blancatypa (Enghoff, 1992) comb,

nov. from Tenerife, which also has short legs and is a soil-dweller (Enghoff 1992).

Discussion

It is perhaps not surprising that the first new Dolichoiulus species to be found after Enghoff (1992)

are from the underground of Gran Canaria. Although caves are scarce on Gran Canaria, there are

many deep artificial old galleries such as Mina de los Roques which have produced an interesting

subterranean fauna (Naranjo et al. 2009). Also, the mesovoid (or mesocavernous) shallow stratum

(MSS) is richly represented, and recent collecting activities here have revealed a rich endemic fauna of,

e.g., Lagynochthonius Beier, 1951 pseudoscorpions, Symploce Hebard, 1916 cockroaches and Oromia

Alonso-Zarazaga, 1987 weevils (Oromi et al. 2010; Mahnert, 2011). The abundant artificial caves and

MSS spots not yet prospected suggest that the underground of Gran Canaria still has not been exhausted

in terms of new millipede species.

Acknowledgements

Thanks to Pedro Oromi and Heriberto Lopez (La Laguna) and to Manuel Naranjo (Las Palmas) for

placing these specimens (and numerous other interesting Canarian millipedes) at my disposal, to GE

Tebexcorade-La Palma and MELANSIS for collections during the project “Invertebrate fauna from the

caves of Gran Canaria”, and to Isabel Sanmartin, Madrid, for access to her unpublished phylogenetic

analysis.

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Manuscript received: 28 March 2012

Manuscript accepted: 9 May 2012

Published on: 24 May 2012

Topic editor: Rudy Jocque

In compliance with the ICZN , printed versions of all papers are deposited in the libraries of the institutes

that are members of the EJT consortium: Museum National d’Histoire Naturelle, Paris, France; National

Botanic Garden of Belgium, Meise, Belgium; Royal Museum for Central Africa, Tervuren, Belgium;

Natural History Museum, London, United Kingdom; Royal Belgian Institute of Natural Sciences,

Brussels, Belgium; Natural History Museum of Denmark, Copenhagen, Denmark.

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