European Journal of Taxonomy 231: 1-19

http://dx.doi.org/10.5852/ejt.2016.231

BY

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

ISSN 2118-9773

www. europeanj ournaloftaxonomy. eu

2016 • Bonato L. et al.

Research article

urn:lsid:zoobank.org:pub:4C501A9D-C90B-4132-BDQ7-C71A9076E7D0

An unusually elongate endogeic centipede

from Sardinia (Chilopoda: Geophilidae)

Lucio BONATO >•*, Marzio ZAPPAROLI 2 , Leandro DRAGO 3 & Alessandro MINELLI 4

1 3 4 Dipartimento di Biologia, Universita di Padova, viaUgo Bassi 58B, 1-35131 Padova, Italy.

2 Dipartimento per la Innovazione nei sistemi Biologici, Agroalimentari e Forestali (DIBAF),

Universita della Tuscia, via S. Camillo de Lellis s.n.c., 1-01100 Viterbo, Italy.

* Corresponding author: [email protected]

2 E-mail: [email protected]

3 E-mail: leandro.dragol @ gmail.com

4 E-mail: [email protected]

1 urn:lsid:zoobank.org:author:DEFC50E9-439C-45Al-8EFE-EFFE745D391F

2 urn:lsid:zoobank.org:author:30E6BE85-ClDA-4215-A07D-652C9770AEC9

3 urn:lsid:zoobank.org:author:84133787-C29F-496D-B3D5-lE4372093FD4

4 urn:lsid:zoobank.org:author:D9FA639F-7538-441C-9572-032FD6C4EEDE

Abstract. Endogeophilus ichnusae gen. et sp. nov. (Chilopoda: Geophilidae sensu stricto ) is described

based on three specimens from two localities in south-western Sardinia, examined by light and scanning

electron microscopy. The new centipede resembles the rare Ibero-Pyrenean genus Galliophilus Ribaut &

Brolemann, 1927 in some features, especially in the forcipular segment, and the temperate European

species Geophilus electricus (Linnaeus, 1758) in other features, especially in the ultimate leg-bearing

segment. However, the true affinities of E. ichnusae gen. et sp. nov. are uncertain, because the new

species departs significantly from the majority of geophilids for the higher number of legs (91-107

pairs in the specimens examined), the slender trunk segments (the stemites being longer than wide),

the relatively stout legs (the tarsus being only about twice as long as wide) and the very short setae

(< 15 pm) scattered on the body surface. All these features are probably derived and suggest adaptation

to a more strictly endogeic habit than other geophilids.

Key words. Geophilidae, morphology, taxonomy, Sardinia, endogeic.

Bonato L., Zapparoli M., Drago L. & Minelli A. 2016. An unusually elongate endogeic centipede from Sardinia

(Chilopoda: Geophilidae). EuropeanJournal of Taxonomy 231: 1-19. http://dx.doi.org/10,5852/eit.2016,231

Introduction

Sardinia is a large Mediterranean island, hosting a high number of soil-dwelling arthropod taxa. A

significant portion of the pedofauna is exclusive of the island, or of the Corsica-Sardinia complex, and

many of these endemic species are deeply divergent from the closest relatives from other areas (e.g.,

AA.VV. 1983). The exploration of the island and the description of its soil fauna are still ongoing, and

1

European Journal of Taxonomy 231 : 1-19 ( 2016 )

recent samplings and investigations are adding significantly to its knowledge (e.g., Cerretti et al. 2009;

Nardi et al. 2011).

Centipedes (Chilopoda) are well represented in the Sardinian soil communities and have been broadly

surveyed in the past (Minelli 1983; Zapparoli 2009, 2011). Recently, however, an unprecedented

sampling effort dedicated to the endogeic fauna in south-western Sardinia has provided a small sample

of specimens of an undescribed geophilid centipede, which is remarkably unusual in many respects

(Zapparoli 2009,2011). The new species is here described and illustrated by means of light and scanning

electronic microscopy, it is compared with other geophilids and its unique morphological features are

tentatively interpreted in terms of ecological adaptation to the endogeic environment.

Material and methods

Specimens were collected by Luca Fancello while sampling endogeic beetles with the following

technique (Fancello et al. 2009; Zapparoli 2011): soil was extracted up to a maximum depth of about

50 cm, the superficial humus layer was removed when present, and detritus was sorted in Berlese-type

funnels modified according to Pace (1996).

Specimens were fixed in 70% ethanol and examined by a Leica DMLB microscope after inclusion

in ethylene glycol in temporary mounts, following standard protocols for geophilomorph centipedes

(Pereira 2000). Measures were taken by means of a micrometre eyepiece. Digital photographs were

taken using a Leica DFC420 camera applied to the microscope and assembled using the image stacking

software CombineZP (Hadley 2008). Line-drawings were produced manually from the photographs.

Scanning electron microscopy was applied to selected body parts of a single specimen, after dehydration

through a graded ethanol series (80%, 95%, 100%, at least 1 hour each step), critical-point drying with

CCf and coating with gold. Sonication was avoided to prevent loss of setae. The samples were examined

with a Cambridge Stereoscan 260 microscope.

The holotype of the new species was compared directly to specimens of another 20 species selected

from different genera of Geophilidae. To control for possible sexual differences, only adult females

were selected. The following measures were taken: head width and length of the longest seta on the

cephalic plate; length and width of a metasternite and of a leg tarsus at about 20% of the total series

of leg-bearing segments. All measures were taken by one person (L.B.) using the same microscope.

Specimens examined: Abatorus allredi Chamberlin, 1965, California, UCRC06; Acanthogeophilus

spiniger Meinert, 1870, Morocco, PD5153; A re nophilus peregrinus Jones, 1989, Great Britain, PD5754;

Chomatophilus sp. Pococlc, 1896, Mexico, UNAM003; Clinopodes flavidus C.L. Koch, 1847, Italy,

PD 1269; Diphyonyx conjungens (Verhoeff, 1898), Turkey, PD 1144; Eurygeophiluspinguis (Brolemann,

1898), Italy, PD4254; Galliophilus beatensis Ribaut & Brolemann, 1927, France (no direct examination,

but data from Brolemann 1927); Geophilus alpinus Meinert, 1870, Italy, PD680; G. carpophagus Leach,

1815, Canary Islands, PD2058; G. electricus (Linnaeus, 1758), Italy, PD569; G. fucorum Brolemann,

1909, Italy, PD3156; G. piae Minelli, 1983, Italy, PD3148; G. pygmaeus Latzel, 1880, Italy, PD638;

G. richardi Brolemann, 1904, Italy, PD5776; G. truncorum Bergsoe & Meinert, 1866, Great Britain,

PD943; Pleurogeophilus mediterraneus (Meinert, 1870), Italy, PD 1154; Sogona sp. Chamberlin, 1912,

Mexico, PD 1382; Stenotaenia sorrentina (Attems, 1903), Italy, PD5758; Tuobaposeidonis (Verhoeff,

1901), Malta, PD947.

For the morphological terminology, we followed Bonato et al. (2010).

For the collections, we used the following abbreviations:

2

BONATO L. et al., A new elongate geophilid centipede from Sardinia

MSNG = Museo civico di Storia naturale “Giacomo Doria”, Genova, Italy

MZ = coll. M. Zapparoli

PD = coll. A. Minelli & L. Bonato

UCR = University of California, Riverside, U.S.A.

UNAM = Instituto de Biologia, Universidad Nacional Autonoma de Mexico, Mexico City, Mexico

Results

Class Chilopoda Latreille, 1817

Order Geophilomorpha Pococlc, 1896

Family Geophilidae Leach, 1815

Endogeophilus gen. nov.

um:lsid:zoobank.org:act:0904F67C-219A-459F-A121-3D0C7E47D469

Figs 1-4; Table 1

Type species

Endogeophilus ichnusae gen. et sp. nov.

Diagnosis

Geophilids with body almost uniform in width, only slightly tapering at both ends; head slightly

longer than wide; clypeus uniformly areolate, without clypeal areas; intermediate part of the labrum

bearing a few tubercles; labral side-parts distinct from the clypeus, well separated from each other,

their posterior margin with bristles; first maxillae with bi-articulated telopodites and two pairs of

short lappets; second maxillary coxosternite with long isthmus, without inner processes, and lacking

distinctly sclerotized ridges; second maxillary telopodites composed of three articles, bearing a simple,

sub-conic claw; forcipular tergite sub-trapezoid, posteriorly about as wide as the subsequent tergite,

the lateral margins distinctly converging anteriorly; forcipular coxosternite slightly wider than long,

without anterior denticles, chitin-lines complete, coxopleural sutures complete and distinctly diverging

anteriorly; forcipules stout and strongly tapering, with two distinct intermediate articles: forcipular

tarsungula abruptly narrowing, relatively short and only weakly curved, with a single small basal

denticle; carpophagus pits shallow; leg-bearing segments with a sub-ovoid/sub-triangular pore-field on

the posterior half of each metasternite; leg claws bearing two slender accessory spines; ultimate leg¬

bearing segment with pleuropretergite entire, metasternite sub-trapezoid wider than long; coxal pores

mainly opening along the ventro-internal, anterior and dorso-internal sides of the coxopleuron, with an

additional isolated ventral pore; telopodites of the ultimate leg pair composed of six articles, slightly

swollen in the male in comparison with the female, bearing a claw; bi-articulated gonopods in the male,

short gonopodal lamina in the female; a pair of anal pores.

Differential characters with respect to the most similar genera are given in Table 1.

Etymology

Composed of the prefix “endo-”, from the classic Greek svbov (inside), and the genus name Geophilus ,

which is from the classic Greek prefix ye©- (earth) and (pilew (to love), referring to the endogeic habitat.

The gender of the genus name is masculine.

3

European Journal of Taxonomy 231 : 1-19 ( 2016 )

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6

BONATO L. et al., A new elongate geophilid centipede from Sardinia

Endogeophilus ichnusae gen. et sp. nov.

um:lsid:zoobank.org:act:BF3DE3C0-542A-4767-A62E-6C9D6A2923AD

Figs 1—4

Geophilidae sp. nov-Zapparoli 2009: 145, 155.

Geophilidae n. sp-Zapparoli 2011: 240.

Diagnosis

As for the genus (see above).

Etymology

From the Latin Ichnusa , which is from the Greek I/vovcraa, ancient name of Sardinia. The species

epithet is a noun in the genitive case.

Type material

Holotype

SARDINIA: adult §, 34 mm long, with 107 leg-bearing segments, Colle di Campanasissa, between

Siliqua (Province of Cagliari) and Nuxis (Province of Carbonia-Iglesias), 280 m; 27 Dec. 2006,

L. Fancello leg., in ethanol, head detached from trunk (MSNG, code 58344).

Paratype A

SARDINIA: adult S, 32 mm long, with 95 leg-bearing segments, same site and date of the holotype, in

ethanol, head detached from trunk, some pieces of trunk gold-coated for SEM examination (MZ).

Paratype B

SARDINIA: adult S, 31 mm long, with 91 leg-bearing segments, Mamenga, Marganai forest, near

Domusnovas (Province of Carbonia-Iglesias), 610 m, 1 Mar. 2006, L. Fancello leg., in ethanol, not

dissected (PD, code 1373).

Description

Female

Description of holotype (see also Figs 1A-F, 2, 4)

General features. Body remarkably narrow, 34 mm long, uniformly 0.5 mm wide for most part of the

trunk, only slightly narrowing anteriorly along ca 30 most anterior leg-bearing segments and backwards

along ca five most posterior leg-bearing segments, both the first and the ultimate leg-bearing segments

being 0.4 mm wide. Legs relatively short, most of them unifo rml y 0.4 mm long, a few tens of most

anterior pairs decreasing gradually, the legs of the first pair being 0.3 mm long. Colour (in ethanol)

almost uniformly pale yellow, only the forcipular condyles and the distal part of the tarsungula distinctly

darker.

Cephalic capsule. Cephalic plate sub-rectangular, 1.2 times as long as wide, 0.9 times as wide as the

forcipular tergite, the anterior margin slightly angulated with a medial notch, the lateral margins slightly

convex, the posterior margin straight; transverse suture absent; setae rare and short, up to ca 15 pm long.

Clypeus uniformly areolate, without clypeal areas and without plagulae; lateral margins complete; a total

of 6 setae, arranged in a longitudinal series of three pairs on the anterior part of the clypeus. Pleurites

uniformly areolate, each with 2-3 setae on the anterior part. Labrurn composed of an intermediate part,

which is continuous with the clypeus and bears two medial short tubercles and two lateral bristles, and

lateral parts bordered anteriorly by a complete sulcus and posteriorly by a row of bristles.

7

European Journal of Taxonomy 231 : 1-19 ( 2016 )

Antennae. Slender, 3.6 times as long as the head. Intermediate articles up to 2.5 times as long as wide.

Article XIV 1.9 times as long as wide, 1.7 times as long as article XIII and only slightly wider than the

latter. Setae gradually denser and shorter from basal to distal articles, both ventrally and dorsally. Apical

sensilla ca 6 pm long, spear-lilce, without projections, only gently narrowing at about mid-length. Club¬

like sensilla ca 11 pm long, only on article XIV, grouped on the distal parts of both internal and external

Fig. 1 . Endogeophilus ichnusae gen. et sp. nov. A-F. Holotype, $ . A. Anterior part of the body, dorsal

view, setae omitted. B. Head, without antennae, and forcipular segment, dorsal view. C. Leg-bearing

segment 21, dorsal view. D. Leg-bearing segment 21, ventral view. E. Head, without antennae, ventral

view. F. Ultimate leg-bearing segment and postpedal segments, ventral view, setae omitted. G. Paratype

B, S, ultimate leg-bearing segment and postpedal segments, ventral view, setae omitted. Line drawings

based on photos taken at the microscope. Scale bars = 200 pm.

8

BONATO L. et al ., A new elongate geophilid centipede from Sardinia

sides. Three longitudinal rows of proprioceptive spine-lilce sensilla at the bases of articles: ca 3 sensilla

in each row on articles II-V, apparently a single sensillum on each side of articles VII-IX and XI-XIII;

only a single, dorsal sensillum on articles VI, X and XIV. Groups of 1-3 sensilla, similar to the apical

ones, ca 6 pm long, on both dorso-extemal and ventro-intemal position, close to the distal margin of

articles V, IX and XIII.

Mandibles. A single pectinate lamella on each mandible.

First maxillae. Coxosternite entire, without mid-longitudinal sulcus, without setae. Coxal projection

sub-triangular, longer than wide, bearing 1-2 basal setae and ca 5-6 more distal spine-lilce sensilla, the

tip covered with fine scales. Telopodite of two articles, the basal one without setae, the distal one with

2 setae and ca 4 spine-like sensilla, the tip covered with fine scales. Coxostemal and telopodital lappets

present, very short.

Second maxillae. Coxosternite entire, the intermediate part uniformly sclerotized as the remaining

parts; anterior margin widely concave; no sclerotized ridges; setae only close to the anterior margin;

metameric pores featuring as transversal slits, close to the postero-medial margin. Telopodite composed

Fig. 2. Endogeophilus ichnusae gen. et sp. nov., holotype, $, forcipular segment, left half. A. Dorsal

view. B. Ventral view. Scale bar =100 pm.

9

European Journal of Taxonomy 231 : 1-19 ( 2016 )

of three articles, only slightly narrowing towards the tip; pretarsus in the shape of a simple claw, sub¬

conic, only slightly bent, with a small dorsal bulge.

Forcipular segment. Tergite sub-trapezoid, the lateral margins distinctly converging anteriorly, 1.8 times

as wide as long, posteriorly almost as wide as the subsequent metatergite. Pleurites without scapular ridge.

Exposed part of the coxosternite 1.2 times as wide as long; anterior margin slightly projecting anteriorly

with an intermediate shallow concavity, without denticles; coxopleural sutures complete, entirely

ventral, only slightly sinuous, and strongly converging posteriorly; chitin-lines distinct, reaching the

condyles, only slightly curved and converging posteriorly. Basal distance between the forcipules 0.1 of

the maximum width of the coxosternite. Trochanteroprefemur 1.3 times as wide as long, the internal side

much shorter than the external side, without denticles. Forcipular intermediate articles distinct, without

denticles, each with a shallow bulge only. Tarsungulum 1.9 times as long as wide, with a basal sub-conic

tubercle; distal part almost straight, the internal side with a shallow longitudinal groove between two

distinct ridges. Poison calyx sub-spherical, located in the distal part of the trochanteroprefemur.

Fig. 3. Endogeophilus ichnusae gen. et sp. nov., paratype A, f. A. Leg-bearing segment 20, ventral

view. B. Posterior part of leg-bearing segment 21 and anterior part of segment 22, ventral view. C. Right

forcipular tarsungulum, ventral view. D. Right tarsungulum, ventro-mesal view. E. Right tarsungulum,

frontal view. SEM micrographs. Scale bars = 50 pm.

10

BONATO L. et al., A new elongate geophilid centipede from Sardinia

Leg-bearing segments. A total of 107 leg-bearing segments. Metatergite 1 wider than the subsequent

one, lateral margins converging posteriorly, without pretergite. No paratergites. Metasternites slightly

longer than wide or approximately as long as wide, uniformly areolate, with dense micropores; setae

sparse, at most ca 10 pm long, approximately arranged into transversal rows; metasternites of some

anterior segments with a very shallow, bilobate socket on the anterior margin, 0.6 times as wide as

the margin of metastemite, resembling a “carpophagus” pit but no corresponding medial peg on the

posterior margin. Glandular pores on the metasternites from 2 to penultimate, up to 20-25 pores on

each metastemite, loosely arranged in a sub-ovoid/sub-triangular pore-field on the posterior half of

each metastemite; each pore with a sub-circular lumen with a diameter of 1.1-1.3 pm, surrounded by

a distinct ring-like uninterrupted ridge. Procoxa distinctly longer than metacoxa, both sclerites without

glandular pores. Leg claws simple, uniformly bent; a pair of accessory spines, reaching approximately

the mid-length of the pretarsus, the anterior spine slightly longer than the posterior one.

Ultimate leg-bearing segment. Pleuropretergite entire, lacking sutures or sulci or notches, 3.4 times as

wide as long. Metatergite sub-trapezoid, 1.3 times as wide as long, lateral margins convex and distinctly

converging posteriorly, posterior margin slightly convex. Prestemite 5.0 times as wide as long, not

medially constricted. Metastemite sub-trapezoid, 1.2 times as wide as long, anteriorly 1.4 times as wide

as posteriorly, lateral margins anteriorly slightly convex, posteriorly almost straight and converging

backwards, posterior margin slightly concave; setae uniformly sparse. Coxopleuron 2.0 times as long as

Fig. 4. Forcipules of Endogeophilus ichnusae gen. et sp. nov. and the two most similar geophilid

species. Left part of the forcipular segment, ventral view, in adult females. A. E. ichnusae gen. et

sp. nov., holotype, §, original drawing. B. Geophilns electricus (Linnaeus, 1758), original drawing

from a representative specimen, § (see ‘Material and methods’). C. Galliophilus beatensis Ribaut &

Brolemann, 1927, holotype, §, redrawn from Brolemann (1927). Scale bars = 200 pm.

11

European Journal of Taxonomy 231 : 1-19 ( 2016 )

wide, 1.5 times as long as the metasternite; setae uniformly sparse. Coxal organs of each coxopleuron

opening through 13-14 independent pores, which are approximately aligned from the dorsal to the

ventral surface through the anterior side of the coxopleuron: most pores covered by pleuropretergite,

metatergite and presternite; 2 pores exposed on the dorso-lateral side; 6-7 partially exposed on the

ventral side, close to the metasternite; a single pore isolated on the postero-central part of the ventral

side. Telopodite 7.6 times as long as wide, 2.0 times as long and 1.4 times as wide as the penultimate

telopodite; 6 articles, all covered dorsally with sparse longer setae, ventrally with dense shorter setae.

Pretarsus in the shape of a pointed claw, similar but only half the length of the penultimate.

Postpedal segments. Genital pleurosternite entire. Gonopods in the shape of a short slightly bilobate

lamina. Anal organs relatively large, anal pores exposed.

Male

Description of the terminal part of the body of the paratype B (see also Fig. 1G). Same as the holotype,

except for the following characters.

Ultimate leg-bearing segment. Coxopleuron 1.8 times as long as wide; setae distinctly denser close to

the postero-mesal margin. Coxal organs of each coxopleuron opening through 14-15 independent pores.

Telopodite 7.0 times as long as wide, 2.0 times as long and 1.7 as wide as the penultimate.

Postpedal segments. Complete sutures between the genital sternite and the pleurites. Gonopods bi-

articulated, well separated, with a sub-conic penis in between.

Distribution and habitat

Endogeophilus ichnusae gen. et sp. nov. has been found only in two localities, about 25 km from each

other, in the Sulcis-Iglesiente area, SW Sardinia, at 280-610 m above sea level (Fig. 5). The type locality

(Colle di Campanasissa) has a granitic substrate covered with maquis consisting of Pistacia spp. (L.

Fancello, pers. com.), whereas the other locality (Mamenga) has a calcareous substrate covered with

Quercus ilex L. wood (Bardiani 2011).

Fig. 5. Tocalities where Endogeophilus ichnusae gen. et sp. nov. has been collected, in the Sulcis-

Iglesiente, Sardinia. The type locality is indicated by a star.

12

BONATO L. et al., A new elongate geophilid centipede from Sardinia

Discussion

Phyletic position and classification

The new centipede species obviously belongs to the family Geophilidae sensu stricto , as traditionally

circumscribed (Bonato 2011). This is suggested by the combination of major features of the labrum (a

narrow intermediate part with tubercles and two wider lateral parts with bristles), the mandibles (bearing

a single, pectinate lamella), the second maxillae (with sub-conic claws without filaments) and the

gonopods (an undivided lamina in the female). However, a sound phylogenetic analysis to elucidate the

position of the new species within Geophilidae is unfeasible for the time being, because only incomplete

and sometimes unreliable information are available on the morphology of many geophilid genera.

Nevertheless, the new species fits within a subgroup of geophilids that has been variously distinguished

as a subfamily Geophilinae (Attems 1926, 1929) or tribe Geophilini (Brolemann 1930) or subtribe

Geophilina (Brolemann 1909). This subgroup, possibly non monophyletic, differs from the remaining

Geophilidae in the combination of the following characters: head as long as wide or only slightly longer

than wide, second maxillary coxosternite with a continuous broad connection between the two halves

and without paired sclerotized ridges (either statuminia or circumforaminal rings; see Bonato et al.

2011), forcipular tergite as wide as the subsequent metatergite or only slightly narrower than it, and

forcipules not reaching the anterior margin of the head.

Considering all known genera in this subgroup of geophilids (Table 1), the new species differs from them

in one or more characters that are traditionally given taxonomic value at the genus rank, above all in

the forcipular coxosternite (features of coxopleural sutures, chitin-lines, denticles), trunk metasternites

(presence of carpophagus pit and pattern of pore-fields), and ultimate leg-bearing segment (shape of the

metasternite, arrangement of coxal pores and form of the pretarsi). As a consequence, following current

practice in geophilid taxonomy, the new species is classified here in a distinct genus.

Endogeophilus ichnusae gen. et sp. nov. shares most traits with Geophilus electricus (Linnaeus, 1758),

which is the type species of Geophilus Leach, 1814 (a large, heterogeneous genus including ca 140

species inhabiting the entire Holarctic region; see Bonato 2011), and Galliophilus beatensis Ribaut

& Brolemann, 1927, which is the only species in the genus Galliophilus Ribaut & Brolemann, 1927

(recorded only once in the Pyrenees, with two additional records from the central Iberian peninsula:

Brolemann 1927; Garcia Ruiz & Serra 2000) (Table 2; Fig. 4). In particular, E. ichnusae gen. et sp. nov.

resembles G. electricus in the forcipular coxosternite and the ultimate leg-bearing segment, especially

in the arrangement of the coxal pores, which is otherwise uncommon in geophilids (Table 1). However,

E. ichnusae gen. et sp. nov. differs from G. electricus , and from all the other species of Geophilus , in

other characters including the shape of the forcipular tarsungula (distinctly longer and more curved in

all Geophilus species), the shape of the sternal pore-fields (never medially elongated in a sub-triangular

area in Geophilus) and the number of legs (up to 89 pairs recorded in Geophilus species, but usually

much less). Additionally, E. ichnusae gen. et sp. nov. differs from most species of Geophilus other than

G. electricus also in the chitin-lines (incomplete in most Geophilus species) and the arrangement of

coxal pores (all in ventral position in most Geophilus species). On the other hand, E. ichnusae resembles

Galliophilus beatensis in other features that are infrequent or even unknown in other geophilids, i.e., the

stout forcipular tarsungula (Fig. 4) and the elongated trunk segments. However, while broad pore-fields

are easily detected in all three specimens of E. ichnusae , the holotype of G. beatensis was described and

illustrated as lacking recognisable sternal pore-areas: even after adequate preparation for microscopic

examination of the cuticle of a representative anterior metasternite, only rare pores were observed,

along the margins of the metasternites, with at most two paired posterior small clusters (Brolemann

1927, 1930). Further, at difference with E. ichnusae gen. et sp. nov., the coxal pores of the holotype

of G. beatensis were described and illustrated as sparse on the whole ventral and lateral sides of the

13

European Journal of Taxonomy 231: 1-19 (2016)

Table 2. Differences between Endogeophilus ichnusae gen. et sp. nov. and the most similar known species

of Geophilidae. Information for Galliophilus beatensis Ribaut & Brolemann, 1927 is from Brolemann

(1927, 1930). * = Garcia Ruiz & Serra (2000) reported the following values for two specimens from the

central Iberian Peninsula referred to Galliophilus beatensis : 76 (sic) in a S, 73 in a §.

Character

Endogeophilus ichnusae

sp. nov.

Geophilus electricus

(Linnaeus, 1758)

Galliophilus beatensis

Ribaut & Brolemann, 1927

antennal articles

elongate

short

forcipular tarsungulum

narrow, internal margin

moderately angulate

long, internal margin non

angulate

wide, internal margin

distinctly angulate

sternal pore-fields

distinct, longitudinally

elongate

distinct, transversally broad

no (only rare pores)

number of leg pairs

S 91-95 (n = 2), 9 107 (n = 1)

55-75 (n = many)

$ 85 (n = 1)*

maximum body length

34 mm

45 mm

51 mm

arrangement of coxal pores

mainly along metatergite and

metasternite, plus 1 isolated

ventral

mainly along metatergite and

metasternite, plus 1 isolated

ventral

scattered, on ventral and

lateral surface

pretarsus of ultimate leg

present (both sexes)

absent (at least in female)

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metasfernite log-area

Fig. 6. Variation in the elongation of a single leg-bearing segment in a sample of species of Geophilidae,

including Endogeophilus ichnusae gen. et sp. nov. The length/width ratio was measured on a metasternite

at about 20% of the longitudinal series of leg-bearing segments, and the area of the metasternite (length x

width, both in pm, log-transformed) has been taken as a proxy for body size. Full names of species in

‘Material and methods’.

Ge. p/'ae

•

Endog^philus

Ga. beatensis- T .,

• Ge. fucorum Tu. poseidoms

• • •

Di. conjungens

•

Ac. spiniger

Ar. peregrinus Ge. truncorum

Chomatophilus sp.

•

Ge. alpinus

Ge. richardi *

•

PI. mediterraneus

m

Ge. pygmaeus S ogona sp.

Ge. carpophagus

• •

•

Ge. electricus

S t. sorrentina

•Eu. pinguis

Ab. allredi

• •

Cl. flavidus

14

BONATO L. et al. , A new elongate geophilid centipede from Sardinia

coxopleura, only slightly denser close to the metasternite, without dorsal pores and without a distinctly

isolated posterior pore (Brolemann 1927, 1930).

Derived morphological features: adaptation to a more strictly endogeic habit?

All three specimens of E. ichnusae gen. et sp. nov. have well-developed gonopods, so that their length

(31-34 mm) can be taken as a good estimate of the average size of the species at full growth. Thus, the

body size of E. ichnusae gen. et sp. nov. is well within the range recorded in the other geophilids: most

species are 1-5 cm long, whereas significantly higher values (up to ca 10 cm) have been recorded only

in rare species of Chomatophilus Pocock, 1896, Gnathoribautia Brolemann, 1909, Maoriella Attems,

1903 and Pleurogeophilus Verhoeff, 1901 (Bonato 2011).

However, E. ichnusae gen. et sp. nov. departs from most other geophilids in some major structural

features: supernumerary and elongated trunk segments, stout legs and body covered with sparse, short

setae.

Trunk segmentation. The number of leg-bearing segments in E. ichnusae gen. et sp. nov. (91-107) is

remarkable in comparison to most other geophilids. Among the ca 560 known species in the Geophilidae

sensu stricto (Bonato 2011), the overall range is 29-125 segments, however figures higher than 90

segments have been recorded in a few species only: Chomatophilus smithi Pocock, 1896 (up to 115

segments; Crabill 1968), Maoriella aucklandica Attems, 1903 (up to 91 segments; Attems 1903),

3,8

3,6

3,4

3,2

-c 3,0

-i —*

~o

5

S 2,8

a>

c

0

« 2,6

=3

~ 2,4

2,2

2,0

1,8

1,6

30 40 50 60 70 80 90 100 110 120

number of leg pairs

Fig. 7. Variation in the elongation of legs and the number of legs in a sample of species of Geophilidae,

including Endogeophilus ichnusae gen. et sp. nov. The length/width ratio of a leg tarsus was measured

at about 20% of the longitudinal series of legs. Full names of species in ‘Material and methods’.

9 Eu. ping uis

Ge. truncorum

Ar. peregrinus

Ge. piae Chomatophilus sp.

•

Cl. fl avid us

Ge. electricus Ge. carpophagus

Ge. alpinus _ , * * *Ab. allredi

_ a 9%Ge.fucorum

Ge. pygmaeus

St. sorrentma

~ . . .. 9 PI. mediterraneus

0Ge. richardi

• 0Sogona sp.

Tu. poseidonis

-

Endoc^philus

qAc. spiniger

^Di. conjungens

15

European Journal of Taxonomy 231 : 1-19 ( 2016 )

Ribautia taeniata Ribaut, 1923 (up to 125 segments; Ribaut 1923), Stenotaenia sturanyi (Attems, 1903)

(up to 115 segments; Bonato & Minelli 2008), Tampiyapylorus Chamberlin, 1912 (125 segments in the

single specimen known; Chamberlin 1912), and Telocricus multipes Chamberlin, 1915 (113 segments

in the single specimen known; Chamberlin 1915).

Elongation of trunk segments. E. ichnusae gen. et sp. nov. departs from most other geophilids also in

the relative elongation of the single leg-bearing segments. By comparing the anterior part of the trunk

in a sample of geophilids (Fig. 6), the metastemites are wider than long in the majority of the species,

whereas they are longer than wide in E. ichnusae gen. et sp. nov. and at least in another species, namely

Geophilus piae Minelli, 1983. The latter is similar to E. ichnusae gen. et sp. nov. in body width but

remarkably different in body length: G. piae has less than 40 pairs of legs and does not surpass 10 mm

in total length ( Min elli 1983).

Elongation of legs. When comparing geophilids with regard to the elongation of the legs (Fig. 7),

E. ichnusae gen. et sp. nov. belongs to a minor group of species with relatively stout legs, contrasting

with a majority of species with more elongate legs. Besides Endogeophilus gen. nov., quite stout legs

are present also in Acanthogeophilus Minelli, 1982 and Diphyonyx Bonato, Zapparoli & Minelli, 2008.

Coverage of setae. The longest setae on the cephalic plate of E. ichnusae gen. et sp. nov. (ca 15 pm long)

are distinctly shorter than those measured in most other species of a sample of geophilids (50-120 pm)

140

120

100

£ 80

O)

c

0)

X

OJ

E 60

ro

<D

U)

40

20

0

200 400 600 800 1000 1200 1400 1600

head width (|jm)

Fig. 8. Variation in the length of setae in a sample of species of Geophilidae, including Endogeophilus

ichnusae gen. et sp. nov. The longest seta on the cephalic plate was measured, and the maximum width

of the cephalic plate has been taken as a proxy for body size. Full names of species in ‘Material and

methods’.

Cl. fl avid us

Ge. pygmaeus

Sogona sp. Chomatophilus sp.

• •

Ge. truncorum pinguis

Tu. poseidonis

Ge. fucorum

Ac. spiniger • • Di - conjungens

Ar. peregrinus • *Ge. electricus

* Jst. sorrentina

Ge. alpmus»» Ge . richardl •

>P/. mediterraneus

Ab. allredi

Ge. carpophagus

Ge. piae

r Endogeophilus

16

BONATO L. et al., A new elongate geophilid centipede from Sardinia

(Fig. 8). Comparably short setae (not surpassing 20 pm) are also found in Geophilus piae , which is

similar to E. ichnusae gen. et sp. nov. in body width but remarkably different in body length (see above).

Geophilomorph centipedes spend most of their life in the soil, inclusive of leaf litter, even though

some species show up on the surface either occasionally or regularly (pers. obs.). Blindness and lack of

pigmentation are common to the entire clade and are usually regarded as adaptations to a predominantly

endogeic lifestyle. The unusual habitus of Endogeophilus gen. nov. (very slender shape, supernumerary

longitudinal articulations, stout legs and short sparse setae) is suggestive of further specialization to

living in the micro-crevices of fine-grain soils. A somehow similar habitus distinguishes strictly endogeic

lineages in distantly related arthropod groups. In particular, among the staphylinid beetles, the endogeic

Leptotyphlinae are remarkably elongate and short-legged, however much smaller in length (less than

2 mm; Coiffait 1959) than Endogeophilus gen. nov. A parallel morphological syndrome is recurrent also

in the aquatic interstitial fauna, e.g., in isopods (Microparasellidae, Microcerberidae), in the harpacticoid

copepods, and even in ciliated protozoans (Delamare Deboutteville 1960). That E. ichnusae gen. et

sp. nov. may have a more strictly endogeic lifestyle than other geophilids is suggested also by the fact

that it has been discovered only recently, when specific sampling techniques for deep-soil fauna has

been employed, while it has escaped detection in all previous surveys, which usually employed manual

sampling on litter and high soil strata (see Zapparoli 2011 for further comments), or were instead specific

for caves (e.g., Zapparoli 2009). No other specimens of E. ichnusae gen. et sp. nov. were found by one

of us (L.B.) when sampling geophilomorphs in the same part of Sardinia (Sulcis-Iglesiente), however

with conventional methods and thus focussing on the higher soil levels.

Acknowledgments

We are grateful to Tuca Fancello (Cagliari, Italy) for allowing us to study the centipedes he collected in

Sardinia and for information on the sampled sites, Douglas Yanega (University of California, Riverside)

and Fabio G. Cupul-Magana (University of Guadalajara, Mexico) for loan of specimens, and Achille

Casale (formerly, University of Sassari, Italy) for suggestions on the soil fauna. The research has been

supported by the Italian Ministry of the University and Scientific Research MIUR (PRIN 20085YJMTC).

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Manuscript received: 1 March 2016

Manuscript accepted: 28 April 2016

Published on: 21 September 2016

Topic editor: Rudy Jocque

Desk editor: Kristiaan Hoedemakers

Printed versions of all papers are also deposited in the libraries of the institutes that are members of the

EJT consortium: Museum national d’Histoire naturelle, Paris, France; Botanic Garden 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; Naturalis Biodiversity Center, Leiden, the Netherlands.

19