European Journal of Taxonomy 222: 1-13

http://dx.doi.org/10.5852/ejt.2016.222

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2016 • Bosselaers J. & Van Keer J.

Research article

urn:lsid:zoobank.org:pub:9135A958-6E83-4E24-878B-388BD206E49B

A redescription of Harpactea dufouri (Thorell, 1873)

(Araneae, Dysderidae), its occurrence outside the Balearic Islands,

and some notes on the corticalis group of the genus

Jan BOSSELAERS 1 * & Johan VAN KEER 2

’Royal Museum for Central Africa, B-3080 Tervuren, Belgium.

2 Bormstraat 204, bus 3, B-1880 Kapellen op den Bos, Belgium.

* Corresponding author: [email protected]

2 Email: [email protected]

1 urn:lsid:zoobank.org:author:D6AD7414-3540-4F06-8631-8873450AA90C

2 urn:lsid:zoobank.org:author:33B65250-686B-44B7-B5E4-6B63203F6F3C

Abstract. Harpactea dufouri (Thorell, 1873) was collected in the Gavarres protected natural area in

Catalonia, Spain. The specimens were compared with specimens from Mallorca, Balearic Islands,

and found to be conspecific. The female of the species is described here for the first time. The new

finding proves that Harpactea dufouri occurs outside the Balearic Islands. The species, however, may

be endemic to Catalonia.

Keywords. Catalonia, corticalis group, biodiversity, endemism, Harpacteinae.

Bosselaers J. & Van Keer J. 2016. A redescription of Harpactea dufouri (Thorell, 1873) (Araneae, Dysderidae), its

occurrence outside the Balearic Islands, and some notes on the corticalis group of the genus. European Journal of

Taxonomy 222: 1-13. http://dx.doi.org/10.5852/ejt.2016.222

Introduction

The genus Harpactea Bristowe, 1939 (type species: Harpactea hombergi (Scopoli, 1763)) is the second

V _

most speciose genus of the family Dysderidae (Rezac 2008). The World Spider Catalog (2016) lists

175 valid and accepted species for the genus, which is almost one third of the 534 species described for

the family. The focus of distribution of the genus is the Mediterranean area, extending to the Caspian

region in the East, and, for some species, to central and northern Europe in the North. Interestingly,

almost all species of Harpactea have a relatively small distribution area. Only H. hombergi (Scopoli,

1763), H. lepida (C.L. Koch, 1838), H. rubicunda (C.L. Koch, 1838) and H. saeva (Herman, 1879) are

widespread species. The majority of Harpactea species have been described after the publication of

Roewer’s first spider catalogue (1942). Indeed, Roewer (1942) lists only 26 species that are presently

considered as valid species belonging to Harpactea. Most of the additional Harpactea species have

since been described by Alicata, who described 12 species (Alicata 1966a, 1966b, 1966c, 1973, 1974),

Brignoli, who described 35 species (Brignoli 1974, 1976, 1977, 1978a, 1978b, 1978c, 1979a, 1979b,

1979c, 1979d, 1980a, 1980b, 1984), Bosmans and coworkers, who described 31 species (Beladjal &

European Journal of Taxonomy 222 : 1-13 ( 2016 )

Bosmans 1997; Bosmans & Beladjal 1989, 1991; Russell-Smith & Bosmans 2011; Van Keer & Bosnians

2009), Dunin, who described 23 species (Dunin 1989, 1991, 1992a, 1992b, 1993), and Dimitrov,

Lazarov and Naumova, who described 11 species (Dimitrov 1997; Dimitrov & Lazarov 1999; Lazarov

2006a, 2006b, 2008a, 2008b, 2009a, 2009b, 2010; Lazarov & Naumova 2010). Among the 172 species

presently known, 111 are known from Europe (including Ukraine) and the eastern Mediterranean

(Turkey, Lebanon and Israel), 26 from the Caucasus and the region around the Caspian sea, and 35

from North Africa (Algeria and Tunisia). Among the European species, 16 have been reported from the

Iberian peninsula (including the Balearic Islands): H. aeruginosa Barrientos, Espuny & Ascaso, 1994,

H. algarvensis Ferrandez, 1990,//. blasi Ribera & Ferraridez, 1986,//. dufouri (Thorell, 1873),//. fageli

Brignoli, 1980, H. gaditana Pesarini, 1988, H. hispana (Simon, 1882), H. hombergi (Scopoli, 1763),

H. magnibulbi Machado & Ferrandez, 1991, H. minoccii Ferrandez, 1982, H. ortegai Ribera & De Mas,

2003, H. proximo Ferrandez, 1990, H. skiakyi Pesarini, 1988, H. serena (Simon, 1907), H. stalitoides

Ribera, 1993 and H. subiasi Ferrandez, 1990. Thirteen of these species are endemic (Cardoso & Morano

2010: 18).

The authors made a one week field trip to the Gavarres protected natural area in Catalonia, a 300 km 2 hill

massif in the northernmost part of the Catalonian coastal range, characterised by an amazing biodiversity

(Bosselaers 2004, Bosselaers in prep.). On this occasion, the junior author collected six specimens of

Harpactea dufouri , formerly only known from the Balearic Islands. This rare species, including the

hitherto unknown female, is redescribed and discussed below.

Material and methods

Specimens were observed, photographed and drawn using Euromex MIC465 and Olympus SZX9 stereo

microscopes. For stereo microscope photographs, a Praktica DC440 digital camera was used. Details

of the male palp and the vulva were studied with a Zeiss Axio Imager A1 microscope equipped with

a Canon Powershot G6 digital camera. The image series obtained were stacked with Zerene Stacker

version 1.04 build T201411272115. The vulva was cleaned for microscopy with trypsin (Sigma) for 24 h

at room temperature an subsequently cleared in methyl salicylate for observation. The male palp was

imm obilized in Schwarzkopf “Freezing Gel Extreme 5”, a transparent vinylpyrrolidone-vinylacetate

copolymer (CAS 25086-89-9) / carbomer gel (Schroder et al. 2000; Schulze zur Wiesche 2006). All

measurements are in millimetres. The format for leg spination follows Platnick & Shadab (1975),

amended for ventral spine pairs according to Bosselaers & Jocque (2000: 307). Leg spination is also

illustrated in a schematic representation (Fig. 1E-G) where pi, do, rl and ve sides of leg articles are

flattened as a folding net (Durer 1525).

Abbreviations

AER

ALS

avg

CJVK

CRB

fi-

ICS

MOQ

= anterior eyes

= anterior eye row

= anterior lateral spinnerets

= average

= personal collection Johan Van Keer

= personal collection Rop Bosmans

= dorsal

= femur

= frontal

= intercoxal sclerites - intercoxal sclerites are six small triangular or elongated sclerites

surrounding the sternum, their tips penetrating between the coxae of the legs - they may be

free, or fused with the sternum (Bosselaers & Jocque 2002: fig. IK)

= median ocular quadrangle

BOSSELAERS J. & VAN KEER J., A redescription of Harpactea dufouri (Thorell, 1873)

PCT =

PER =

PLE =

PLS =

plv =

PME =

PMS =

rlv

Collections

BMNH =

CJVK =

CRB

IZPAN =

MCZ =

MNHN =

MRSN =

MSNG =

NRM =

ZMUC =

metatarsus

number of specimens

patella

pleural bars - pleural bars are narrow, horizontal sclerites between coxae and carapace, one

above each coxa (“pieces epimeriennes” of Simon (1892: 11, fig. 29)) - they may be fused

among each other (Bosselaers & Jocque 2002: fig. IP), with intercoxal sclerites and/or with

carapace

precoxal triangles - precoxal triangles are small triangular sclerites surrounding the sternum,

their tips facing the bases of the coxae (Penniman 1985: 16) - they may be free, or fused

with the sternum (Bosselaers & Jocque 2002: fig. IK)

posterior diverticulum

posterior eyes

posterior eye row

patellar indentation - the patellar indentation is a slit-like membranous indentation on the

rl side of the pa. It may be very narrow or rather wide (Simon 1892: 22; Ledoux & Canard

1991: fig. 15a-b)

prolateral

posterior lateral eyes

posterior lateral spinnerets

prolateral ventral

posterior median eyes

posterior median spinnerets

retrocoxal hymen - the retrocoxal hymen is a weak spot, in most cases hyaline and lens- to

dome-shaped, on the retrolateral face of coxa I (Raven 1998; Bosselaers & Jocque 2002)

retrolateral

retrolateral ventral

standard deviation

tarsus

transversal bar

tibia

ventral

ventral terminal

British Museum of Natural History, London, UK

personal collection Johan Van Keer

personal collection Rop Bosnians

Museum and Institute of Zoology PAS, Warsaw, Poland

Museum of Comparative Zoology, Harvard University, Cambridge, MA, USA

Museum national d’ Histoire naturelle, Paris, France

Museo Regionale di Scienze Naturali, Torino, Italy

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

Naturhistoriska Riksmuseet, Stockholm, Sweden

Zoologisk Museum Kopenhagen, Denmark

European Journal of Taxonomy 222 : 1-13 ( 2016 )

Results

Class Arachnida Cuvier, 1812

Order Araneae Clerck, 1757

Family Dysderidae C.L. Koch, 1837

Genus Harpactea Bristowe, 1939

Harpactea dufouri (Thorell, 1873)

Figs 1-2

Harpactes dufourii Thorell, 1873: 561.

Harpactea dufouri -Denis 1961: 236, fig. 1. —Alicata 1966a: 211, fig. 61. —Brignoli 1979c: 114, figs

2-3. — Le Peru 2011: 268, fig. 400.

Diagnosis

Harpactea dufouri differs from H. corticalis (Simon, 1882) by the more slender bulbus of the male palp,

the more straight conductor and embolus and the larger number of spines on femora III and IV.

Harpactea dufouri somewhat resembles H. he Ee re ns is Bosmans & Beladjal, 1991, but differs from it

by the knife-shaped, pointed conductor with very sharp teeth, the posterior diverticulum of the vulva

consisting of a small ventral and a larger dorsal lobe and by the larger number of spines on femorae III

and IV.

Material examined

SPAIN: 2 33 , 4 §§, Catalonia, Baix Emporda, Mont Ras, alt. 103 m, 41°54'55.1" N, 3°8'25.6" E,

former quarry converted into lake and picnick area, under stones, 23 Oct. 2013, Van Keer leg. (CJVK)

(Fig. 1H); 2 33 , 2 Balearic Islands, Mallorca, Banyalbufar W., Font de Sant Merita, Platanus

forest, alt. 150 m, sifting litter, 1 Apr. 2003, R. Bosmans leg. et det. (CRB).

Description

Male

Largest specimen Mont Ras, total length 5.50 (avg (n = 4) 4.48, sd 1.08). Carapace length 2.16, width

1.60, yellowish brown with faint grey radial striae, more reddish brown in cephalic region, smooth.

Cephalic region slightly wider than half the carapace width, three times as wide as the eye group

(Fig. 2A-B). Fovea weak and thin, length 0.32, anterior end 1.42 from front end of carapace.

MOQ length 0.22, anterior width 0.26, posterior width 0.16, AER width 0.26, PER width 0.34. All six

eyes ringed with black, PE clear pearly white, almost touching, diameter of PLE 1.5 times PME. AE as

large as PLE, pearly white, subquadratic, separated by half their diameter. PER procurved in both do and

fr view. Clypeus vertical, 0.05, equal to half the diameter of AE.

Chilum single, triangular, brown and sclerotised. Chelicerae brown, slightly rugose, anterior rim with

two teeth close to fang tip, the bigger one furthest from tip. Posterior rim with two small, widely spaced

teeth. Labium twice as long as wide, with small anterior notch. Endites long and parallel, with serrula

and bluntly pointed anterior end. Exterior margin notched.

Sternum yellow, rebordered, oval, narrowed frontally and protruding between all coxae. Sternum

length 1.45, width 1.05, surface smooth, but with tiny, evenly spaced punctures. Four pairs of heavily

sclerotised, sharply pointed, dark brown PCT. ICS and PB fused and surrounding coxae.

BOSSELAERS J. & VAN KEER J., A redescription of Harpactea diifonri (Thorell, 1873)

Fig. 1 . Harpactea dufouri (Thorell, 1873), Mont Ras specimens. A. Male palp, prolateral view. B. Tip of

male embolus and conductor, retrolateral view. C. Endogyne, external ventral view, uncleared. D. Cleared

vulva, ventral view. E. Leg spination scheme, legend. F. Leg spination scheme of male. White dots are

spines present on one leg and absent on the other. G. Leg spination scheme of female. White dots are

spines present on one leg and absent on the other. Scale bars: A, C = 0.5 mm; B, D = 100 pm.

European Journal of Taxonomy 222 : 1-13 ( 2016 )

Fig. 2. A-G. Harpactea dufouri (Thorell, 1873), Mont Ras specimens, except F-G. A. dorsal view.

B. dorsal view. C. Tip of male embolus and conductor, retrolateral view, Nomarski interference

contrast. D. Tip of male embolus and conductor, retrolateral view, bright field. E. Cleared vulva, ventral

view. Left: bright field, middle: phase contrast, right: Nomarski interference contrast. F. Tip of male

embolus and conductor, Banyalbufar specimen. G. Endogyne, external ventral view, Banyalbufar

specimen. — H. Map showing the Mont Ras site of Harpactea dufouri (red square), the Gavarres are

delimited by a black line. Scale bars: A-B = 1 mm; C-F =100 pm; G = 0.5 mm ; H = 5 km

BOSSELAERS J. & VAN KEER J., A redescription of Harpactea dufouri (Thorell, 1873)

Legs orange yellow, shiny. Leg formula 1423. No RH, no trochanter notch. PI very narrow, extending

along the whole length of pa EH, but restricted to the basal third in pa I, II and IV. Ventral terminal

preening brush present on mt III and IV. Tarsi with three claws, no claw tufts or tenent hairs. Leg

spination (Fig. IF): fe: I pi 0-0-1; II pi 0—1—1; III do 2-2-2-2; IV do 1—1—1—0; ti: III pi 1-0-1 do 1-1-1

rl 1-1-0 ve 1-1-2; IV pi 1-0-1 do 2-1-2 rl 1-1-1 ve 1-1-2; mt: III do 2-2-1 ve 1-1-2; IV pi 1-1-1

do 2-2-1-2 rl 1-1-0 ve 0-0-1.

Leg measurements:

Total

1.89

1.32

1.84

1.66

0.55

7.26

1.58

1.05

1.45

1.34

0.50

5.92

III

1.53

0.53

1.00

1.26

0.45

4.76

1.97

0.82

1.71

1.92

0.58

7.00

Abdomen grey, covered with thin, pointed grey setae becoming coarser on anterior rim (Fig. 2A). ALS

subcylindrical, stout, separated by A of their length. PMS thin and conical, PLS thin, subcylindrical,

separated by half their length.

Male palp as illustrated (Figs 1A-B, 2C-D, F), with spindle-shaped bulbus, straight, black, simple,

pointed embolus and flat, hyaline conductor with a broadened, sharply toothed tip. Both male specimens

captured in Mont Ras had the right palp missing. The loss of the palp occurred quite some time before

capture, as the scar had completely healed and was sclerotised.

Female

Largest specimen Mont Ras, total length 5.75 (avg (n = 6) 4.70, sd 0.78). Carapace length 2.10, width

1.56, dark yellowish brown with faint grey radial striae, smooth (Fig. 2B). Cephalic region as in male.

Fovea weak and thin, length 0.34, anterior end 1.32 from front end of carapace.

MOQ length 0.21, anterior width 0.25, posterior width 0.18, AER width 0.25, PER width 0.37. Eyes as

in male. Clypeus vertical, 0.03, equal to one third diameter of AE.

Chiluin, chelicerae, labium and endites as in male.

Sternum, PCT, ICS and PB as in male. Sternum length 1.32, width 1.03.

Legs orange yellow, shiny. Leg formula 4123. No RH, no trochanter notch. PI, ventral terminal preening

brush and tarsi as in male. Leg spination (Fig. 1G): fe: I pi 0-0-1; II pi 0-1-1; III do 0-1-2-2; IV do

1—1—1—0; ti: III pi 1-0-1 do 1-1-1 rl l-(0,l)-0 ve 1-1-2; IV pi 1-0-1 do 2-1-2 rl 1-0-1 ve 1-1-2; mt:

III do 2-2-(l,2) ve l-l-(0,2); IV pi 1-1-1 do 2-2-1-2 rl 1-1-0 ve 0-0-1; ta: III ve 0-(0,l)-0.

Leg measurements:

Total

1.58

1.05

1.34

1.18

0.45

5.60

1.39

0.92

1.21

1.05

0.39

4.97

III

1.18

0.50

0.79

1.00

0.34

3.81

1.71

0.79

1.47

1.71

0.55

6.23

Abdomen pinkish grey, covered with thin, pointed grey setae (Fig. 2B). ALS conical, touching. PMS

thin and conical, PLS thin, subcylindrical, separated by half their length.

European Journal of Taxonomy 222 : 1-13 ( 2016 )

Epigyne essentially absent, all sclerotised parts internal (endogyne, Jocque & Dippenaar-Schoeman

2006: 120), but TB (“valva posterodorsale” ofAlicata (1964: 4), “transversal bar” of Chatzaki & Arnedo

(2006: 5)) clearly visible from the outside, as illustrated (Figs 1C, 2G). Vulva with pronounced TB,

PD (Chatzaki & Arnedo 2006: 6) consisting of an irregular ventral lobe and a globular dorsal lobe,

both with a perforated, glandular aspect (Figs ID, 2E). Genital atrium (Alicata 1964: 4; Chatzaki &

Arnedo 2006: 5) connected on both posterolateral ends to glandular structures. Additional membranous,

transparent pouches also present. Spermatheca sausage-shaped, connected to anterior end of genital

atrium (Figs ID, 2E).

Distribution

Only known from the Balearic islands and the Northern part of the Catalonian coastal range (Fig. 1H).

Discussion

The genus Harpactea has a quite homogeneous somatic morphology, but there is considerable genitalic

diversity, suggesting the existence of a number of subgenera, or even the possibility that several related

genera are involved. Simon (1893: 318) was the first to suggest a division of Harpactes Templeton, 1835

(preoccupied as the bird name Harpactes Swainson, 1833 (Trogonidae) and replaced by Harpactea by

Bristowe 1939: 5) in three subgroups. However, only one of these groups concerns species presently

considered as true Harpactea , the other two being composed of species belonging to Dasumia Thorell,

187 5, Harpactocrates Simon, 1914 and Parachtes A1 icata, 1964.

The first to suggest a subdivision of Harpactea as presently delimited is Alicata (1966a: 192). He bases

his subdivision on genitalic characteristics and recognises three large groups: one consisting of species

reminiscent of Dasumia ; another, consisting of two subgroups, having a male bulbus with a complex

set of apophyses and including H. hombergi ; and a third group, characterised by a simple, elongated

bulbus with 1-3 apophyses (embolus, conductor, pseudoconductor (Brignoli 1978b: 481)), consisting

of four subgroups and including H. corticalis (Simon, 1882). Brignoli (1978b: 481) recognises two

groups: a corticalis group with a simple set of apophyses on the bulbus (consisting of six subgroups),

and a hombergi group with complex apophyses (consisting of four subgroups). Deeleman-Reinhold

(1993: 130) recognises four groups, based on a combination of genitalic and leg spination characters: a

corticalis group, largely similar to Alicata’s and Brignoli’s group with a simple male bulbus, a hombergi

group characterised by a complex male bulbus, a lepida group with a lamellar conductor and a wide PD,

and a rubicunda group with a large, entirely membranous PD. Beladjal & Bosmans (1997: 24) follow

the subdivision of Brignoli (1978b), but add a third group, the auriga group, characterised by a globular

bulbus and a long, filiform embolus.

None of these subdivisions is entirely satisfactory, as they are purely phenetic and no cladistic analysis

was performed. Although the hypothesis sounds plausible, there is no proof that the corticalis group

is “le groupe le plus primitif’ (Beladjal & Bosmans 1997: 24), as no phylogenetically valid outgroup

comparison (Watrous & Wheeler 1981; Maddison et al. 1984) is available. There is even no solid proof

that this group actually exists, id est , that it is not polyphyletic.

Nevertheless, provisory as they are, the subdivisions proposed to date have some practical advantage,

facilitating an overview of the genitalic diversity in the large genus Harpactea. Moreover, it has to be

admitted that the corticalis group is indeed “abbastanza omogeneo” and “abbastanza ben delimitato

dagli altri” (Brignoli 1978b: 481), so it can be retained for the time being, until the enormous work of a

cladistic analysis of Harpactea has been undertaken.

Scrutiny of the available literature suggests that such a corticalis group should at least include H. acuta

Beladjal & Bosmans, 1997, H. angustata (Fucas, 1846), H. arguta (Simon, 1907), H. blasi Ribera &

BOSSELAERS J. & VAN KEER J., A redescription of Harpactea dufouri (Thorell, 1873)

Ferrandez, H. carusoi Alicata, 1974,//. corticalis { Simon, 1882),//. dufouri { Thorell, 1873),//. dumonti

Bosnians &Beladjal, 1991,//. gaditana Pesarini, 1988, H. gennargentu Wunderlich, 1995,//. gridellii

(Caporiacco, 1951), H. heizerensis Bosnians & Beladjal, 1991, H. innupta Beladjal & Bosnians, 1997,

H. kalaensis Beladjal & Bosnians, 1997, H. kareli Bosnians & Beladjal, 1991, H. major (Simon, 1911),

H. mertensi Bosmans & Beladjal, 1991, H. minuta Alicata, 1974, H. mitidjae Bosmans & Beladjal,

1991, H. monicae Bosmans & Beladjal, 1991, /A muscicola (Simon, 1882), H. oranensis Bosmans &

Beladjal, 1991, H. ouarsenensis Bosmans & Beladjal, 1991, H. piligera (Thorell, 1875), H. punica

Alicata, 1974,//. senalbensis Beladjal & Bosmans, 1997, H. sicula Alicata, 1966,//. sinuata Beladjal &

Bosmans, 1997, H. undosa Beladjal & Bosmans, 1997, H. yakourensis Beladjal & Bosmans, 1997,

and H. zannonensis Alicata, 1966. These 31 species (one fifth of the genus) all occur in the western

Mediterranean and they all have small distribution areas.

Three of the 16 Iberic Harpactea species can be attributed to the corticalis group with some confidence:

H. blasi, H. dufouri and H. gaditana.

The specimens collected in Mont Ras were at first considered to be a new species by the authors. Indeed,

the illustrations of the male palp of H. dufouri in Denis (1961), Alicata (1966a, drawn after Denis) and

Le Peru (2011, a synthetic computer drawing) are insufficient to identify the species with certainty.

Nevertheless, Brignoli (1979c: fig. 2) illustrates a toothed conductor that is quite similar to the one of

the Mont Ras specimens (Fig. IB, 2C-D), as rightly pointed out by Crespo when refereeing a previous

version of this paper. Unfortunately, the type specimen of H. dufouri is not present in NRM, where it

would normally be kept, given the fact that it was collected by the Swede F. Soderlund (Thorell 1873:

561). The type could also not be traced in BMNH, IZPAN, MCZ, MNHN, MRSN, MSNG or ZMUC.

As a result, the type of H. dufouri must be considered lost.

However, four adult specimens of H. dufouri collected by Bosmans in Mallorca could be studied.

They turned out to be conspecific with the Mont Ras material (Figs 1B-C, 2F-G). It can be concluded

that H. dufouri is not an island endemic, but also occurs in the Catalonian coastal region. The female,

described here for the first time, has a genitalic morphology typical for the corticalis group of Harpactea.

Acknowledgements

The authors are grateful to Monique Luyten for hospitality and daily care and to Joan Botey i Serra

for encouragement and the permission to collect on his private property in the Gavarres. Luis Carlos

Crespo and an anonymous referee are thanked for their interesting comments that helped improve the

manuscript. Many thanks are also due to the collection curators and arachnologists that were helpful

in trying to locate the holotype of Harpactea dufouri : Gonzalo Giribet, Laura Leibensperger (MCZ);

Pietro Passerin d’Entreves, Elena Gavetti (MRSN); Maria Tavano (MSNG); Christine Rollard (MNHN);

Jerzy Proszynski, Wioletta Wawer (IZPAN); Nikolaj Scharff (ZMUC); Janet Beccaloni (BMNH); Kjell

Arne Johanson, Torbjorn Kronestedt, Karin Sindemark Kronestedt (NRM). Rop Bosmans is gratefully

acknowledged for the specimens of Harpactea dufouri from Mallorca.

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World Spider Catalog 2016. World Spider Catalog. Natural History Museum, Bern. Available from

http://wsc.nmbe.ch. version 17.0 [accessed on 11 Mar. 2016]

Manuscript received: 4 January 2015

Manuscript accepted: 24 March 2016

Published on: 16 August 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.