European Journal of Taxonomy 210: 1-21

http://dx.doi.org/10.5852/ejt.2016.210

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2016 • Pedroso D.R. et al.

Research article

urn:lsid:zoobank.org:pub:E239C584-B41B-4AC7-89El-E3D8A2EB921A

Redescription and synonymies of Diplura macrura (C.L. Koch, 1841)

and D. lineata (Lucas, 1857), with notes on the genus

(Araneae, Dipluridae)

Denis Rafael PEDROSO 1 *, Pedro DE SOUZA CASTANHEIRA 2 &

Renner Luiz CERQUEIRA BAPTISTA 3

'Museu Nacional, Universidade do Brasil (UFRJ). Quinta da Boa Vista, Sao Cristovao, 20.940-040,

Rio de Janeiro, RJ, Brazil.

2 ’ 3 Instituto de Biologia, Universidade do Brasil (UFRJ). Avenida Brigadeiro Trompowski s/n,

Ilha do Fundao, 21944-970, Rio de Janeiro, RJ, Brazil.

* Corresponding author: [email protected]

2 E-mail: [email protected]

3 E-mail: baptistr@gm ail. com

1 um:lsid:zoobank.org:author:E67DAD82-2E69-4E76-B4F0-7BA09E13A68F

2 urn:lsid:zoobank.org:author:BFEC9541-C4F5-4FBF-90AB-AE4885A943D5

3 urn:lsid:zoobank.org:author:CF514365-072A-4248-AF23-C112FC360305

Abstract. Diplura C.F. Koch, 1850 is a mygalomorph genus with putative records from Central and

South America. The type-species Diplura macrura (C.F. Koch, 1841), originally described from West

Indies, is poorly known and represented only by its holotype. Most of the 20 species currently included

in the genus lack modern taxonomic descriptions, as D. lineata (Fucas, 1857), from Rio de Janeiro

state, Brazil. Males and females of Diplura macrura and D. lineata are herein redescribed. New junior

synonyms of D. macrura are identified ( Linothele bicolor (Simon, 1889), Diplura uniformis Mello-

Feitao, 1923, and the two junior synonyms of the latter species, Thalerothele minensis Mello-Feitao,

1926 and T. aurantiaca Mello-Feitao, 1943). Also, two junior synonyms are established for D. lineata :

Diplura fasciata (Bertkau, 1880) and Diplura nigridorsi (Mello-Feitao, 1924). The type-locality of

D. macrura is corrected to Sao Joao del Rei, Minas Gerais state, Brazil. D. macrura is restricted to the

state of Minas Gerais and D. lineata to the state of Rio de Janeiro. The type-locality of D. parallela

(Mello-Feitao, 1923) is also corrected from Argentina to Parana state, Brazil. The distribution of Diplura

is now restricted from south Panama to north Argentina, excluding previous erroneous records for Cuba

and West Indies. The six synonymies herein established help to clarify the genus composition, which

includes now 17 valid species.

Keywords. Atlantic Forest, biodiversity, Brazil, Diplurinae, Mygalomorphae.

Pedroso D.R., Castanheira P. de Souza & Baptista R.L.C. 2016. Redescription and synonymies of Diplura macrura

(C.L. Koch, 1841) and D. lineata (Lucas, 1857), with notes on the genus (Araneae, Dipluridae). European Journal

of Taxonomy 210: 1-21. http://dx.doi.org/10.5852/ejt.2016.21Q

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European Journal of Taxonomy 210 : 1-21 ( 2016 )

Introduction

Diplura C.L. Koch, 1850 is a Neotropical mygalomorph genus, including twenty species of variable

size and color pattern. Nowadays, its accepted distribution goes from Cuba to Argentina (World Spider

Catalog 2015). Most species were described from southeastern and southern Brazil, with species

recorded also from the following countries: Bolivia, Ecuador, Paraguay and Venezuela.

Raven (1985) included Diplura in the subfamily Diplurinae, alongside Trechona C.L. Koch, 1850 and

Linothele Karsch, 1879. The traditional diagnostic character fox Diplura is the simple lyra, at the internal

side of the maxilla (coxa) of the palp, formed by a single row of clavate setae (Raven 1985). In contrast,

Linothele has no lyra and Trechona has a complex lyra, with multiple rows of numerous setae, arranged

in a black shield. However, Harmonicon F.O. Pickard-Cambridge, 1896, included in Diplurinae by

Marechal & Marty (1998), also has a simple lyra. In the revalidation of Harmonicon , Marechal &

Marty (1998) proposed a separation from Diplura based on the shape and number of setae on lyra. In

Harmonicon , the lyra is formed by only 5 setae, with a flattened and curved tip, whereas in Diplura it

has more setae and a different tip. However, Pedroso & Baptista (2014) pointed out that the number

and shape of the setae are variable in Diplura , where some species may have just a few setae (down

to 2) and setae tip curved and sometimes a bit flattened. The number of setae is not a reliable character

for Harmonicon , as already suggested by Drolshagen & Backstam (2011). Nevertheless, the strongly

curved and flattened setal tip is found only in Harmonicon (Pedroso & Baptista 2014). Several other

characters have previously been used to separate the genera included in Diplurinae, as the tarsal scopulae

and segmentation (Raven 1985) and leg formulae and size (Marechal & Marty 1998). A preliminary

discussion on the diagnostic characters used in Diplurinae is found in Pedroso & Baptista (2014), where

several traits are dealt with. For example, the short and thickened palp tibia of males in Diplura (see

Figs 5 and 17 in the plates below) strongly contrasts with the long and thin tibia in other Diplurinae,

as Trechona (Pedroso et al. 2008: fig. 5), Harmonicon (Pedroso & Baptista 2014: fig. 7) and Linothele

(Duperre & Tapia 2015: fig. 28). On the other hand, the rigid setae near lyra found in most Harmonicon

species (Pedroso & Baptista 2014: fig. 6) is diagnostic at least to a large subset of the genus.

In this paper, the composition and distribution of Diplura are discussed. The type-species Diplura

macrura (C.L. Koch, 1841) is redescribed, based on the holotype and recently collected specimens.

Also, the female is newly described and new synonymies are presented. The type-locality of the species

is corrected to Minas Gerais state, Brazil, implying in the exclusion of Cuba and West Indies from its

geographical range. Additionally, Diplura lineata (Lucas, 1857) is redescribed, two synonymies are

given and the species distribution is widened. The type-locality of D. parallela (Mello-Leitao, 1923) is

corrected from Argentina to Brazil.

Material and methods

The description of color pattern is based on specimens preserved in 75% ethanol. The female genitalia

was cut off and clarified with clove oil. Habitat information and photos of living animals were added,

when available. Observations, photographs and measurements were made with aLeicaDFC295 camera,

attached to a Leica M205C stereo microscope. Samples for scanning electron were coated with gold-

palladium and observed under high vacuum on a JEOL JSM-6510 microscope. All photos were edited in

the software Photoshop CS5 and plates were mounted in the software CorelDraw X7. Measurements are

given in millimeters, unless otherwise noted. Carapace length was measured from anterior margin of the

clypeus to the posterior border. Total length was measured from the anterior margin of the clypeus to the

posterior border of the anal tubercle, not including the spinnerets. Each article of the pedipalp and legs

was measured in retrolateral view, from the basal condylus to the distal one. Geographical coordinates

for localities were obtained from Geonames (2015). The distribution map (Fig. 26) was elaborated using

ESRIARCGIS 10 software.

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PEDROSO D.R. et al.. Redescription of Diplura macrura and D. lineata

Abbreviations

Institutions

IBSP = Institute Butantan, Sao Paulo, Brazil

MfN = Museum fur Naturkunde, Berlin, Germany (formerly Zoologisch Museum fur Naturkunde

der Humboldt-Universitat)

Mello-Leitao’s Private Collection, now at Museu Nacional, Universidade do Brasil/

Universidade Federal do Rio de Janeiro, Brazil

Museum National d'Histoire Naturelle, Paris, France

Museu Nacional, Universidade do Brasil/Universidade Federal do Rio de Janeiro, Brazil

Museu de Zoologia da Universidade de Sao Paulo, Brazil

MFPC =

MNHN =

MNRJ =

MZSP =

Structures

AFE = anterior lateral eyes

AME = anterior median eyes

ITC = inferior (or unpaired) tarsal claws

PEE = posterior lateral eyes

PLS = posterior lateral spinnerets

PME = posterior median eyes

PMS = posterior median spinnerets

STC = superior (or paired) tarsal claws

Spines (or macroseta)

ap = apical

p = prolateral

pld = prolaterodorsal

plv = prolateroventral

r = retrolateral

rid = retrolaterodorsal

rlv = retrolateroventral

v = ventral

Results

Class Arachnida Cuvier, 1812

Order Araneae Clerclc, 1757

Family Dipluridae Simon, 1889

Genus Diplura C.L. Koch, 1850

Diplura macrura (C.L. Koch, 1841)

Figs 1-13, 26

Mygale macrura C.L. Koch, 1841: 38, fig. 715 (f).

Diplura bicolor Simon, 1889: 215 ($) syn. nov.

Thalerothele uniformis Mello-Leitao, 1923: 105, fig. 4 (f) syn. nov.

Thalerothele minensis Mello-Leitao, 1926: 314, figs 1-3 (f) syn. nov.

Thalerothele aurantiaca Mello-Leitao, 1943: 255 (§) syn. nov.

Diplura bicolor - Mello-Leitao 1937: 3, fig. 3 (f). —Biicherl 1957: 385, figs 12, 12a.

Diplura macrura-CD. Koch 1850: 75. —Raven 1985: 74, figs 18-23.

Thalerothele uniformis - Biicherl, Timotheo & Lucas 1971: 128, figs 12-14 (syn. T. minensis,

T. aurantiaca).

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European Journal of Taxonomy 210: 1-21 (2016)

Diplura uniformis - Platnick 1993: 89 (transfer). — Silva-Moreira et al. 2010: 32.

Linothele bicolor - Platnick 1998: 120 (transfer).

Diagnosis

Both sexes of this species have a strongly contrasting color pattern, with a reddish brown carapace

and a dark brown abdomen, shared only with Diplura paraguayensis (Gerschman & Schiapelli, 1942).

In mature D. macrura , the abdomen is uniformly dark brown, without spots or any visible markings.

However, according to its original description (Gerschman & Schiapelli 1942), D. paraguayensis has

a blackish brown reticulate on abdomen dorsum. Moreover, the lyra of D. paraguayensis has 13 setae

(Gerschman & Schiapelli 1942: pi. ix; Schiapelli & Gerschman 1968, fig. 7), in contrast with the 7-9

setae in D. macrura. The bulb of D. macrura (Figs 6-7) has an embolic base much larger than in

D. paraguayensis (Schiapelli & Gerschman 1968: figs 14-15). Also, the embolus is about 2* longer

than the bulb in D. macrura and 3x longer in D. paraguayensis. The spermathecae of D. macrura has

a longer and thinner stalk (Fig. 13) than the females of D. paraguayensis from Argentina (Goloboff

1982: 1). Also, the distal lobes are larger and spread over the distal third of the stalk in D. macrura , while

they are smaller and concentrated in the apex in D. paraguayensis.

Type material

Mygale macrura: BRAZIL: S, holotype, Minas Gerais, Sao Joao del Rei, [1830], Sellow, Friedrich W.

(MIN 2083), examined.

Diplura bicolor: BRAZIL: 2 §§, syntypes, Minas Gerais, Catas Altas, Cara 9 a (coll. E. Simon, MNHN

AR 4932 B337), examined (photos).

Thalerothele uniformis: BRAZIL: holotype, Sao Paulo, E. of Garbe (MZSP?, MNRJ?), not located.

Thalerothele minensis: BRAZIL: S, holotype, Minas Gerais, Ouro Preto, Magalhaes Gomes (Mello-

Leitao collection 880, MNRJ 1360), not located.

Thalerothele aurantiaca: BRAZIL: §, holotype, Minas Gerais, Ouro Preto/Mariana: Itacolomi,

O. Leonardos (MNRJ 53945), not located.

Material examined

BRAZIL: 1 S, Minas Gerais, Parque Nacional Serra do Cipo, R. Bertani, R. Martins, C.S. Fukushima &

M.P Pavani leg. (MNRJ); 1 §, 2 juvs, Caete, 11—19 Feb. 1961, P. Vanzolini, H. Britski & N. Menezes

Fig. 1. Original (right) and added (left) labels from the holotype of Diplura macrura (C.L. Koch, 1841).

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PEDROSO D.R. et al.. Redescription of Diplura macrura and D. lineata

Figs 2-13. Diplura macrura (C.L. Koch, 1841). 2. §, habitus dorsal. — 3-9. left leg I. 3. Tibial

spur, retrolateral. 4. Tibial spur and metatarsal clasper, ventral. 5. Left palp, retrolateral. 6-9. Left bulb.

6. Retrolateral. 7. Prolateral. 8. Ventral. 9. Dorsal. — 10-13. §. 10. Maxilla with lyra. 11. Lyra detail,

SEM. 12. Maxilla with lyra, SEM. 13. Vulva, dorsal. Scale = 1 mm, unless otherwise noted.

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European Journal of Taxonomy 210: 1-21 (2016)

leg. (MZSP 6576); 1 S, Catas Altas, Reserva Particular de Patrimonio Natural Serra do Cara 9 a, 24 Apr-

1 May 2002, Equipe Biota (IBSP13148); 2 1 juv., Lima Duarte, Parque Estadual de Ibitipoca, 21 Mar.

1997, A. Oliveira & B. Souza leg. (IBSP 8390); 1 S, Parque Estadual de Ibitipoca, Apr. 1997, A. Oliveira

& B. Souza leg. (IBSP 8432); 1 §, juv., Ouro Preto, Esta 9 ao Ecologica de Tripui, R. Bertani, R. Martins,

C.S. Fukushima & M.P Pavani leg. (MNRJ4467 and MNRJ 4490, respectively); 1 $, Esta 9 ao Ecologica

de Tripui, 1220 m, 20°22'92.7" S, 43°33'29.3" W, R. Bertani, R. Martins, C.S. Fukushima & M.P. Pavani

leg. (MNRJ 4502); 1 $, Esta 9 ao Ecologica de Tripui, 1236 m, 20°22'93.2" S, 43°33'28.7" W, R. Bertani,

R. Martins, C.S. Fukushima & M.P. Pavani leg. (MNRJ 4496); 1 ? (MNRJ 4487), 1 $ (MNRJ 4497),

1 $ (MNRJ), Esta 9 ao Ecologica de Tripui, 1246 m, 20°22'93.2" S, 43°33'28.7" W, R. Bertani, R. Martins,

C.S. Fukushima & M.P. Pavani leg.; 1 $, Esta 9 ao Ecologica de Tripui, 20°22'6.62" S, 43°32'5.74" W,

R. Bertani, R. Martins, C.S. Fukushima & M.P. Pavani leg. (MNRJ 4509); 1 juv., Esta 9 ao Ecologica

de Tripui, 20°22'9.70" S, 43°33'1.04" W, R. Bertani, R. Martins, C.S. Fukushima & M.P. Pavani leg.

(MNRJ 4498); 1 $, Esta 9 ao Ecologica de Tripui, 20°22'915" S, 43°33'227" W, R. Bertani, R. Martins,

C.S. Fukushima & M.P. Pavani leg. (MNRJ 4489); 1 Esta 9 ao Ecologica de Tripui, 20°22'93.2" S,

43°33'26.7" W, R. Bertani, R. Martins, C.S. Fukushima & M.P. Pavani leg. (MNRJ 4505); 1 Esta 9 ao

Ecologica de Tripui, 20°22'93.2" S, 43°33'28.7" W, R. Bertani, R. Martins, C.S. Fukushima & M.P. Pavani

leg. (MNRJ 4479); 1 juv., Esta 9 ao Ecologica de Tripui, 20°22'93.2" S, 43°33'28.7" W, R. Bertani, R.

Martins, C.S. Fukushima & M.P. Pavani leg. (MNRJ); 1 juv., Esta 9 ao Ecologica de Tripui, 20°22'970" S,

43°33T04" W, R. Bertani, R. Martins, C.S. Fukushima & M.P. Pavani leg. (MNRJ 4492); 1 c?, Esta 9 ao

Ecologica de Tripui, 20°22'92,2" S, 43°33T3" W, 8 Jan. 2006, R. Bertani, R. Martins, C.S. Fukushima

& M.P. Pavani leg. (MNRJ ); 1 ?, Esta 9 ao Ecologica de Tripui, 20°22'93.2" S, 43°33'287" W, 7 Jan.

2005, R. Bertani, R. Martins, C.S. Fukushima & M.P. Pavani M. leg. (MNRJ 4333); Ouro Preto: 1 juv.,

Parque Estadual do Itacolomi, R. Bertani, R. Martins, C.S. Fukushima & M.P. Pavani leg. (MNRJ);

1 juv., Parque Estadual do Itacolomi, R. Bertani, R. Martins, C.S. Fukushima & M.P. Pavani leg. (MNRJ

4515); 1 S, Parque Estadual do Itacolomi, 1,306 m, 20°22'662" S, 43°32'574" W, R. Bertani, R. Martins,

C.S. Fukushima & M.P. Pavani leg. (MNRJ 4495); 1 $, Parque Estadual do Itacolomi, 20°25'638" S,

43°30'399" W, R. Bertani, R. Martins, C.S. Fukushima & M.P. Pavani leg. (MNRJ 4507).

Description

Male (MNRJ 4496) (Figs 3-9)

Carapace: 6.4 long, 5.0 wide. Abdomen: 6.6 long. Spinnerets: PMS 1.3 long; PLS, total length 6.8,

basal article 2.0, middle 2.2, distal 2.6, respectively. Legs: see Table 1. Carapace: length/width 1.3; flat,

cephalic area slightly raised, thoracic furrows shallow and wide. Fovea: short, deep, recurved. Carapace

covered with short, thin setae, interspersed with some longer and thicker setae; border with abundant

long and thick setae pointing outwards, increasing in number towards posterior angles. Clypeus almost

totally hidden by the bulging eye tubercle, frontal margin bearing 5 thick, long, erect setae. Eye tubercle:

0.6 long, 1.0 wide, area between posterior eyes covered with thin setae and bearing 4 thicker, longer

setae. AME 0.3, almost spherical, but a bit longer than wide, set apart by 0.7x their diameter. ALE

elliptical, much longer than wide, its length about 0.6x the AME diameter. PME small, with flattened

lens, longer than wide, its length about 0.5x AME diameter. PLE elliptical, much longer than wide,

its length a bit less than 0.7x AME diameter. PME and PLE clearly set apart by around 0.4x the PME

length. Anterior eye row slightly recurved, posterior eye row recurved. Eye rows with similar width.

Chelicerae: promargin with 11 teeth on left and 9 on right chelicera. Plectrum with 5 thick, long setae.

Labium: length/width 0.8, no cuspules. Labio-stemal groove deep with elongated sigilla. Sternum:

about 45% longer than wide, very similar to the sternum of D. lineata. Posterior angle in a blunt point,

not separating coxae IV. Sigilla: three pairs, spherical, with a subtle increase in size from anterior to

posterior, all near margin. Palp (Fig. 5): relatively short, without retrolateral spines, femur: d 1-2-0,

pl0-0-l, tibia pi 0-1-0, v 1-2-0. Tibia: length 2.3, width 0.9, short, incrassated, thinner at the basis and

apex. Maxillae: length/width 1.5. Cuspules: 13 spread over ventral inner heel. Lyra at the ventral side

of the maxilla, formed by 8 modified thick, long setae, increasing in size from basis to apex of the lyra.

6

PEDROSO D.R. et al ., Redescription of Diplura macrura and D. lineata

Table 1. Diplura macrura (C.L. Koch, 1841), c^- Length of left leg articles (dorsal view).

Leg I

Leg II

Leg III

Leg IV

Fe

5.5

5.1

4.7

5.9

Pa

3.2

2.9

2.4

2.5

Ti

4.2

3.8

3.2

4.4

Mt

4.0

4.1

4.4

6.2

Ta

3.3

3.2

3.1

3.6

Total

20.2

19.1

17.8

22.6

weakly curved at apical portion. Legs: Leg formula 4123. Legs covered with short, thin, horizontal black

setae and with some longer, thicker, erect black setae. All tarsi with thin scopula, throughout the length

of the article, divided by two series of thicker setae at the middle line of the ventral face. Metatarsus

I with undivided thin scopula, covering the distal half of the retrolateral side. Metatarsus II similar to

I, but with scopula on both sides. Metatarsi III-IV without scopula. All tarsi provided with numerous

small cracks covering almost all the ventral and lateral faces, except by the basis and tip of the article.

Leg I (Pigs 3^1): tibia I with a relatively short distal retrolateral spur, curved and blunt, placed at its

ventral comer. Megaspine pointed, slightly sinuous, especially on the apex, almost 2x longer than the

spur. Metatarsus I with a distinct retrolateral tubercle placed ventrally at the beginning of the median

third, conical, pointed and facing towards the tip of article. Ventrally, there is one spine near the apex and

another one on the median third, placed much ahead of the tubercle. Lringe formed by many thick and

long spiniform setae and spines (clasper) covering almost all prolateral side of the metatarsus (Fig. 4).

Spines: leg I: femur d 1-2-0, pld0-0-l, rld0-0-l, patella p0-0-l left, pO right, rO left, 0-1-0 right, tibia

p0-2-0, vO-1-1 ap (apophysis), metatarsus p0-2-0, vO-1-1 ap; leg II: femur d 1-2-0, pld 1-0-1 left,

pld0-0-l right, rld0-0-l left, rldO right, patella p0-l-l left, p0-0-l right, tibia p0-2-0, vl-l-2ap left,

vl-0-2ap right, metatarsus p 1—1—1 left, p 1—1—0 right, vl-2-2ap left, vl-2-lap; leg III: femur dl-2-0,

pldO left, pld0-l-l right, rld0-l-l left, rldO-2-1 right, patella p0-l-l left, p0-l-l right, rO-l-O left,

rl-1-0 right, tibia p0-2-0 left, pO-2-1 right, r 1—1—1, vl-l-2ap; metatarsus p2-2-l left, p0^1-l right,

rl-1-1 left, r 1—2—1 right, v0-4-3ap left, v0-3-3ap right; leg IV: femur d2-l-0 left, dl-2-0 right, rldO-

0-1 left, rld0-0-2 right, patella r0-l-0, tibia pO-l-O left, p0-2-0 right, rl-2-1 left, rl-1-1 right, v2-

l-2ap left, vl-0-2ap right, metatarsus dl-0-0 left, dl-1-0 right, p 1—2—1, rl-2-1, vl-5-3ap. Claws:

ITC without teeth. Teeth at STC: all claws with a small spur at the basis in both sides leg I : inner 6-7

and outer rows 5-6; leg II : inner row 9, outer row 8-9; leg III : inner row 6-7, outer row 6; leg IV : inner

and outer rows 6-7. Bulb (Figs 6-9): piriform and conical in a retrolateral view, with length about 3 A of

its width. Embolus about 2x longer than the bulb itself, with a wide base in relation to the globose part

of the bulb and tapering towards the apex. Embolus clearly curved since its base and tapering from the

basal third to the apex, clearly seen both on retrolateral and prolateral views. However, in dorsal view,

the embolus is just slanted, almost straight. On ventral view, spermatic duct large at the base, tapering

regularly towards the tip, but much thinner from the distal third on.

Female (MNRJ 4487) (Figs 2, 10-13)

Total length 15.3. Carapace: 7.3 long, 5.8 wide. Abdomen: 8.0 long, 5.0 wide. Spinnerets: PMS 1.4 long;

PLS, total length 5.9, basal article 2.3, middle 1.6, distal 2.0, respectively. Legs: see Table 2. Females

are very similar to males except by its bigger size and the following characteristics: carapace length/

7

European .Journal of Taxonomy 210: 1-21 (2016)

Table 2. Diplura macrura (C.L. Koch, 1841), $. Length of left leg articles (dorsal view).

Leg I

Leg II

Leg III

Leg IV

Fe

5.6

4.8

4.4

5.2

Pa

3.4

3.0

2.6

2.9

Ti

3.6

3.3

3.0

4.2

Mt

4.0

3.5

4.2

5.6

Ta

3.0

2.8

2.8

3.0

Total

19.6

17.4

17.0

20.9

width 1.3. Clypeus very narrow, with the frontal margin carrying five thick, long setae, facing forward.

Eye tubercle with a thick seta on its anterior margin. Area between eyes with two longer and thicker

setae. AME separated by around their diameter. PME and PLE less separated. Chelicera with 10-11

promarginal teeth, on the right and left chelicera, respectively. Plectrum with 6 thick, long setae. Maxilla

with 16 (left) or 19 (right) cuspules. Lyra (Figs 10-11) as in male, formed by 7 modified setae. Tarsi

I—II with scopula similar to male, but tarsus III with a very thin scopula and tarsus IV without scopula.

Metatarsi I—II with undivided thin scopula, covering both sides and the ventral face. All tarsi provided

with few small cracks covering only the median area of the ventral and lateral faces. Spines: leg I:

femur d 1-2-0, pld0-0-l; patella 0; tibia p0-0-l, v0-0-2ap (apophysis); metatarsus v0-3-2ap; leg II:

femur dl-1-0, pld0-0-l; patella p0-0-l; tibia p0-0-l, v0-0-2ap; metatarsus v0-3-2ap; leg IIP femur

d 1-0-0, rld0-0-2 left, rld0-0-l right; patella p0-l-l left, r0-l-0; tibia p 1-1-0 left, p0-2-0 right,

r0-2-0, v0-0-lap left, v0-0-2ap right; metatarsus p 1-2-1 left, pl-3-1 right, rl-1-1, v0^1-3ap; leg IV:

femur dl-1-0 left, dl-2-0 right, rld0-0-l; patella r0-l-0; tibia p 1—1—0, r2-l-0, v0-0-lap left, v-0-

0-2ap right; metatarsus pO-2-1 left, pl-2-1 right, r2-l-2 left, rl-1-2 right, vl-3-2ap. Spermathecae

(Fig. 13): separated by about 70% of its length, with a thick stem, keeping a similar width up to the distal

lobules. Stem curved forward from the median part on, bearing three large apical lobules of variable

sizes.

Color pattern

Both sexes with carapace reddish brown, covered by abundant setae of a weakly coppery hue. Thoracic

furrows and cephalic area darker. Eye area black. Legs, palps, chelicera and spinnerets dark brown, with

a clear contrast to the color of carapace. Labium, sternum and leg coxae orange, with darker sigilla.

Dorsum of the abdomen uniformly dark brown without markings or stripes, venter usually lighter

colored than dorsum, but with a dark hue in some males.

Variation

Chelicera with 8-11 promarginal teeth. Lyra at the ventral side of the maxilla, formed by 7 to 9 modified

setae (Figs 10-12). The maxillary cuspules may vary from 13 to 17 in males and from 16 to 19 in females.

Synonymy and notes

When describing Mygale macrura , C.L. Koch (1841: 39) indicated that the type-locality of the species

was “Westlndien, St. Juan”. Nine years later, he transferred the species to his new genus Diplura ,

without mentioning the type-locality (C.L. Koch 1850: 75). The type-locality indicated by Koch 1841

was cited differently by subsequent authors, for example, “Westindien” (Ausserer 1871: 178), “St. Jean,

8

PEDROSO D.R. et al, Redescription of Diplura macrura and D. lineata

Antilles” (Simon 1903: 963), “West Indies” (Petrunkevitch 1911: 60), or “Saint John, U.S. Virgin

Islands” (Muchmore 1993: 32). All those subsequent authors did not mention any additional specimen

of D. macrura and were certainly restating the information given by C.L. Koch (1841).

Banks (1909: 155) was the first arachnologist to record D. macrura in Cuba, from Pinar Del Rio, as

“Ischnothele macrura Koch”, but he made no comments on the implicit genus transfer, nor did he

include any description of the specimens he examined. Lutz (1915: 77) and Petrunkevitch (1926: 27)

also pointed out that D. macrura had previously been recorded from Cuba. While Lutz expressly cited

Banks (1909), Petrunkevitch did not mention its source. However, he was probably also citing Banks

(1909), as he stated that “ Diplura macrura C. Koch, has been also reported from Cuba”. In his paper on

endemic spiders from Cuba, Alayon (2000: 38) also included D. macrura without any further comment.

Currently, the only record for D. macrura included in the World Spider Catalog (2015) is Cuba, without

any mention of Koch's type-locality.

In her paper on Cuban spiders though, Bryant (1940: 260) considered Banks’ record a misidentification

of Ischnothele longicauda Lranganillo, 1936. She based her opinion on the relative abundance of the

latter species in Cuba. The family Dipluridae is scarcely represented in Cuba, with only I. longicauda

and two species of Masteria L. Koch, 1873 recorded for the country (World Spider Catalog 2015), not

considering the erroneous D. macrura citation by Banks. Therefore, the only specimen of D. macrura

undoubtedly cited for Cuba, West Indies or any other Caribbean locality is the holotype.

Surprisingly, an analysis of the holotype of D. macrura indicates that the type-locality was misinterpreted

by Koch. The holotype label clearly indicates “Koch *. typ. S. Joao d. Rey. Sello” (Tig. 1, right). On

the other hand, there is an additional label (Tig. 1, left) inside the vial that contains the type indicating

“Macrura N. Koch. * fg. 715. Cameta Sieber”, referring to Cameta, a locality in the state of Para,

northern Brazil. This additional label was probably wrongly added later in the vial, as there are no

records for D. macrura from Para. Up to now, all the specimens of D. macrura we examined came only

from the state of Minas Gerais. Therefore, it is clear that the type-locality is really Sao Joao del Rei,

central Minas Gerais state, Brazil. Lriedrich Sellow (or Sello, 1789-1831), a famous Prussian naturalist,

collected a large number of zoological specimens in Minas Gerais state, such as insects (Papavero 1973)

and birds (Rego et al. 2013). Purthermore, most of Sellow s specimens are also deposited at MIN.

Diplura bicolor was described from Cara 9 a, Minas Gerais State, Brazil, based on two female syntypes

(Simon 1889: 215). It is, currently, placed in the genus Linothele, although it was considered a Diplura

by former authors (Mello-Leitao 1937; Biicherl 1957). This species was implicitly transferred to its

current genus following Raven (1985: 182): “all alyrate Diplurinae [... ] are transferred to Linothele ”. As

Simon (1889: 215) had not cited a lyra in his description, Diplura bicolor was regarded as a Linothele

in all later spider catalogs (e.g., World Spider Catalog 2015). Notwithstanding the fact that the first

description of a lyra in the maxilla of Dipluridae was made by Blackwall (1867), this structure was not

mentioned again until the reevaluation of its form and function by Pocock (1896). So it is clear that

Simon and other former authors had not dissected the maxilla of their specimens in search for a lyra.

We have only been able to analyze photos of one syntype of Linothele bicolor (Dipluridae Contributors,

2016), as it is currently in loan to another researcher, but the photos clearly show a lyra formed by 8

setae. Simon (1889: 216) clearly cited the characteristic contrasting color pattern found in D. macrura

in the original description of D. bicolor. However, he also cited some light brown spots scattered on

the posterior half of the abdomen. It is possible that the syntypes cited by Simon are immature or small

mature females (15.3 mm long) since isolated light brown spots on the sides of the abdomen are usually

found only in immatures of D. macrura , disappearing in older males and females. Besides the syntype

photos, the examination of other specimens of D. bicolor from its type-locality (Cara 9 a, Catas Altas) and

nearby areas of Minas Gerais state allow the clear recognition of the species. The examined specimens

9

European Journal of Taxonomy 210: 1-21 (2016)

agree very well to the original description of D. bicolor in color and size, except by the presence of a

lyra. Furthermore, despite several collections on Cara£a and other nearby localities, including extensive

search for Mygalomorphae, we and other researchers had not been able to find any other Diplura or

Linothele species from the same region. The comparison of the above cited specimens with D. macrura

from central Minas Gerais state allowed us to establish this synonymy, based on the similar color pattern

and lyra.

The male of D. bicolor was supposedly described by Mello-Leitao (1937: 3, fig. 3) and Biicherl

(1957: 385, fig. 12-12a). However, both specimens clearly belong to other species. We were able to

examine the “allotypus” of D. bicolor described by Mello-Leitao, from Gustavo da Silveira, Minas

Gerais state (IBSP 3452). It is a male of an undetermined species of Nemesiidae, with a distinctive

copulatory bulb and a very setose pedipalp (Mello-Teitao 1937: fig. 3). Tikewise, judging by the figures

12 and 12a by Biicherl 1957, the male he identified as D. bicolor is actually a Theraphosidae, with the

typical two lobed tibial spur in leg I (Bucherl 1957: fig. 12a) and a copulatory bulb also characteristic of

that family (Bucherl 1957: fig. 12).

Diplura uniformis (Mello-Teitao, 1923) was originally described as Thalerothele uniformis by Mello-

Leitao (1923: 105). He stated that the male holotype was collected in the state of Sao Paulo: Sao Paulo,

by E. Garbe, and should be deposited in Museu de Zoologia de Sao Paulo (MZSP, old number 321).

However, Bucherl et al. (1971: 119, 122) examined a male specimen from MNRJ, collected in Ouro

Preto, labeled as type of I uniformis by Mello-Leitao [MNRJ 192, MLPC 1056]. Since they were not

able to find any type material of the species in MZSP, they considered the MNRJ specimen as the holotype

of T. uniformis. They also stated that the type-locality and repository cited in the description were wrong

(Silva-Moreira et al. 2010: 32). Since a later examination of the MZSP collection by R. Baptista also

did not produced any evidence of the holotype, we follow Bucherl et al. (1971) and Silva-Moreira et

al. (2010) in considering the MNRJ specimen as the actual holotype of T. uniformis. The transfer of

T. uniformis to Diplura was made by Platnick (1993) in his catalog, following Raven (1985).

Bucherl et al. (1971: 123) also considered T. minensis Mello-Leitao, 1926 and T. aurantiaca Mello-

Leitao, 1943, both described from Ouro Preto (Mello-Leitao 1926: 105 and 1943: 255, respectively) as

synonyms of T. uniformis. Bucherl et al. (1971: 119) examined the female holotype of T. aurantiaca ,

collected in Itacolomi, Ouro Preto, by Othon Leonardos (MNRJ 53945), but not the male holotype of T

minensis. In the description of Thalerothele minensis , it is stated that the holotype received the number

MLPC 880. However, this number belongs to a type specimen of Opiliones, as Mello-Leitao wrote

down in the catalogue of the part [or whole?] of his private collection purchased by MNRJ in 1929

(Kury & Baptista 2004). Probably the wrong number in the original description was a misprint or a lapse

(Silva-Moreira et al. 2010: 32).

Unfortunately, we were not able to find any Diplura material from Ouro Preto in MNRJ collection,

despite several attempts. Thus, the MNRJ specimen Bucherl et al. (1971) considered as the holotype of

T. uniformis and the female holotype of T. aurantiaca had also been probably lost later on. We follow

Bucherl et al. (1971) on the synonymies of T. minensis and T. aurantiaca with D. uniformis , considering

that all species were described from Ouro Preto and the original descriptions do not allow the recognition

of any diagnostic trait. T. minensis was diagnosed in relation to T. uniformis (= D. macrura) by Mello-

Leitao (1926: 11) based on the uniformly dark color of the abdomen, 7 setae on lyra and only 5 teeth on

the promargin of chelicera. Nevertheless, the coloration of the venter in D. macrura males vary from

a light hue to a dark color similar to dorsum, the number of setae on lyra varies between 7 to 9 and

the number of cheliceral teeth varies from 8 to 11. The last character is a not reliable one, as it may be

variable even in one specimen, as the 9 and 11 teeth on different sides of the male herein redescribed

demonstrate. The illustrations of the holotype of T. minensis by Mello-Leitao (1926: figs 1-3) are poorly

10

PEDROSO D.R. et al. , Redescription of Diplura macrura and D. lineata

done and not diagnostic. The similarity in color, size and lyra structure indicates that it really is a

synonym of D. macrura.

T. aurantiaca is just the female of T. uniformis (= D. macrurai), with the same color pattern and similar

size as the males. Among the characters mentioned in the original description, only the 6 setae on lyra

fall out of the range for D. macrura (7 to 9). However, the first seta in the lyra is smaller and thinner

than the others (Fig. 10) and the number of setae increases over age. Therefore, it is probable that the

6 setae may represent a normal variation or indicate that the holotype was a small young female. The

holotype vulva poorly illustrated by Biicherl et al. (1971: fig. 12) does not seem to be fully developed.

Apart from the reasons mentioned above, only one species has been collected in Ouro Preto, in spite of

many collecting trips resulting in abundant specimens of D. macrura. Notwithstanding the loss of the

types of the three species from Ouro Preto, the synonymies also avoid the proliferation of nomina dubia.

We compared specimens of Ouro Preto with specimens of D. macrura and this examination indicated

that they all belong to the same species, as they share the same color pattern, structure of lyra and shape

of male and female genitalia.

Habitat notes

D. macrura specimens have been found under fallen logs and rocks in relatively dry areas of Atlantic

Forest or Cerrado vs Atlantic Forest ecotones. The spiders do not make funnel-webs, but applies silk to

the ground or log cavities, sometimes building small entrance silk tubes.

Distribution (Fig. 26)

Known only from localities in central and south Minas Gerais state, Brazil.

Diplura lineata (Fucas, 1857)

Figs 14-26

Mygale lineata Fucas, 1857: 14, fig. la-c (<$)■

Thalerothele fasciata Bertlcau, 1880: 24, pi. 1, f. 6 ($) syn. nov.!

Harmonicon nigridorsi Mello-Feitao, 1924: 186 ( 5 ) syn. nov.!

Diplura fasciata - Simon 1889: 183.

Thalerothele fasciata- Simon 1903: 963. —Mello-Feitao 1923: 101, f. 10.

Thalerothele lineata - Mello-Feitao 1926: 309.

Harmonicon nigridorsi - Mello-Feitao 1926: 315, fig. 4. —Biicherl et al. 1971: 122, figs 15-16 (f).

Paraharmonicon nigridorsi - Mello-Feitao 1926: 316 (tentative superfluous name only).

Type material

Mygale lineata: BRAZIF, Rio de Janeiro: Rio de Janeiro, [no date], near Rio [“environs de Rio”,

probably Tijuca] (S holotype, MNHN, not located).

Thalerothele fasciata'. BRAZIF, Rio de Janeiro, Tijuca [no date], van Beneden leg. ($ holotype, IRSNB,

not located).

Harmonicon nigridorsi'. BRAZIF, Rio de Janeiro, [no date], W.S. Bristowe leg. ($ holotype + $

probably mixed later, MNRJ 17, MFPC 847, examined).

Material Examined

BRASIF: 1 §, 1 juv., Rio de Janeiro, Casimiro de Abreu: Barra de Sao Joao, Morro de Sao Joao,

21-24 Mar. 2003, Expedition Arachne (MNRJ 4322); 1 $, 24 Mar. 2003, Expedition Arachne (MNRJ

11

European Journal of Taxonomy 210: 1-21 (2016)

Table 3. Diplura lineata (Lucas, 1857), <$■ Length of left leg articles (dorsal view).

Leg I

Leg II

Leg III

Leg IV

Fe

6.7

6.4

6.0

7.6

Pa

4.0

3.3

2.8

3.2

Ti

5.6

5.3

4.7

6.1

Mt

5.3

5.9

6.6

9.1

Ta

5.0

4.6

4.0

5.0

Total

26.6

25.5

24.1

31.1

4335); Mangaratiba: 1 c?, 1 ?, Hha de Itacuruga, Aguas Lindas, 24-25 Mar. 2007, R.L.C. Baptista,

C. S. Costa & A.R. Oliveira leg. (LJFRJ 0332); 1 juv., Reserva Ecologica Rio das Pedras, 11-

12 Nov. 2004, A.P.L. Giupponi leg. (MNRJ 4312); 1 juv., Nova Iguagu: Parque Municipal de Nova

Iguagu, 21 Jul. 2004, C. Lima leg. (MNRJ); 1 A, PM Nova Iguagu, (MNRJ 4311); 1 §, Rio de Janeiro,

Parque Estadual da Pedra Branca, (MNRJ 4310); 1 §, Camorim, Agude, 31 Mar. 2014, P. Castanheira

leg. (MNRJ 6820); 1 A, Serra do Mendanha, 485 m. Pitfall, 12 Dec. 2008, J.A.L. Pontes leg. (MNRJ

18434); 1 $, Floresta da Tijuca, 20 Apr. 1986, R.L.C. Baptista leg. (MNRJ 3112 [CRB T005]); 1 ?,

1 juv., Floresta da Tijuca, 400 m, 27 Nov. 1987, A.P.L. Giupponi leg. (MNRJ 3410); 1 $, Floresta

da Tijuca, 7 Jan. 1998, A.P.L. Giupponi leg. (MNRJ 1854); 1 §, Floresta da Tijuca, 400 m, 3 Mar.

2001, A.P.L. Giupponi, D.R. Pedroso & D.F. Almeida leg. (MNRJ 1856); 1 A, Parque Nacional da

Tijuca, 25 May 2001, A.P.L. Giupponi, D.R. Pedroso & R.L.C. Baptista leg. (MNRJ 3568); 1 §, Parque

Nacional da Tijuca, 10 Jan. 2005, D.R. Pedroso & R.L.C. Baptista leg. (MNRJ 4302 [AER]); 1 §, Parque

Nacional da Tijuca, 25 Aug. 2004, D.R. Pedroso & A.P.L. Giupponi leg. (MNRJ 4308); 1 $, Parque

Nacional da Tijuca, Archer, 22 Jan. 2005, D.R. Pedroso & A.P.L. Giupponi leg. (MNRJ 4309); 1 juv.,

Parque Nacional da Tijuca, Gavea, 18 Jan. 2005, D.R. Pedroso leg. (MNRJ 4307 [AER]); 2 $ §, Parque

Nacional da Tijuca, Gavea, 18 Jan. 2005, D.R. Pedroso leg. (MNRJ 4304); 2 Parque Nacional da

Tijuca, Pai Ricardo, 21 Jan. 2005 D.R. Pedroso leg. (MNRJ 4305 [AER]); 1 §, Parque Nacional da

Tijuca, Pai Ricardo, 21 Jan. 2005, D.R. Pedroso leg. (MNRJ 4306 [AER]); 1 §, Parque Nacional da

Tijuca, Pai Ricardo, 21 Jan. 2005, D.R. Pedroso leg. (MNRJ [AER]); 1 §, Parque Nacional da Tijuca,

Sumare, 22 Jan. 2005, D.R. Pedroso leg. (MNRJ 4303 [AER]).

Diagnosis

Both sexes of this species have a characteristic color pattern, similar to the Amazonian D. sanguinea

(F.O. Pickard-Cambridge, 1896). The abdomen of both species have a dark brown dorsum bearing

beige broad transversal stripes. In D. lineata , the stripes are short and broad, with irregular outline,

covering only the side margins of the dorsum. At the sides, there are several small beige spots among

and under the stripes (sometimes fused with them). On the other hand, D. sanguinea have longer and

a bit thinner stripes, without connecting beige spots at the sides. The copulatory bulb is very similar in

both species, but the spermatic duct is very constricted at the basis of the embolus, becoming almost

filiform afterwards, in D. lineata (Fig. 20), while in D. sanguinea the constriction is small, and the

basis of embolus harbors a large duct, which tapers regularly towards the apex. The spermathecae in

D. lineata (Fig. 25) have a thick stem and three distal lobules, while D. sanguinea have a thin stem and

six to seven distal lobules.

12

PEDROSO D.R. et al.. Redescription of Diplura macrura and D. lineata

Figs 14-25. Diplura lineata (Lucas, 1857). 14. $, habitus, dorsal. — 15-21. S, left leg I. 15. Tibial

spur, retrolateral. 16. Tibial spur and metatarsal clasper, ventral. 17. Left palp, retrolateral. 18-21. Bulb

detail. 18. Retrolateral. 19. Prolateral. 20. Ventral. 21. Dorsal. — 22-25. §. 22. Maxilla with lyra. 23.

Lyra detail, SEM. 24. Maxilla with lyra, SEM. 25. Vulva, dorsal. Scale = 1 mm, unless otherwise noted.

13

European Journal of Taxonomy 210: 1-21 (2016)

Description

Male (MNRJ 4311)

Total length 18.5. Carapace: 7.4 long, 6.0 wide, chelicerae 2.7. Abdomen: 7.6 long, 3.9 wide. Spinnerets:

PMS 0.9 long; PLS, total length 10.0, basal article 3.3, middle 3.3, distal 3.3, respectively. Legs: see

Table 3. Carapace: length/width 1.2; flat, cephalic area slightly raised, thoracic furrows shallow and

wide. Fovea: short, deep, slightly recurved. Carapace covered with short, thin setae, interspersed with

some longer and thicker setae; border with abundant long and thick setae pointing outwards, increasing

in number towards posterior angles. Clypeus narrow and small, but clearly visible, not totally hidden

by the eye tubercle, frontal margin bearing 6 thick, long, erect setae. Eye tubercle: 0.5 long, 1.1 wide,

one thick, long seta on tubercle anterior margin, area between posterior eyes covered with thin setae and

bearing 3 thicker, longer setae. AME 0.3, almost spherical, but a bit longer than wide, set apart by half

its diameter. ALE elliptical, much longer than wide, just a bit longer than the AME diameter. PME small,

with flattened lens, longer than wide, its length around 0.6x AME diameter. PLE elliptical, much longer

than wide, its length a bit less than 0.9x AME diameter. PME and PLE closely spaced by around 0.2x

the PME length. Anterior eye row slightly recurved, posterior eye row recurved. Eye rows with similar

width. Chelicerae: formed by 11 teeth on promargin on both left and right chelicera. Plectrum with 4

thick, long setae. Labium: length/width 0.8, no cuspules. Labio-sternal groove deep, with elongated

sigilla. Sternal groove deep, with elongated sigilla. Sternum slightly longer than wide. Posterior angle

in a blunt point, not separating coxae IV. Sigilla: three pairs, elliptical, increasing in size from anterior

to posterior, all near margin. Palp (Fig. 17): relatively short, femur: dl-3-0, pl0-0-l, rl0-0-l, tibia pi

0-2-0, vO. Tibia: 3.1 long, 1.0 wide, short, incrassated, thinner at the basis and apex. Maxillae: length/

width 1.5. Cuspules: 15 spread over ventral inner heel. Lyra: located at the ventral side of the maxilla,

formed by 6 modified thick setae, the basa l m ost much thinner and shorter than the others, which slightly

increase in size from basis to apex of the lyra, all setae wealdy curved at apical portion, with rounded

apex. Legs (Figs 15-16): Leg formula 4123. Legs covered with short, thin, horizontal black setae and

with some longer, thicker, erect black setae. All tarsi with thin scopula, throughout the length of the

article. Tarsi I—III with scopula almost undivided, with only some isolated thicker setae at the middle

line of the basal third of the ventral face. Tarsi IV with scopula partially divided, with thicker setae

covering the basal half of the ventral face. Metatarsi I—II almost without scopula, with only a few setae

near the apex. Metatarsi III-IV without scopula. All tarsi provided with numerous small cracks covering

almost all the ventral and lateral faces, except by the basis and tip of the article. Tibia I around 4.3 x

longer than wide. Retrolateral distal spur placed at the ventral corner of tibia I, with a wide-base, bearing

at its apex a pointed and almost erect megaspine, with similar length of the spur. Metatarsus I relatively

long and a little sinuous on ventral view, with a protruding retrolateral tubercle, placed ventrally on its

basal third, faced towards the apex of the article. Ventral side with an apical spine and two spines on the

median third, the basalmost located in advance to the tubercle. Fringe formed by many spiniform setae

and spines (clasper) covering the median portion of the prolateral side of metatarsus I (Fig. 16). Spines:

leg I : femur d 1-2-1, pldO-2-1, rid 1-1-1 left, rldO-1-2 right; patella 0 left, 0-0-1 right; tibia p0-l-l,

v0-l-lap (apophysis); metatarsus p0-l-0, vO-2-lap; leg II : femur dl-2-1 left, dl-2-0 right, pldO-

2-1, rld0-2-0 left, rldO-2-1 right; patella 0-1-1 left, 0-0-1 right; tibia p0-l-l, vl-l-2ap; metatarsus

pl0-2-0, vl-2-2; leg III : femur dl-2-0, pldO-2-1, rldO-2-1 left, rldl-1-1 right; patella pldO-1-O left,

pld0-l-l right, rld0-l-0; tibia d 1-1-0 left, dO-l-O right, pO-l-O left, p 1-1-0 right, r0-l-l left, r 1-2-0

right, v2-2-2ap; metatarsus d3—1—1 left, d2-3-2 right, p0-2-0 left, p 1-1-0 right, r0-l-0; vl-3-3ap;

leg IV : femur dl-2-0, pldO-2-1, rid 1-2-1 left, rldO-2-1 right; patella 0-1-0 left, 0-0-1 right, tibia

d0-l-0, p0-l-l left, p0-2-0 right, r2—1-1, v2-2-2; metatarsus d2-2-2, p 1—1—1, rl-1-0, v2-3-3ap.

Claws: ITC without teeth. Teeth at STC: aft claws with a wide and high spur at the basis in both sides

leg I : inner and outer rows 9-10; leg II : inner row 9, outer row 8-9; leg III : inner row 6-7, outer row 6;

leg IV : inner and outer rows 6-7. Bulb (Figs 18-21): globose, slightly wider than long, with embolus

moderately long, around 2x the bulb size. Bulb with an abrupt curve near the base of the embolus and a

14

PEDROSO D.R. et al ., Redescription of Diplura macrura and D. lineata

Table 4. Diplura lineata (Lucas, 1857), §. Length of left leg articles (dorsal view).

Leg I

Leg II

Leg III

Leg IV

Fe

7.5

7.0

6.6

7.4

Pa

4.6

4.4

3.7

4.1

Ti

5.5

5.2

4.6

5.9

Mt

5.5

5.5

5.9

6.9

Ta

4.1

4.0

3.6

3.5

Total

27.2

26.1

24.4

27.8

strong constriction on ventral view. Embolus with a relatively thin base, gradually tapering towards the

apex, either on prolateral or retrolateral view. Embolus on ventral view slightly curved at its beginning,

straight through most of its length, with the apex bended retrolaterally. Also on ventral view, a strong

bulge near the base of the embolus and spermatic duct wide, abruptly tapering near the base of the

embolus, becoming thinner, a lm ost filiform.

Female (MNRJ 6820) (Fig 14)

Total length 15.3. Carapace: 6.7 long, 5.4 wide, chelicerae 2.6. Abdomen: 8.6 long, 4.9 wide. Spinnerets:

PMS 1.1 long; PLS, total length 6.9, basal article 2.5, middle 2.2, distal 2.2; respectively. Legs: see

Table 4. Females are very similar to males except by its bigger size and the following characteristics:

carapace length/width 1.2. Clypeus narrow, around A AME diameter, frontal margin with 7 setae. Eye

tubercle with 4 thick setae on anterior margin, area between posterior eyes bearing 5 thick setae. 13 teeth

on promargin of chelicera. Maxilla with 19 (left) and 16 (right) cuspules. Lyra (Fig. 22-24) with 8 setae.

All tarsi with thin scopula, throughout the length of the article. Tarsi I—II with scopula divided by two

series of thicker setae at the middle line of the ventral face, but tarsi III-IV with many setae arranged in

several rows covering most of the ventral face throughout the article. Metatarsi I—II with undivided thin

scopula, covering both sides and the ventral face. Metatarsi III-IV without scopula. All tarsi provided

with few small cracks covering only the median area of the ventral and lateral faces. Spines: leg I: femur

d2—1—1 left, dl-2-0 right; patella 0; tibia p0-0-l, r0-0-l left, rO right, v0-0-2; metatarsus v0-3-2ap;

leg II: femur dl-2-0, pld0-0-l; patella 0; tibia p0-0-l, v0-0-2ap; metatarsus vl-2-2ap; leg III: femur

dl-0-0 left, dO right, pld0-0-l, rld0-0-l left, rld0-0-2 right; patella r0-0-l left, rO-l-O right, pldO-

0-1 right; tibia p 1-2-0 left, pO-l-O right, rl-l-lleft, r 1-2-0 right, v0-0-2ap; metatarsus p 1-3-1 left,

p2-3-2 right, r 1-2-1, vl-3-3ap left, vl-5-3ap right; leg IV: femur dl-2-0, rld0-0-2, patella p0-0-l,

r0-l-0, tibiap0-2-0, r 1-1-1, v0-l-2ap; metatarsus p 1-1-0, rl—1-0, vl-2-lap. Spermathecae (Fig. 25)

separated by about 50% of its length, with a thick stem and similar width up to the distal lobules. Stem

strongly bent forward at the distal third, bearing one large internal lobe just before the curvature and two

apical lobules, all with similar size.

Color pattern

Both sexes with carapace reddish brown, thoracic furrows and cephalic area slightly darker, sometimes

all the carapace dark reddish brown. Eye area black. Chelicera reddish brown; labium, sternum and leg

coxae reddish orange brown with darker sigillae. Legs orange brown. Abdomen dark brown, bearing

short and large light brown transversal stripes with irregular outlines, placed only over the flanks, many

light brown spots between and underneath the stripes. Venter pale brown.

15

European Journal of Taxonomy 210: 1-21 (2016)

Variation

Total length may vary from 16.2 to 18.5 in males and from 15.3 to 26 in females. Cheliceral teeth on

promargin varying from 8 to 11 in both males and females. The maxillary cuspules may vary between

12 and 15 in males and 14 and 19 in females. In females, the lyra is composed by 6 to 8 setae.

Synonymy and notes

Tucas 1857 described Mygale lineata based on a male specimen from the surroundings of the city of

Rio de Janeiro. The collecting locality was probably Tijuca Forest, as many foreigners had houses or

farms in the area and collecting trips to other localities were not encouraged by Brazilian government.

Considering the common practice in that time and the illustration of the habitus (Lucas 1857: fig. 1),

the male holotype was probably dried and pinned. The small abdomen seems shrunk and folded at both

sides of the dorsum, what gives an impression of a median light longitudinal stripe. The holotype may be

lost, as it was not found at the MNHN collection during a visit by the first author and it was not located

by the MNHN curator afterward.

Bertkau (1880) described Thalerothele fasciata based on a female from Tijuca. The holotype of this

species is probably lost, like most Bertkau’s type material (e.g., Levi 1969: 71, 1991: 203, 210; Hofer &

Brescovit 2000: 332). The description given by Bertkau (1880: 24) for Thalerothele fasciata , including

an illustration of the female habitus (Bertkau 1880: fig. 6), is extensive. However, it does not give details

of the genitalia, as Bertkau considered that it was probably an immature female (Bertkau 1880: 25).

On the other hand, the illustration depicts the typical color pattern of D. lineata. The 14.5 mm body

50°0'0"W 48°0'0"W

46°0'0"W 44°0'0"W 42°0'0"W 40°0'0"W

D. lineata

D. macrura

---

50°0'0"W 48°0'0"W 46°0'0"W 44°0'0"W 42°0'0"W o 40 ‘ 0 '°" w

-26°0'0"S

24°0'0"S

•24°0'0"S

26°0'0"S

20°0'0"S

-20°0'0"S

•22°0'0"S

22°0'0"S

Fig. 26. Map showing the known records of Diplnra macrura (C.L. Koch, 1841) and Diplura lineata

(Lucas, 1857).

16

PEDROSO D.R. et al, Redescription of Diplura macrura and D. lineata

length of the immature female holotype is just a little less than the 15.3 to 23.5 mm found in D. lineata

fully grown females. There are 8 teeth on the promargin of the chelicera, what is similar to the 8 to 11

teeth in D. lineata. Compared to the range of 14-19 cuspules in D. lineata , the smaller number (6-7) of

T. fasciata holotype is probably due to its small age, as immature D. lineata have less cuspules on maxilla

and there is a well-known increase in number of cuspules with age and size in many Mygalomorphae.

Mello-Leitao (1924: 186) established Harmonicon nigridorsi (now Diplura nigridorsi) for one female

holotype from the city of Rio de Janeiro. Furthermore, the redundant description of the same species

(Mello-Leitao 1926: 10) also mentioned only the female holotype (“typo”). Both descriptions of

Harmonicon nigridorsi (Mello-Leitao 1924, 1926) are short, without mention to genitalia or any clear

diagnostic trait in relation to D. lineata. When examining the type vial (MNRJ 17, MLPC 847), we

found a male from the same species, besides the female holotype. Biicherl et al. (1971: 119) considered

the male “tipo” (holotype) and the female as a “sintipo” (paratype), in spite of clear indication by Mello-

Leitao that the holotype was a female, without mention to any additional specimen. The female holotype

is badly preserved and darkened, with chelicerae, legs and most of the pedipalps separated from the

body. Only the coxa (maxilla) and trochanter of the right pedipalp are still connected to the holotype

body, and the maxilla of the left pedipalp is missing. Dissection of the genital area revealed an small

genitalia, with lobes not completely developed, indicating that the female was not old, but show the

pattern found in D. lineata (Fig. 25). The color pattern is barely visible, probably due to former events

of desiccation. The original description mentioned a large dark median band on the back of abdomen,

with sinuous margins, delimited by light stripes at each side. Also, there were light spots scattered over

the sides. To date, the shrunk abdomen still displays the light brown transversal stripes with irregular

outlines and some light brown spots underneath the stripes also found in typical D. lineata. The male

(total length 19.5) was clearly erroneously included in the holotype vial and seems to have been collected

later, as it is in a better state than the female, although also shrunk and darkened. The chelicerae, coxae

and trochanter II-IV and left femur IV are still attached to the body. The severely shrunk abdomen still

shows the light brown stripe as in the female. The copulatory bulb, lyra with 6 setae, tibia I retrolateral

spur and metatarsus I tubercle are also similar to D. lineata. The illustration of the copulatory bulb by

Biicherl et al. (1971: fig. 15) is misleading, as it depicts a very elongated and wide embolus, compared

to smaller and thinner embolus found in the vial, similar to D. lineata (Figs 17-21). On the other hand,

the tibia and metatarsus I in fig. 16 are accurate and similar to D. lineata (Fig. 16).

In relation to the original description, there are two numbers for the total length of the holotype of

D. nigridorsi'. 28 mm (Mello-Leitao 1924) and 23 mm (Mello-Leitao 1926). Probablly both numbers

are wrong, but 23 mm falls in the range observed for females of D. lineata (15.3-26) and is closer to

the current shrunk size of the holotype (13.5). Mello-Leitao cited only 8 cheliceral teeth, but there are

also 3 small teeth he may have overlooked. He also cited 12-14 cuspules on the maxilla of the holotype.

The inner corner of the right maxilla has 14 cuspules, but there is a scar on that area, what may indicates

that the original number of cuspules was higher. Anyway, the number of teeth and cuspules both fall

within the range of D. lineata , but there is a lot of variation in the number of those structures throughout

species of Diplura. There are 5 setae in the lyra of the holotype of D. nigridorsi , just one less than the

minimal number found in D. lineata. This is obviously just intraspecific variation, as the female is a

young mature specimen. Finally, only one species of Diplura has been found in the dozens of collection

trips to forested areas in Rio de Janeiro city. Following the reasons above, the three described species

are considered synonyms, and D. lineata prevails as the senior synonym.

Habitat

Specimens of D. lineata have been found under fallen logs in humid areas of the Atlantic forest. The

spiders do not make funnel-webs, but apply silk to the tunnels or log cavities. Sometimes they build

small entrance silk tubes or connect some tubes in a small silk network inside log cavities.

17

European Journal of Taxonomy 210: 1-21 (2016)

Distribution (Fig. 26)

Known only from the state of Rio de Janeiro, Brazil, restricted to the municipalities of Casimiro de

Abreu, Nova Igua 9 u, Rio de Janeiro and Mangaratiba. The records for Venezuela and Colombia (Simon

1889: 188) and for the state of Santa Catarina, Brazil (Mello-Leitao 1923: 101-102) are probably

erroneous. The specimens cited by Simon may belong to D. sanguinea (F.O. Pickard-Cambridge, 1896)

or a related species, as we have examined specimens from Colombia belonging to that species-group.

Discussion

Currently, the accepted distribution of Diplura includes Cuba, Venezuela, Ecuador, Brazil, Bolivia,

Paraguay and Argentina (World Spider Catalog 2015). As discussed above, the records for D. macrura

from Cuba or any other Caribbean or West Indies localities are erroneous. We have been able to examine

a male of Diplura sanguinea from Arraijan: Cerro Silvestre, belonging to personal collection of Roberto

Miranda. Therefore, the norther nm ost record known for Diplura is from a locality in central Panama,

near Panama city. As pointed out above, Colombia should also be included in the distribution range for

Diplura.

One additional remark on the distribution of Diplura is the correct type-locality of Diplura parallela

(Mello-Leitao, 1923). This species is erroneously cited for Argentina in the World Spider Catalog (2015)

and older catalogs. However, the holotype and only known specimen of D. parallela was collected from

an unknown locality in the state of Parana, Brazil, as Mello-Leitao (1923: 100) stated in the original

description. This mistake has probably arisen from the homonymy of Parana, a locality in the Entre-Rios

province, northern Argentina, and the state of Parana, in southern Brazil.

Taking into account the synonymies we established in this paper, 17 valid described species remain in

Diplura. Ten of the described species are recorded from Brazil, two species each from Argentina and

Bolivia, and just one for Ecuador, Paraguay and Venezuela. Additionally, there are records of unidentified

species from Colombia and Panama. In summary, the revised distribution range of Diplura goes from

south Panama to north Argentina.

Acknowledgments

We thank Osvaldo Villarreal Manzanilla (MNRJ) and Adriano Brilhante Kury for the aid with the map,

Marcelo Henrique de Oliveira Sales (UFRJ) by technical support on SEM photos, Ricardo Monteiro and

Margarete Macedo, of the Laboratorio de Ecologia de Insetos (UFRJ), and Sergio Potsch de Carvalho-

e-Silva, of the Laboratorio de Herpetologia (UFRJ), for allowing the use of equipment for photos and

measurements. We also thank Rogerio Bertani (Instituto Butantan) for the live specimen photographs

and additional samples. This work was supported by a Ph.D. grant from Funda 9 ao de Amparo a Pesquisa

do Estado do Rio de Janeiro (FAPERJ).

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Published on: 7 July 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;

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