A. K. Hundsdoerfer Museum of Zoology




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Abbreviations. PM = population mean(s); P.no. = popula- tion number, referring to Appendix: Table A1.
Order Notostraca Sars, 1867

Family Triopsidae Keilhack, 1909

Genus Triops Schrank, 1803



Fig. 7 Triops baeticus n. sp., adult male (holotype). a Telson, dorsal view. b Distal part of 2nd trunk limb. c Proximal region of



5th endite of 2nd trunk limb


Triops baeticus Korn n. sp.
(Fig. 7)

Note. This species refers to the ‘S.Iberian’ phylogenetic lineage.

Etymology. The specific epithet refers to the Roman

province Baetica, named after the river Baetis (modern Guadalquivir). The delta region of that river represents the barycentre of the new species’ geographic distribution, and its diversity hotspot. The epithet is to be treated as a noun in apposition for the purposes of nomenclature.

Material examined. Holotype ♂, Museum of Zoology

Dresden (MTD Crus 3434); Spain, Sevilla, Coto de Doñana National Park, Caño Travieso, P.no. 007, leg. Hugues Lefranc 6 December 2006. Paratypes: 6 ♂♂, 3 ♀♀, Museum of Zoology Dresden (MTD Crus 3299–3301,

3430–3433, 3435–3436); same data as holotype. Other material: P.no. 001, 1♂, 1♀; P.no. 002, 4♂♂, 3♀♀; P.no.

005, 4♂♂, 5♀♀; P.no. 008, 4♂♂, 3♀♀; P.no. 010, 2♂♂; P. no. 013, 3♂♂, 1♀♀; P.no. 014, 1♂; P.no. 015, 2♀♀; P.no.

019, 2♀♀; P.no. 022, 5♂♂, 7♀♀; P.no. 025, 1♂, 1♀; P.no.

026, 2♀♀; P.no. 027, 1♂; P.no. 030, 1♂; P.no. 032, 7♂♂,

4♀♀; P.no. 033, 9♂♂, 4♀♀; P.no. 034, 6♂♂, 6♀♀; P.no.

035, 1♀; P.no. 036, 5♂♂, 6♀♀; P.no. 037, 5♂♂, 3♀♀; P. no. 038, 7♂♂, 7♀♀; P.no. 039, 2♂♂, 2♀♀; P.no. 041,

5♀♀; P.no. 042, 1♂, 1♀; P.no. 045, 4♂♂, 3♀♀; P.no. 046,

6♂♂, 4♀♀; P.no. 048, 1♂, 1♀; P.no. 049, 10♂♂, 4♀♀; P. no. 053, 1♀; P.no. 054, 1♀; P.no. 056, 2♂♂, 3♀♀; P.no.

057, 4♂♂; P.no. 058, 3♀♀; P.no. 059, 2♂♂, 1♀♀; P.no.

060, 6♂♂, 9♀♀; P.no. 061, 4♀♀; P.no. 063, 3♂♂, 3♀♀; P. no. 064, 6♂♂, 6♀♀; P.no. 065, 6♂♂, 5♀♀; P.no. 066,

2♀♀; P.no. 069, 7♂♂, 6♀♀; P.no. 076, 1♀; P.no. 080,

2♂♂; P.no. 081, 2♂♂, 2♀♀.

Diagnosis. Adult male: 10th trunk limb: 7–13 (PM

10.0–11.9) anterior meshwork spines on 3rd endite, 7–15 (PM 10.4–13.5) submarginal spines on 4th endite; 2nd trunk limb: proportional endopodite length 72.9–95.5% (PM 77.3–89.6%; standardized values), proportional max- imum length of spines on 5th endite 4.5–7.8% (PM 5.5–

7.0%; standardized values); telson length ratio 0.35–0.57 (PM 0.39–0.53), furcal spine width / telson width 0.11–0.20 (PM 0.12–0.16), length of telson posterior incision / telson width 0.024–0.098 (PM 0.039–0.079); apodous abdominal segments 5.4–7.7 (PM 6.2–7.2). Adult female: 10th trunk limb: 8–15 (PM 10.0–13.7) anterior meshwork spines on

3rd endite, 8–17 (PM 10.7–14.7) submarginal spines on 4th endite; telson length ratio 0.33–0.53 (PM 0.38–0.48), furcal spine width / telson width 0.11–0.18 (PM 0.12–0.16), length of telson posterior incision / telson width 0.029–

0.098 (PM 0.040–0.086); apodous abdominal segments

4.8–6.7 (PM 5.1–6.4). Resting egg: diameter up to 552 μm

(PM 438–499 μm).

Description of holotype. Adult male, body rings 32; total number of trunk limbs 55; 10th trunk limb: 11 anterior meshwork spines on 3rd endite, 12 submarginal spines on

4th endite; 2nd trunk limb: length of endopodite 2,558 μm, length of 5th endite 4,178 μm, longest spine on 5th endite

136 μm; length of carapace 22.52 mm, 24 dorsal carina spines, sulcus with 26 spines, length of nuchal organ

0.49 mm; 2nd maxilla well developed, with long terminal

setae; telson width 2.73 mm, telson length ratio 0.42, furcal spine size ratio 2.19, posterior marginal spines of medium size, 6 median spines (the 4 anteriormost very small); approximately 7 furcal spines on both sides (not clearly separated from a row of small spines covering posterior margin of ventral side of telson), on one side, main spine not much bigger than first of the following spines; apodous abdominal segments 6.6, supernumerary spines 1.

Range. Guadalquivir and Guadiana River valleys and adjacent areas. Northernmost records are from Tagus River catchment. In south central Portugal recorded from area north of Serra de Monchique and Serra do Caldeirão.

Triops gadensis Korn & García-de-Lomas n. sp. (Fig. 8)

Note. This species refers to the ‘Cadiz’ phylogenetic

lineage.




Fig. 8 Triops gadensis n. sp., adult male (holotype). a Telson, dorsal view. b Distal part of 2nd trunk limb. c Proximal region of



5th endite of 2nd trunk limb



Etymology. The specific epithet refers to Gades, the Roman name for the town of Cádiz. The epithet is to be treated as adjectival for the purposes of nomenclature.

Material examined. Holotype ♂, Museum of Zoology

Dresden (MTD Crus 3089); Spain, Cádiz, Barbate, P.no.

083, leg. Juan García-de-Lomas 3 April 2006. Paratypes: 3

♂♂, 2 ♀♀, Museum of Zoology Dresden (MTD Crus

3060–3061, 3086–3088); same data as holotype. Other material: P.no. 082, 6♂♂, 14♀♀; P.no. 084, 1♂, 4♀♀; P. no. 085, 2♂♂, 2♀♀; P.no. 086, 1♂; P.no. 087, 5♂♂, 8♀♀; P.no. 090, 1♂; P.no. 091, 3♀♀.

Diagnosis. Adult male: 10th trunk limb: 9–11 (PM 9.8–

10.2) anterior meshwork spines on 3rd endite, 9–13 (PM

10.2–12.3) submarginal spines on 4th endite; 2nd trunk limb: proportional endopodite length 73.3–85.1% (PM

79.4–84.7%; standardized values), proportional maximum length of spines on 5th endite 4.3–6.1% (PM 4.9–5.5%; standardized values); telson length ratio 0.40–0.52 (PM

0.42–0.47), furcal spine width / telson width 0.10–0.19 (PM 0.13–0.17), length of telson posterior incision / telson width 0.034–0.099 (PM 0.055–0.068); apodous abdominal segments 5.7–7.2 (PM 6.2–6.8). Adult female: 10th trunk limb: 9–14 (PM 10.8–12.7) anterior meshwork spines on

3rd endite, 8–16 (PM 10.3–13.6) submarginal spines on 4th endite; telson length ratio 0.33–0.53 (PM 0.39–0.46), furcal spine width / telson width 0.12–0.18 (PM 0.12–0.15), length of telson posterior incision / telson width 0.033–

0.082 (PM 0.058–0.064); apodous abdominal segments

4.4–6.5 (PM 5.1–5.7). Resting egg: diameter up to 479 μm

(PM 448 μm).

Description of holotype. Adult male, body rings 31; total number of trunk limbs 50; 10th trunk limb: 11 anterior meshwork spines on 3rd endite, 11 submarginal spines on 4th endite; 2nd trunk limb: length of endopodite 2,674 μm, length of 5th endite 3,641 μm, longest spine on 5th endite 131 μm; length of carapace

19.05 mm, 48 dorsal carina spines, sulcus with 39 spines, length of nuchal organ 0.51 mm; 2nd maxilla

well developed, with long terminal setae; telson width

2.37 mm, telson length ratio 0.46, furcal spine size ratio 2.07, posterior marginal spines long, 2 median spines; 3 + 4 furcal spines, main spines considerably bigger than following spines; apodous abdominal segments 6.3.

Range. Western lowlands of Cádiz province, between El

Puerto de Santa Maria and Tarifa.
Triops vicentinus Korn, Machado, Cristo & Cancela da

Fonseca n. sp.


(Fig. 9)

Note. This species refers to the ‘Portuguese’ phyloge- netic lineage.

Etymology. The specific epithet refers to the new species’ barycentre of distribution along the Costa Vicentina in south- west Portugal. The epithet is to be treated as a noun in apposition for the purposes of nomenclature.

Material examined. Holotype ♂, Museum of Zoology Dresden (MTD Crus 3158); Portugal, Sagres, Parque Natural do Sudoeste Alentejano e Costa Vicentina PNSACV, pond G3, P.no. 095, leg. Margarida Machado, Margarida Cristo and Luís Cancela da Fonseca 3 February

2006. Paratypes: 9 ♂♂, 13 ♀♀, Museum of Zoology Dresden (MTD Crus 3124–3137, 3153–3157, 3159–3162); same data as holotype. Other material: P.no. 094, 5♂♂,

6♀♀; P.no. 096, 9♂♂, 3♀♀; P.no. 097, 6♂♂, 6♀♀; P.no.

101, 6♂♂, 5♀♀.

Diagnosis. Adult male: 10th trunk limb: 11–14 (PM

11.7–12.4) anterior meshwork spines on 3rd endite, 12–17 (PM 13.3–15.7) submarginal spines on 4th endite; 2nd trunk limb: proportional endopodite length 79.5–94.3% (PM 82.1–86.9%; standardized values), proportional max- imum length of spines on 5th endite 4.3–7.0% (PM 4.6–

5.9%; standardized values); telson length ratio 0.32–0.49

(PM 0.35–0.42), furcal spine width / telson width 0.11–0.16 (PM 0.12–0.14), length of telson posterior incision / telson width 0.040–0.122 (PM 0.071–0.099); apodous abdominal



Fig. 9 Triops vicentinus n. sp., adult male (holotype). a Proxi- mal region of 3rd endite of 10th trunk limb, anterior view; meshwork spines (and associat- ed row of spinules) shown in black, submarginal spines and edge of endite in grey. b Detail of 10th trunk limb, anterior view (EN3, 4 = 3rd, 4th endite; MWSP = meshwork spines; SMSP = submarginal spines). c Distal part of 2nd trunk limb. d Proximal region of 5th endite of

2nd trunk limb. e Telson, dorsal

view. f Detail of 4th endite of

10th trunk limb, posterior view; only proximalmost parts of pos- terior row of meshwork spines shown, anterior row of mesh- work spines in grey


segments 5.7–7.3 (PM 6.4–7.0). Adult female: 10th trunk limb: 11–17 (PM 13.3–15.1) anterior meshwork spines on

3rd endite, 14–19 (PM 14.0–17.7) submarginal spines on

4th endite; telson length ratio 0.29–0.45 (PM 0.34–0.43), furcal spine width / telson width 0.11–0.18 (PM 0.12–0.17), length of telson posterior incision / telson width 0.048–

0.134 (PM 0.080–0.099); apodous abdominal segments

5.3–6.5 (PM 5.5–6.4). Resting egg: diameter up to 583 μm

(PM 524–532 μm).

Description of holotype. Adult male, body rings 33; total number of trunk limbs 57; 10th trunk limb: 12 anterior meshwork spines on 3rd endite, 17 submarginal spines on

4th endite; 2nd trunk limb: length of endopodite 2,891 μm, length of 5th endite 4,108 μm, longest spine on 5th endite

119 μm; length of carapace 23.26 mm, 47 dorsal carina spines, sulcus with 45 spines, length of nuchal organ

0.49 mm; 2nd maxilla well developed, with long terminal

setae; telson width 2.97 mm, telson length ratio 0.34, furcal



Fig. 10 Triops emeritensis n. sp., adult male (holotype). a Distal part of 2nd trunk limb. b Telson, dorsal view. c Proximal region of 3rd endite of 10th trunk limb, anterior view; meshwork spines and associated row of spinules shown in black, submarginal spines and edge of endite in grey






spine size ratio 1.72, posterior marginal spines long, 1 median spine; 7 furcal spines on both sides, main spine considerably bigger than following spines; apodous ab- dominal segments 6.6.

Range. Costa Vicentina to south central Algarve.

Triops emeritensis Korn & Pérez-Bote n. sp. (Fig. 10)

Note. This species refers to the ‘Gitanilla’ phylogenetic lineage.

Etymology. The specific epithet refers to the geographic proximity of the type locality to the Roman town Emerita Augusta, the modern Mérida. The epithet is to be treated as adjectival for the purposes of nomenclature.

Material examined. Holotype ♂, Museum of Zoology Dresden (MTD Crus 3109); Spain, Badajoz, La Albuera, Laguna de la Gitanilla, P.no. 103, leg. José L. Pérez-Bote, reared from sediments). Paratypes: 16 ♂♂, 12 ♀♀, Museum of Zoology Dresden (MTD Crus 2640, 2764–

2773, 3100–3102, 3104–3108, 3110–3118); same data as holotype. Other material: P.no. 102, 1♂, leg. Miguel Alonso April 2006.

Diagnosis. Adult male: 10th trunk limb: 8–10 (PM 9.2) anterior meshwork spines on 3rd endite, 9–14 (PM 10.5) submarginal spines on 4th endite; 2nd trunk limb: propor- tional endopodite length 69.8–80.5% (PM 74.0%; standard- ized values), proportional maximum length of spines on 5th endite 3.7–6.2% (PM 4.8%; standardized values); telson length ratio 0.38–0.51 (PM 0.43), furcal spine width / telson width 0.13–0.19 (PM 0.16), length of telson posterior incision / telson width 0.063–0.104 (PM 0.084); apodous abdominal segments 6.2–7.5 (PM 6.8). Adult female: 10th trunk limb: 9–12 (PM 11.4) anterior meshwork spines on 3rd endite, 10–14 (PM 12.2) submarginal spines on 4th endite; telson length ratio 0.36–0.49 (PM 0.42), furcal spine width / telson width 0.14–0.21 (PM 0.17), length of telson posterior incision / telson width 0.58–0.119 (PM 0.086); apodous abdominal segments 5.5–6.4 (PM 6.0). Resting egg: diam- eter up to 593 μm (PM 553 μm).

Description of holotype. Adult male, body rings 32; total number of trunk limbs 55; 10th trunk limb: 10 anterior meshwork spines on 3rd endite, 10 submarginal spines on

4th endite; 2nd trunk limb: length of endopodite 2,660 μm, length of 5th endite 4,342 μm, longest spine on 5th endite

110 μm; length of carapace 19.85 mm, 35 dorsal carina

spines, sulcus with 33 spines, length of nuchal organ

0.69 mm; 2nd maxilla well developed, with long terminal setae; telson width 2.51 mm, telson length ratio 0.45, furcal spine size ratio 1.74, posterior marginal spines short, 2 median spines; 4 furcal spines on both sides, main spine only slightly bigger than following spine; apodous abdom- inal segments 6.6.

Range. Extremadura, Guadiana valley, recorded in two ponds near La Albuera.


Key to the southern Iberian species of Triops
This key uses population means of character states from both adult sexes and resting eggs; it is not designed for the identification of individual specimens. Couplet 1 includes values for the furcal spine size ratio in order to avoid possible confusion with short-spined taxa, even though these have not been reported from southern Iberia.
1. Furcal spine size ratio ≥0.34; 10th trunk limb with

≥11.7 (males), 13.3 (females) anterior meshwork spines on 3rd endite and with ≥13.3 (males), 14.0 (females) submarginal spines on 4th endite; length of telson posterior incision / telson width ≥0.074; diameter of resting eggs >500 μm; geographic range: Costa Vicentina to south central Algarve ....................................

..................................................... Triops vicentinus n. sp.

– Furcal spine size ratio ≥0.34; 10th trunk limb either with <11.7 (males), 13.3 (females) anterior meshwork spines on 3rd endite or with <13.3 (males), 14.0 (females) submarginal spines on 4th endite, or counts of both types of spines below the given values

......................................................................................... 2

2. Proportional endopodite length of 2nd trunk limb in males (size standardized) <75%; length of telson posterior incision / telson width >0.080; diameter of resting eggs >500 μm; geographic range: Extremadura, Guadiana valley ...................... Triops emeritensis n. sp.

– Proportional endopodite length of 2nd trunk limb in males (size standardized) 77–90%; length of telson posterior incision / telson width <0.080; diameter of resting eggs ≤500 μm .................................................. 3

3. Proportional spine length of 5th endite of 2nd trunk limb in males (size standardized) ≤5.5%; geographic range: western lowlands of Cádiz province, between El Puerto de Santa Maria and Tarifa .....................................

........................................................ Triops gadensis n. sp.

– Proportional spine length of 5th endite of 2nd trunk limb in males (size standardized) ≥5.5%; geographic range: Guadalquivir and Guadiana River valleys and adjacent areas, in Cádiz province north of Guadalete River, in south central Portugal north of Serra de Monchique and Serra do Caldeirão ............................

........................................................ Triops baeticus n. sp.

Acknowledgements We are very grateful to Miguel Alonso (Barcelona), Federico Marrone (Palermo), Yasar Al-Khalili (Pest Management Consultants, Dubai) and Ernst-Gerhard Burmeister (München) for providing samples for this investigation, to Hugues Lefranc (Sevilla) for his help in taking samples, and to David Paz and Hector Garrido (Sevilla) for assistance in the field. We are greatly indebted to


Christian Kehlmaier and Anke Müller (Dresden) for their dedicated engagement in the sequencing work. Katja Frohberg (Dresden) further contributed to the sequencing work in this study. We are grateful to two anonymous reviewers, as well as to Olaf R.P. Bininda-Emonds (Chief Editor) and Martin V. Sørensen (Associate Editor) for their most helpful suggestions that greatly improved the manuscript. We would further like to thank Ian J. Kitching (London) for advice on the taxonomy section.



Appendix

Table A1 Geographic origin, museum specimen tissue voucher numbers (MTD-TW) and short names of 12S and 16S haplotypes (indicating respective main phylogenetic lineage and individual haplotype) of Triops mauritanicus and T. c. cancriformis specimens used for genetic investigations and for molecular determinations of samples investigated morphologically in the present study. For samples from a preceding study (Korn et al. 2006), GenBank accession numbers (all beginning with AM18-) are given (in parentheses) instead of tissue voucher numbers. Abbreviations: Pop. no. = number code identifying population in present study; G = Gitanilla, M = Morocco, P = Portugal, S = Spain




Pop. no.

Voucher numbers (GenBank accession numbers AM18-)

Geographic origin

Haplotypes 12S, [16S]

001

423, 796-800, 1998-2003

S, Doñana National Park

S.Iberia 1, 2, 3, 17, 18, 22

002

418, 459-463, 1943-1948

S, Doñana National Park

S.Iberia 2, 17, 18, 24, [1]

003

3164-3169

S, Doñana National Park

S.Iberia 1, 2, 3

004

3170-3175

S, Doñana National Park

S.Iberia 3, 17, 20, 24, 26

005

1918-1923

S, Doñana National Park

S.Iberia 2, 3, 17, 18

006

3176-3181

S, Doñana National Park

S.Iberia 1, 2, 3, 24, 27

007

3185, 3227-3231

S, Doñana National Park

S.Iberia 1, 3

008

1955-1960

S, Doñana National Park

S.Iberia 1, 2, 17

009

557, 558, 984-987

S, Doñana National Park

S.Iberia 17, 18, 28

010

419, 952-956

S, Almonte

S.Iberia 1

011

801-806

S, N. Doñana Natural Park

S.Iberia 2

012

778, 922-926

S, N. Doñana Natural Park

S.Iberia 14, 15

013

836-841

S, N. Doñana Natural Park

S.Iberia 9

014

853-858

S, Doñana National Park

S.Iberia 1

015

426, 859-863

S, Doñana National Park

S.Iberia 2, [1]

016

427, 791-795

S, Doñana National Park

S.Iberia 2

017

807-812

S, Doñana National Park

S.Iberia 2

018

842-847

S, W. Doñana Natural Park

S.Iberia 2

019

414, 947-951

S, W. Doñana Natural Park

S.Iberia 1

020

819-824

S, W. Doñana Natural Park

S.Iberia 2

021

776, 903-907

S, Doñana National Park

S.Iberia 1, 2, 19

022

813-818

S, Doñana National Park

S.Iberia 3, 19

023

779-784

S, E. Doñana Natural Park

S.Iberia 1, 3

024

428, 962-966

S, E. Doñana Natural Park

S.Iberia 1, 4, [1]

025

785-790

S, E. Doñana Natural Park

S.Iberia 1, 4

026

424, 912-916

S, N. Doñana Natural Park

S.Iberia 2, 7, 21, [2]

027

830-835

S, N. Doñana Natural Park

S.Iberia 18

028

422, 957-961

S, Villamanrique

S.Iberia 4, 7, [1]

029

621, 848-852

S, Villamanrique

S.Iberia 4

030

2004-2009

S, El Cuervo

S.Iberia 1, 13

031

413, 917-921

S, Fuentes de Andalucía

S.Iberia 1, 25

032

429, 932-936

S, Cañada Rosal

S.Iberia 1

033

444, 967-971

P, Castro Marim

S.Iberia 12, [1]

034

1982-1987

P, Serpa

S.Iberia 1
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