Bonn zoological Bulletin 67 (1): 1-3

May 2018

In memoriam Dr. Hans Ulrich (1934-2017)

Wolfgang Bohme & Rainer Hutterer

Department of Vertebrates, Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany

Email: w.boehme@leibniz-zfmk. de

On May 24, 2017, the long-term curator of the Diptera

section of Museum Koenig, Dr. Hans Ulrich, passed

away at the age of 83 years. With this obituary, we want

to commemorate and celebrate his life and his scientific

activities.

Hans Ulrich was born as the third child of the former

Reichsbank inspector and later Bankrat (retired) Ferdi-

nand Ernst Hans Hugo Ulrich and his wife Matilde Mar-

garethe Marie Ulrich, born Lollke, on July 22, 1934 at

Stendal, Saxony-Anhalt. In 1938 the family moved to

Ludwigsburg in Baden-Wutrttemberg. In December 1944,

an aerial bomb attack destroyed most of the household.

They moved to relatives at Nordhausen (Harz) where in

two new aerial attacks in April 1945 the family lost again

all they had. Shortly after the end of World War II and

before the occupation of Thuringia by the Soviet army

they returned to Wirttemberg and found a first quarter at

Mundelsheim on the Neckar, then in Ludwigsburg and

finally in Stuttgart.

Hans Ulrich entered basic school in spring 1940 and

moved to high school in summer 1944. He finished his

maturation at the Friedrich Schiller High School in Lud-

wigsburg. He started his university studies in biology,

chemistry and geology at the Technical University Stutt-

gart and at the University of Tubingen in summer semes-

ter 1953 and finished in spring 1960 with an examination

(state examen) for school teachers. On March 12, 1962

he received his doctoral degree with a histological dis-

sertation on hippoboscid flies (Diptera: Hippoboscidae),

at the Faculty of Natural Sciences of the Technical Uni-

versity of Stuttgart, under the tutorship of Prof. Dr. Otto

Pflugfelder (1904-1994).

From May 1957 to April 1962, with one year inter-

ruption, Hans Ulrich served the Zoological Institute of

the Technical University Stuttgart and the nearby Ag-

ricultural University Hohenheim. Subsequently, from

May 1962 to June 1963, he became part of a team at

the Landesanstalt fiir Pflanzenschutz (State institute for

plant protection) within a focus program of the Deutsche

Forschungsgemeinschaft (German Science Foundation,

DFG) called “Integrative plant protection”. Next station

of his career was a temporary employment as a scien-

tific assistant at the Staatliches Museum fur Naturkunde

in Stuttgart, Department of Phylogenetic Research, from

July 1963 to December 1965. These two and a half years

were particularly important for him, since he proved to

be the only assistant of the world-famous phylogeneticist

Prof. Willi Hennig (1913-1976), and he became strongly

influenced by Hennigian thinking for all his further sci-

entific work.

A further station on his way was a temporary employ-

ment, within a DFG sponsored project at the Institut fur

Biologische Schadlingsbekampfung (Institute for bio-

logical combat against noxious organisms) in Darmstadt,

combatting the noxious tortricid moth Cydia pomonel-

la Linnaeus, 1758 (Lepidoptera: Tortricidae). And only

2 Wolfgang Bohme & Rainer Hutterer

thereafter, being employed again by the DFG at the Zo-

ologische Staatssammlung Munchen (Zoological State

Collection Munich), he finally found his long-term spe-

cial field of research, viz. the taxonomy and anatomy of

two dipteran families: Dolichopodidae and Empididae.

In December 1971, he entered the Zoologisches For-

schungsmuseum Alexander Koenig as a curator of dip-

terology and took responsibility for a collection which

had been brought to international reputation by his prede-

cessor Dr. Bernhard Mannheims (1909-1971), mainly in

respect to blepharicerids, tipulids and limontids. Interna-

tionally important was also the collection of phorid flies.

Until the employment of a curator of hymenopterology

in 1986, Hans Ulrich took also care of the Hymenoptera

collection. In 1974 he became a preliminary civil servant

which was extended to permanency in 1977.

Since 1974 he had an own assistant in his section, Mrs.

Christa GroBe-Streuer, who helped him in all respects

including his activity as editor of two ZFMK journals

“Bonner zoologische Beitrage” (Festschrift H. E. Wolt-

ers 1975 and volumes 31-35, 1980-1984) and “Bonner

zoologische Monographien” (nos 12—16, 1979-1982).

Hans Ulrich worked as a curator until 1999, and then

continued as a volunteer until about 2012, his visits be-

came rare. He spent quite some money on fossil Diptera,

most of which he donated to the museum’s collections.

He also donated his books, and finally his house. He was

a really generous curator.

Taxon names coined by H. Ulrich and colleagues

Amphithalassius Ulrich, 1991

Amphithalassius latus Ulrich, 1991

Amphithalassius piricornis Ulrich, 1991

Microphorella similis Brooks & Ulrich, 2012

Plesiothalassius flavus Ulrich, 1991

Plesiothalassius natalensis Ulrich, 1991

(all Diptera: Dolichopodidae)

Taxa named after H. Ulrich

Ulrichophora Brown, 2007 (Diptera: Phoridae)

Metopina ulrichi Disney, 1979 (Diptera: Phoridae)

Microphorella ulrichi Gatt, 2003

(Diptera: Dolichopodidae)

Publications List

Ulrich H (1963) Vergleichend histologische und zyklische Un-

tersuchungen an den weiblichen Geschlechtsorganen und

den innersekretorischen Drtisen adulter Hippobosciden (Dip-

tera Pupipara). Deutsche entomologische Zeitschrift (Neue

Folge) 10: 28-71

Bonn zoological Bulletin 67 (1): 1-3

Ulrich H (1965) Der Fang- und Greifapparat von Mantispa —

ein Vergleich mit Mantis. Natur und Museum 95: 499-508

Ulrich H (1966) Zum Aufbau der Ovarien von Glossina mor-

sitans (Diptera). Deutsche entomologische Zeitschrift (neue

Folge) 13: 379-382

Ulrich H (1968) Versuche tiber die Empfindlichkeit von 7ri-

chogramma (Hymenoptera, Chalcidoidea) gegentiber Fun-

giziden. Anzeiger fur Schadlingskunde 40: 101—106

Ulrich H (1968) Ein verbesserter Kafig fiir die Massenaufzucht

des Eiparasiten Trichogramma (Hymenoptera, Chalci-

doidea). Entomophaga 13: 233-236

Ulrich H (1971) Zur Skelett- und Muskelanatomie des Thorax

der Dolichopodiden und Empididen (Diptera). Verdffen-

tlichungen d. Zoologischen Staatssammlung Munchen 15:

1-44

Ulrich H (1972) Der Gattungsname Anoplomerus Rondani

(Diptera, Dolichopodidae). Opuscula zoologica 111: 14

Ulrich H (1972) Zur Anatomie des Empididen-Hypopygiums

(Diptera). Veroffentlichungen der Zoologischen Staatssam-

mlung Munchen 16: 1—27

Steyskal G C, Robinson H, Ulrich H, Hurley R L (1973) Ay-

drophorus Fallén, 1823 (Insecta, Diptera, Dolichopodidae):

Request for suppression under the plenary powers of the des-

ignation by Macquart, 1827 of H. jaculus Fallén as type of

the genus in favour of H. nebulosus Fallén in order to con-

serve consistent usage. Bulletin of zoological Nomenclature

30: 118-120

Ulrich H (1974) Das Hypopygium der Dolichopodiden (Dip-

tera): Homologie und Grundplanmerkmale. Bonner zoolo-

gische Monographien 5: 1-60

Ulrich H (1975) Das Hypopygium von Chelifera precabunda

Collin (Diptera, Empididae). Bonner zoologische Beitrage

26: 264-279

Ulrich H (1978) Abwandlungen im Bau des Dolichopo-

diden-Hypopygiums (Diptera) — I: Medetera truncorum Mei-

gen. Bonner zoologische Beitrage 28: 412-420

Ulrich H (1980) Subfamilies of Dolichopodidae (Diptera). Ab-

stracts of the 16" International Congress of Entomology: 24

Ulrich H (1980) Zur systematischen Gliederung der Dolichopo-

diden (Diptera). Bonner zoologische Beitrage 31: 385-402

Ulrich H (1984) Skelett und Muskulatur des Thorax von Micro-

phor holosericeus (Meigen) (Diptera, Empidoidea). Bonner

zoologische Beitrage 35: 351-398

Ulrich H (1988) Das Hypopygium von Microphor holosericeus

(Meigen) (Diptera, Empidoidea). Bonner zoologische Beit-

rage 39: 179-219

Ulrich H (1991) Two new genera of parathalassiine-like flies

from South Africa (Diptera, Empidoidea). Bonner zoolo-

gische Beitrage 42: 187-216

Ulrich H, Schmelz R M (1998) Enchytraeidae (Oligochaeta) as

prey of Dolichopodidae, recent and in Baltic amber. Abstract

4" international Congress on Diptera (Oxford): 231

Ulrich H, Schmelz R M (2001) Enchytraeidae as prey of Dol-

ichopodidae, recent and in Baltic amber (Oligochaeta; Dip-

tera). Bonner zoologische Beitrage 50: 89-101

Ulrich H (2003) How recent are the Empidoidea of Baltic am-

ber? [Wie rezent sind die Empidoidea des Baltischen Bern-

steins?]. Studia dipterologica 10 (1): 321-327

Ulrich H (2005) Predation by adult Dolichopodidae (Diptera):

a review of literature with an annotated prey—predator list.

[Dolichopodiden (Diptera) als Pradatoren im Imaginalstadi-

um: eine Literaturtibersicht mit einem kommentierten Beu-

te-Rauber- Verzeichnis. ] Studia dipterologica 11 (2): 369-403

Brooks S E, Ulrich H (2012) Microphorella similis sp. nov.

from Switzerland, a close relative of the type species, Micro-

©ZFMK

In memoriam Dr. Hans Ulrich (1934-2018) 3

phorella praecox (Loew) (Diptera: Dolichopodidae: Parath-

alassiinae). Zootaxa 3489: 45-57

REFERENCES

Brown B V (2007) A further new genus of primitive phorid fly

(Diptera: Phoridae) from Baltic amber and its phylogenetic

implications. Contributions in Science 531: 1-14

Bonn zoological Bulletin 67 (1): 1-3

Disney R H L (1979) The British Metopina (Diptera: Phoridae)

with description of a new species. Zoological Journal of the

Linnean Society 67: 97-113

Gatt P (2003) New species and records of Microphorella Beck-

er (Diptera: Empidoidea, Dolichopodidae) from the Mediter-

ranean region. Revue suisse de Zoologie 110: 669-684

Schriftleitung (1994) Wir stellen vor. Heute: Sektion Dipter-

ologie, Leiter: Dr. Hans Ulrich. Tier und Museum (Bonn) 4

(1): 22-24

©ZFMK

BHL

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Bonn zoological Bulletin 67 (1): 5-13

May 2018

Loktak Lake, Manipur, revisited: A Ramsar site as the rotifer

(Rotifera: Eurotatoria) biodiversity hot-spot of the Indian sub-region

Bhushan Kumar Sharma"* & Sumita Sharma!

' Department of Zoology, North-Eastern Hill University, Shillong-793 022, Meghalaya, India

“Corresponding author. E-mail: profbksharma@gmail.com

Abstract. The plankton and semi-plankton samples collected at Loktak Lake in Manipur state of northeast India (NED), be-

tween February 2015 and January 2017, revealed 180 rotifer species belonging to 42 genera and 22 families. Our report raises

the total richness of the phylum known from this Ramsar site to 189 species and thus assigns this floodplain wetland the status

of the most species-rich rotifer hot-spot of the Indian sub-region. Two species are new to the Oriental region and 25 are new

records from Manipur. The rotifer fauna is characterized by a number of species of global (~18% of species) and regional

biogeographic (~15% of species) interest, high richness and common occurrence of ‘tropic centered’ Lecane, relative paucity

and scarceness of Brachionus, Filinia, Hexarthra and Conochilus species, and littoral-periphytic assemblages with a number

of small-sized species. The present study is interesting for the rotifer ecosystem diversity update of this well-sampled flood-

plain wetland of the Indian sub-region. We estimate 270+ Rotifera species from this lake system pending specific analysis of

periphytic, colonial, and benthic taxa as well as of likely cryptic diversity of certain species complexes.

Key words. Biogeography, ecosystem diversity, floodplain wetland, heterogeneity.

INTRODUCTION

Rotifera have been documented and described from dif-

ferent parts of India for more than a century but there is

still paucity of information on their ecosystem diversity

in different aquatic biotopes because of limited studies,

inadequate sampling, incomplete inventories and lack of

taxonomic expertise (Sharma & Sharma 2017). Never-

theless, our investigations over the last two decades have

highlighted the global importance of the floodplain lakes

(locally termed ‘beels’ or ‘pats’) of northeast India (NEI)

(Sharma & Sharma 2014a, 2014b, 2017) regarding Roti-

fera biodiversity and biogeography. Deepor beel, a Ram-

sar site located in this region, has been reported to be

one of the most rotifer speciose individual ecosystems of

India (Sharma & Sharma 2015). Sharma et al. (2016) ex-

tended the investigations to Loktak Lake — another Ram-

sar site — and provided a notable update on the rotifer

fauna reported earlier by Sharma (2009). The present in-

tensive sampling culminated in more biodiverse assem-

blage characterizing this floodplain wetland of NEI as a

rotifer hot-spot of the Indian sub-region. An inventory of

the rotifer species recorded till date from Loktak Lake is

presented. Remarks are made on salient features of the

composition and species richness of Rotifera, and on new

records and taxa of biogeographic importance. Various

interesting species reported in this study are illustrated.

Received: 07.11.2017

Accepted: 22.03.2018

MATERIAL & METHODS

Loktak Lake (93° 46’ — 93° 55’ E, 24° 25’ — 24° 42’ N;

area: 286 km’; max. depth: 4.58 m, mean depth: 2.07 m),

Bishnupur/Imphal districts of Manipur (NEI), is an inter-

esting floodplain lake system (Figs 1 A—C) characterized

by its floating mats of vegetation called “Phumd1” which

are inhabited by endangered brow-antlered deer (Rucer-

vus eldi eldi). This Ramsar site is reported to have a di-

verse assemblage (233 species) of aquatic macrophytes

(Tunginba Singh, 2013)

The qualitative plankton and semi-plankton samples

were collected between February 2015 and January

2017 at three sampling sites: Loktak A (93°45’56.3” E;

24°32713.5” N; alt. 726 m asl), Loktak B (93°477°58.1”

E; 24°30°39.1” N; alt. 714 m asl) and Loktak Barrage

(93°45’43.5” E; 24°32’46.9” N; alt. 718 m asl). In addi-

tion, others from different parts (pats) of Loktak Lake ba-

sin are analyzed. The samples were collected by towing a

nylobolt plankton net (#50 um mesh size) and preserved

in 5% formalin. All samples were screened with a Wild

stereoscopic binocular microscope; individual rotifers

were isolated and mounted in Polyvinyl alcohol-lacto-

phenol, and were observed with a Leica (DM 1000) ste-

reoscopic phase contrast microscope fitted with an image

analyzer. The rotifer taxa were identified using Koste

(1978), Segers (1995), Sharma (1978a, 1978b, 1983,

1998), Sharma & Sharma (1987, 1997, 1999, 2000, 2008,

2013), and Jersabek and Leitner (2013). The voucher col-

lections are in the holdings of the Department of Zoolo-

gy, North-Eastern Hill University, Shillong.

Corresponding editor: B. Huber

6 Bhushan Kumar Sharma & Sumita Sharma

NORTHEAST INDIA

Fig. 1A—C. A. Map of India showing northeast India; B. Map of northeast India indicating Manipur state and location of Loktak

Lake; C. Loktak Lake — a Ramsar site showing typical ‘Phumdi‘

Bonn zoological Bulletin 67 (1): 5-13 ©ZFMK

Loktak Lake — a rotifer biodiversity hot-spot a

RESULTS

A total of 180 species representing 42 genera and 22 fam-

ilies were identified in our collections from Loktak Lake.

Our study raises the total number of rotifers known from

this Ramsar site to 189 species. The following is a de-

tailed systematic list of the recorded taxa.

Systematic list of Rotifera

recorded from Loktak Lake, Manipur

Phylum: Rotifera

Class: Eurotatoria

Subclass: Monogononta

Order: Ploima

Family: Brachionidae

1. Anuraeopsis fissa (Gosse, 1851)

2. Brachionus angularis Gosse, 1851

3. B. bidentatus Anderson, 1889

4. B. calyciflorus Pallas, 1766

5. B. caudatus Barrois & Daday, 1894

6. B. diversicornis (Daday, 1883)**

7. B. durgae Dhanapathi, 1974

8. B. falcatus Zacharias, 1898

9. B. forficula Wierzejski, 1891**

10. B .kostei Shiel, 1983

Il. B. mirabilis Daday, 1897

12. B. quadridentatus (Hermann, 1783) s. lat

13. Keratella cochlearis Gosse, 1851

14. K. edmondsoni Ahlstrom, 1943**

15. K. lenzi Hauer, 1953

16. K. tecta (Gosse, 1851)

17. K. tropica (Apstein, 1907)

18. Platyias leloupi (Gillard, 1967)

19. P. quadricornis (Ehrenberg, 1832)

20. Plationus patulus (O.F. Muller, 1786)

Family: Epiphanidae

21. Epiphanes brachionus (Ehrenberg, 1837)

Family: Euchlanidae

22. Beauchampiella eudactylota (Gosse, 1886)

23. Dipleuchlanis propatula (Gosse, 1886)

24. Euchlanis dilatata Ehrenberg, 1832

25. E. incisa Carlin, 1939

26. E. meneta Myers, 1930

27. E. semicarinata Segers, 1993#

28. E. triquetra Ehrenberg, 1838

29. Tripleuchlanis plicata (Levander, 1894)

Family: Mytilinidae

30. Lophocharis salpina (Ehrenberg, 1834)

31. Mytilina acanthophora Hauer, 1938

32. M. brevispina (Ehrenberg, 1830)**

Bonn zoological Bulletin 67 (1): 5-13

33. M. bisulcata (Lucks, 1912)

34. M. lobata Pourriot, 1996

35. M. michelangellii Reid & Turner, 1988

36. M. ventralis (Ehrenberg, 1832)

Family: Trichotriidae

37. Macrochaetus danneelae Koste & Shiel, 1983

38. M. longipes (Myers, 1934)

39. M. sericus (Thorpe, 1893)

40. Trichotria tetractis (Ehrenberg, 1830)

41. Wolga spinifera (Western, 1894)

Family: Lepadellidae

42. Colurella adriatica (Ehrenberg, 1837)

43. C. obtusa (Gosse, 1886)

44. C. sulcata (Stenroos, 1898)

45. C. uncinata (O.F. Miller, 1773)

46. Lepadella acuminata (Ehrenberg, 1834)

47. L. apsicora Myers, 1934

48. L. apsida Harring, 1916

49. L. benjamini Harring, 1916

50. L. bicornis Vasisht & Battish, 1971

S51. L. biloba Hauer, 1958**

52. L. cristata (Rousselet, 1893)**

53. L. costatoides Segers, 1992

54. L. dactyliseta (Stenroos, 1898)

55. L. desmeti Segers & Chittapun, 2001

56. L. discoidea Segers, 1993

57. L. ehrenbergi (Perty, 1850)

58. L. eurysterna Myers, 1942

59. L. heterostyla (Murray, 1913)

60. L. lindaui Koste, 1981

61. L. neglecta Segers & Dumont, 1995*

62. L. minuta (Weber & Montet, 1918)

63. L. ovalis (O.F. Miller, 1786)

64. L. patella (O.F. Miller, 1773) s. lat

65. L. quadricarinata (Stenroos, 1898)

66. L. quinquecostata (Lucks, 1912)

67. L. rhomboides (Gosse, 1886)

68. L. triba Myers, 1934

69. L. triptera Ehrenberg, 1830

70. L. vandenbrandei Gillard, 1952

71. Squatinella mutica (Ehrenberg, 1832)

Family: Lecanidae

72. Lecane acanthinula (Hauer, 1938)#

73. L. aculeata (Jakubski, 1912)

74. L. aeganea Harring, 1914

75. L. arcula Harring, 1914

76. L. aspasia Myers, 1917

77. L. batillifer (Murray, 1913)**

78. L. blachei Berzins, 1973

79. L. bifastigata Hauer, 1938**

80. L. bifurca (Bryce, 1892)

81. L bulla (Gosse, 1851) s. lat

L. bulla diabolica (Hauer, 1936)

©ZFMK

8 Bhushan Kumar Sharma & Sumita Sharma

8&2. L. closterocerca (Schmarda, 1898)

83. L. crepida Harring, 1914

84. L. curvicornis (Murray, 1913) s. lat

85. L. decipiens (Murray, 1913)

86. L. dorysimilis Trinh Dang, Segers

& Sanoamuang, 2015**

87. L. doryssa Harring, 1914

88. L. elegans Harring, 1914

89. L. elongata Harring & Myers, 1926**

90. L. flexilis (Gosse, 1886)

91. L. furcata (Murray, 1913)

92. L. hamata (Stokes, 1896) s. lat

93. L. haliclysta Harring & Myers, 1926

94. L. hastata (Murray, 1913)**

95. L. hornemanni (Ehrenberg, 1834)

96. L. inermis (Bryce, 1892)

97. L. inopinata Harring & Myers, 1926

98. L. lateralis Sharma, 1978

99. L. latissima Yamamoto, 195**

100.L. leontina (Turner, 1892)

101.L. ludwigii (Eckstein, 1883)

102.L. luna (O.F. Muller, 1776) s. lat

103.L. lunaris (Ehrenberg, 1832)

104.L. marchantaria Koste & Robertson, 1983*

105.L. monostyla (Daday, 1897)

106.L. nitida (Murray, 1913)

107.L. niwati Segers, Kothetip & Sanoamuang, 2004

108.L. obtusa (Murray, 1913)

109.L. ohioensis (Herrick, 1885)

110. L. papuana (Murray, 1913)

111. L. ploenensis (Voigt, 1902)

112. L. pusilla Harring, 1914

113. L. pyriformis (Daday, 1905)**

114. L. quadridentata (Ehrenberg, 1832)

115. L. rhenana Hauer, 1929

116. L. rhytida Harring & Myers, 1926

117. L. ruttneri Hauer, 1938

118. L. signifera (Jennings, 1896)

119. L. simonneae Segers, 1993

120.L. solfatara (Hauer, 1938)#

121.L. stenroosi (Meissner, 1908)

122.L. superaculeata Sanoamuang & Segers, 1997**

123.L. syngenes (Hauer, 1938)**

124.L. tenuiseta Harring, 1914

125.L. thienemanni (Hauer, 1938)

126.L. undulata Hauer, 1938

127.L. unguitata (Fadeev, 1925)

128.L. ungulata (Gosse, 1887)

Family: Notommatidae

129. Cephalodella forficula (Ehrenberg, 1832)

130.C. gibba (Ehrenberg, 1832)

131.C. mucronata Harring & Myers, 1921

132.C. trigona (Rousselet, 1895)**

133. Monommata longiseta (O.F. Muller, 1786)

134. M. maculata (Harring & Myers, 1924)

Bonn zoological Bulletin 67 (1): 5—13

135. Monommata sp.#

136.Notommata spinata Koste & Shiel, 1991

Family: Scaridiidae

137. Scaridium longicaudum (O.F. Muller, 1786)

Family: Gastropodidae

138. Ascomorpha ecaudis Perty, 1850

139.A. ovalis (Bergendal, 1892)

Family: Trichocercidae

140. Trichocerca abilioi Segers & Sarma, 1993#

141.T. bicristata (Gosse, 1887)

142.T. capucina (Wierzejski & Zacharias, 1893)**

143.T. cylindrica (Imhof, 1891)

144.T: edmondsoni (Myers, 1936)

145.T. elongata (Gosse, 1886)

146.T. flagellata Hauer, 1938

147.T. insignis (Herrick, 1886)

148.T. hollaerti De Smet, 1990

149.T: longiseta (Schrank, 1802)

150.T: maior Hauer, 1936

151.T: pusilla (Jennings, 1903)**

152.T: rattus (O.F. Miller, 1786)

153.T. scipio (Gosse, 1886)

154.T. similis (Wierzejski, 1893)

155.T: sulcata (Jennings, 1894)**

156.T. tenuior (Gosse, 1886)

157.T. weberi (Jennings, 1903)

Family: Asplanchnidae

158. Asplanchna priodonta Gosse, 1850

Family: Synchaetidae

159. Ploesoma lenticulare Herrick, 1855

160. Polyarthra vulgaris Carlin, 1943

161.Synchaeta oblonga Ehrenberg, 1832**

162.8. pectinata Ehrenberg, 1832

Family: Dicranophoridae

163. Dicranophoroides caudatus (Ehrenberg, 1832)

164. Dicranophorus. forcipatus (O.F. Miller, 1786)

Order: Gnesiotrocha

Family: Flosculariidae

165. Floscularia ringens (Linnaeus, 1758)#

166. Sinantherina semibullata (Thorpe, 1893)

167.S. spinosa (Thorpe, 1893)

168.8. socialis (Linnaeus, 1758)

Family: Hexarthridae

169. Hexarthra mira (Hudson, 1871)**

Family: Conochilidae

170.Conochilus unicornis Rousselet, 1892

©ZFMK

Loktak Lake — a rotifer biodiversity hot-spot 9

Family: Testudinellidae

171.Testudinella amphora Hauer, 1938

172.T. brevicaudata Yamamoto, 1951

173.T. emarginula (Stenroos, 1898)

174.T. parva parva (Ternetz, 1892)

T. parva bidentata (Ternetz, 1892)**

175.T. patina (Hermann, 1783)

176.T. tridentata Smirnov, 1931

Family: Trochosphaeridae

177.Filinia brachiata (Rousselet, 1916)#

178.F: camasecla Myers, 1938

179.F: longiseta (Ehrenberg, 1834)

180.F: opoliensis (Zacharias, 1898)

181.F: saltator (Gosse, 1886)

182. Trochosphaera aequatorialis Semper, 1872

Family: Collothecidae

1&3.Collotheca ornata (Ehrenberg, 1832)**

Subclass: Digononta

Order: Bdelloidea

Family: Habrotrochidae

184. Habrotrocha angusticollis (Murray, 1905)#

Family: Philodinidae

185. Dissotrocha aculeata (Ehrenberg, 1832)**

186. Rotaria macroceros (Gosse, 1851)

187.R. neptunia (Ehrenberg, 1832)

1&8.R. rotatoria (Pallas, 1766)**

189.R. tardigrada (Ehrenberg, 1830)#

* New records from India; ** New records from Manipur; # not

observed in this study

Lepadella neglecta (Fig. 2) and Lecane marchantaria

(Fig. 3) are new records from India. Brachionus diver-

sicornis, B. forficula, Cephalodella trigona, Collotheca

ornata, Dissotrocha aculeata, Hexarthra mira, Keratel-

la edmondsoni (Fig. 4), Lecane batillifer (Fig. 5), L. bi-

fastigata (Fig. 6), L. dorysimilis (Fig. 7), L. elongata, L.

hastata, L. latissima (Fig. 8), L. pyriformis, L. superac-

uleata (Fig. 9), L. syngenes, Lepadella biloba, L. crista-

ta (Fig.10), Mytilina brevispina, R. rotatoria, Synchaeta

oblonga, Trichocerca capucina, T: pusilla, T. sulcata and

Testudinella parva bidentata (Fig. 11) are new records

from Manipur state of NEI. Lecanidae (57 species), Lep-

adellidae (30 species), Brachionidae (20 species), and

Trichocercidae (18 species) collectively formed 66.1%

(125 species) of the total rotifer species richness now

known from Loktak Lake. Euchlanidae and Notomma-

tidae included eight species each, while Testudinellidae

and Trochosphaeridae are represented by six species

each.

Bonn zoological Bulletin 67 (1): 5-13

DISCUSSION

Our report of 180 species characterizes the biodiverse

rotifer assemblages of Loktak Lake, raises the total spe-

cies richness known from this wetland to 189 species and

thus categorizes this Ramsar as the most species rich Ro-

tifera hot-spot of the Indian sub-region and one of the

globally most diverse rotifer biotopes. The results affirm

the hypothesis of Segers et al. (1993) on (sub) tropical

floodplains as the world’s richest rotifer habitats and also

endorse the speciose nature of the rotifers of the flood-

plain lakes of NEI (Sharma & Sharma 2014a, 2014b).

The total species richness comprises ~42% and ~76% of

Rotifera species known till date from India (Sharma &

Sharma 2017) and northeast India (BKS, unpublished),

respectively. Our inventory exceeds the highest Indian

report of 171 rotifer species (Sharma & Sharma 2015)

from Deepor beel — a Ramsar site and an important flood-

plain lake of NEI. This study marks a significant rich-

ness update on the taxon from Loktak basin compared

to earlier reports of 120 (Sharma 2009) and 152 species

(Sharma et al. 2016). The listing of 42 genera and 22

families affirms rich higher level diversity of Eurotatoria

as compared with 65 genera and 25 families of the phy-

lum known from India (Sharma & Sharma 2017). The

biodiverse Rotifera fauna is hypothesized to result from

micro-habitat diversity and environmental heterogeneity

of Loktak Lake while high richness in our plankton and

semi-plankton collections is hypothesized to result from

greater habitat diversification due to the influence of the

littoral vegetation (Green 1972; Serafim et al. 2003). We

also attribute the reported high richness to the ‘rotiferol-

ogist effect (cf. Fontaneto et al. 2012).

The Rotifera species richness of Loktak Lake con-

curs with the report of “All Taxa Biological Inventories

(ATBI)” for the rotifer assemblages of the tropical and

subtropical lakes, listing between 123 and 210 species

(Dumont & Segers 1996). Total richness is lower than

207, 230 and 252 species reported from the floodplains

of Africa (Segers et al. 1993), South America (Serafim

Jr. et al. 2003), and Australia (Shiel et al. 1998) while

it broadly corresponds with 184 examined species from

the Upper Parana floodplain (Bonecker et al. 1994, 1998,

2005; Lansac-Toha et al. 1997) of Brazil. On the other

hand, Loktak Rotifera is more diverse than the records of

114 species (Jose de Paggi 2001) from the Rio Pilcomayo

National Park (a Ramsar site), Argentina, 124 species

(Oguta lake) and 136 species (ly1-Efi lake) from the Ni-

ger delta (Segers et al. 1993) of Africa, 130 species from

Lake Guarana, Brazil (Bonecker et al. 1994), 106 taxa

from Thale-Noi Lake, a Ramsar site in Thailand (Segers

& Pholpunthin 1997), 104 species from Laguana Bufeos,

Bolivia (Segers et al. 1998), and 151 (Koste 1974) and

148 species from Rio Tapajos and Lago Camaleao (Koste

& Robertson 1983) of Brazil, respectively.

©ZFMK

Ries

Eat

‘2

Bonn zoological Bulletin 67 (1): 5-13

Bhushan Kumar Sharma & Sumita Sharma

Figs 2-11.

2. Lepadella neglecta Segers

& Dumont (ventral view); 3.

Lecane marchantaria Koste

& Robertson (ventral view);

4. Keratella edmondsoni

Ahlstrom (ventral view); 5.

Lecane_ batillifer (Murray)

(dorsal view); 6. Lecane

bifastigata Hauer (ventral

view); 7. Lecane dorysimilis

Trinh Dang, Segers & Sano-

amuang (ventral view); 8.

Lecane latissima Yamamo-

to (dorsal view); 9. Lecane

superaculeata Sanoamuang

& Segers (ventral view); 10.

Lepadella cristata (Rousse-

let) (ventral view); 11. Te-

studinella parva bidentata

(Ternetz) (ventral view).

Scales = 50um.

©ZFMK

Loktak Lake — a rotifer biodiversity hot-spot 1]

The two Neotropical species (Segers 2007) Lepadella

neglecta and Lecane marchantaria are new to the Ori-

ental Rotifera. The former was described (Segers & Du-

mont 1995) from Lobo (Broa) reservoir, Brazil; it was

confused in the past (Segers & Dumont loc cit.) with L.

quinquecostata. This remark deserved caution as L. ne-

glecta is characterized by its pyriform lorica, slightly nar-

rowed aperture, dorsum domed with seven low but con-

spicuous semi-longitudinal ridges, and posterior margin

projecting and with a minute median notch and thus dif-

fered distinctly in its morphology from the latter. Lecane

marchantaria is deemed to be confused (Segers 1995)

with the L. hamata complex; it is differentiated from the

latter by nearly coincident and broadly V-shaped head

aperture margins. Our collections from Loktak basin add

25 new records to the rotifer fauna of Manipur. Our col-

lections reveal important fractions of species of global

(~18% of species; 34 species) and regional biogeographic

interest (~15% of species; 27 species). The Australasian

Lecane batillifer;, the Neotropical Lepadella neglecta and

L. marchantaria, the Oriental Keratella edmondsoni, Le-

cane latissima and L. superaculeata, the Indo-Chinese

Lepadella dorysimilis, and Cephalodella trigona and

Lecane bifastigata are new to the list of Rotifera from

Loktak. Brachionus kostei, Dissotrocha aculeata, Eu-

chlanis semicarinata, Filinia camascela, Notommata

spinata, Lecane aeganea, L. aspasia, L. dorysimilis, L.

latissima, L. rhenana, L. rhytida, L. niwati, L. solfatara,

L. superaculeata, L. undulata, Lepadella desmeti, L. van-

denbrandei, Monommata maculata, Notommata spinata,

Testudinella amphora, T: brevicaudata, T: parva biden-

tata, Trichocerca abilioi, T: edmondsoni, T: hollaerti, T:

maior and T: sulcata are examples of species of region-

al distribution interest in the Indian Rotifera with their

distribution limited till date to NEI (Sharma & Sharma

2017).

Lecanidae > Lepadellidae > Brachionidae > Trichocer-

cidae collectively form a large fraction (~66% of species)

of the rotifer fauna of Loktak. The higher lecanid rich-

ness compares with the reports from the floodplains of

Africa (Segers et al. 1993, 1998; Green 2003), Argen-

tina (Jose de Paggi 2001), Brazil (Koste 1974; Koste &

Robertson 1983; Bozelli 1992; Bonecker et al. 1998;

Martinez et al. 2000; Serafim Jr. et al.2003; Bonecker et

al. 2005, 2009), Venezuela (Vasquez & Rey 1989), Thai-

land (Sanoamuang 1998), and India (Sharma & Sharma

2014a, 2014b; Sharma et al. 2017). Nevertheless, the rel-

ative paucity of the Brachionidae (20 species) in Loktak

basin is in contrast to the above listed studies and the

report from the Brahmaputra floodplains of NEI (Sharma

& Sharma 2014b).

Our results ascertain the collective importance (~52%

of species) of the littoral-periphytonic Lecane (57 spe-

cies) > Lepadella (25 species) > Trichocerca (18 spe-

cies). The consistency of the importance of these genera

Bonn zoological Bulletin 67 (1): 5-13

in the Loktak basin highlights the possibility of rules for

the periphytic rotifer assemblages as hypothesized by

Green (2003). The relative significance of these taxa con-

curs with the reports from the floodplains of Argentina

(Jose De Paggi 2001), Africa (Segers et al. 1993; Green

loc cit.), Brazil (Koste 1974; Koste & Robertson 1983;

Bonecker et al. 1998), Thailand (Segers & Pholpunthin

1997; Sanoamuang 1998), and Bolivia (Segers et al.

1998) as well as of Assam state of NEI (Sharma 2014;

Sharma & Sharma 2008, 2014a, 2017). High richness

of ‘tropic centered’ Lecane also concurred with several

reports on the tropical rotifer faunas, 1.e., Koste & Shiel

(1983), Dussart et al. (1984), Bozelli (1992), Bonecker et

al. (1994), Segers (1995), and Sharma & Sharma (2008,

2014a, 2017).

Loktak Rotifera indicates a number of small-sized

littoral-periphytonic species of Colurella, Lecane, Lep-

adella and Trichocerca, this feature is hypothesized to

result from predation influence of juvenile fish and inver-

tebrates (Baumgartner et al. 1997) though specific stud-

ies are needed to confirm this hypotheses. Our collec-

tions exhibit the paucity of planktonic rotifers and that of

Brachionus (10 spp.), Filinia, Hexarthra and Conochilus

Species in particular. The occurrence of fewer Brachio-

nus species corresponds with our reports from the flood-

plains of the Majuli River Island (Sharma 2014) and the

Dibru-Saikhowa Biosphere reserve (Sharma et al. 2017)

of upper Assam, NEI. Sharma et al. (2017) proposed the

L/B quotient based on Lecane/Brachionus species ratios

to characterize habitat variations of wetlands of lower

Assam, NEI. Based on overall richness of the two gen-

era, the L/B quotient for Loktak Lake indicates a value

of 5.7 thus ascertaining ‘wetland character’ of this Ram-

sar site; this is affirmed by the littoral-periphytic nature

of the lake system with a lack of permanent open-water

limnetic conditions.

The morphological variability observed in certain

species from Loktak Lake, namely Lepadella ovalis, L.

patella, Lecane bulla, L. curvicornis, L. hamata, L. leon-

tina, L. luna, L. lunaris, L. quadridentata, L. ungulata,

Testudinella emarginula, T. patina, T. tridentata and Pla-

tionus patulus needs attention for cryptic diversity anal-

ysis in light of some interesting studies on such species

complexes (Suatoni et al. 2006; Schroder & Walsh 2010;

Montero-Pau et al. 2011; Mills et al. 2017).

To sum up, the designation of Loktak Lake as a Roti-

fera hot-spot of the Indian sub-region, new records, and

species of global and regional biogeographic interest

highlight the biodiversity and ecosystem diversity im-

portance of this Ramsar site. Our intensive sampling and

the results justify this revisit of Loktak Lake with regards

its importance to biodiversity and biogeography of the

Indian Rotifera. We, however, estimate an occurrence of

270+ species of the phylum from this floodplain lake sys-

tem pending specific analysis of periphytic, colonial and

©ZFMK

| pea Bhushan Kumar Sharma & Sumita Sharma

benthic taxa, the rotifer-macrophytic associations relat-

ing to the characteristic “‘phumd1’, and analysis of cryptic

diversity of certain species complexes.

Acknowledgements. The senior author thanks the Head, De-

partment of Zoology, North-Eastern Hill University, Shillong

for laboratory facilities and various researchers and well-wish-

ers for untiring help in the field work. We thank our anonymous

peers for useful comments and suggestions. We wish to thank

Ralph S. Peters and Bernhard Huber (Bonn zoological Bulletin)

for useful suggestions during the peer review. The authors have

no conflict of interests.

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Bonn zoological Bulletin 67 (1): 15—24

May 2018

New records of snakes (Squamata: Serpentes)

from Hoa Binh Province, northwestern Vietnam

Truong Quang Nguyen'", Tan Van Nguyen '°, Cuong The Pham!’”, An Vinh Ong‘ & Thomas Ziegler*

' Institute of Ecology and Biological Resources, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet Road, Hanoi,

Vietnam

? Graduate University of Science and Technology, Vietnam Academy of Science and Technology, 18 Hoang Quoc Viet Road,

Hanoi, Vietnam

3 Save Vietnam's Wildlife, Cuc Phuong National Park, Ninh Binh Province, Vietnam

* Vinh University, 182 Le Duan Road, Vinh City, Nghe An Province, Vietnam

> AG Zoologischer Garten KéIn, Riehler Strasse 173, D-50735 Cologne, Germany

* Corresponding author. E-mail: nqt2@yahoo.com

Abstract. We report nine new records of snakes from Hoa Binh Province based on a reptile collection from Thuong

Tien, Hang Kia-Pa Co, Ngoc Son-Ngo Luong nature reserves, and Tan Lac District, comprising six species of Colubri-

dae (Dryocalamus davisonii, Euprepiophis mandarinus, Lycodon futsingensis, L. meridionalis, Sibynophis collaris and

Sinonatrix aequifasciata), one species of Pareatidae (Pareas hamptoni) and two species of Viperidae (Protobothrops mu-

crosquamatus and Trimeresurus gumprechti). In addition, we provide an updated list of 43 snake species from Hoa Binh

Province. The snake fauna of Hoa Binh contains some species of conservation concern with seven species listed in the

Governmental Decree No. 32/2006/ND-CP (2006), nine species listed in the Vietnam Red Data Book (2007), and three

species listed in the IUCN Red List (2018).

Key words. New records, snakes, taxonomy, Hoa Binh Province.

INTRODUCTION

In the recent checklist of the herpetofauna of Vietnam,

Nguyen et al. (2009) listed 192 species of snakes. Since

then 13 new country records, one new genus and 15 new

species of snakes have been described from Vietnam

(Ziegler & Nguyen 2010, Uetz et al. 2018). In Hoa Binh

Province, previous studies documented a total of 34 spe-

cies of snakes (Nguyen et al. 2009, Nguyen et al. 2010,

Ziegler et al. 2010, Luu et al. 2011). In this paper, we re-

port nine new records of snakes from Hoa Binh Province

based on newly collected specimens from Thuong Tien

Nature Reserve (Kim Boi District), Hang Kia-Pa Co Na-

ture Reserve (Mai Chau District), Ngoc Son-Ngo Luong

Nature Reserve (Tan Lac and Lac Son districts), and Tan

Lac District.

MATERIAL & METHODS

Field surveys were conducted in Thuong Tien Nature

Reserve (hereafter NR) by V.Q. Luu in March 2009; in

Hang Kia-Pa Co and Ngoc Son-Ngo Luong NR, in April,

May, September, and October 2014 and April 2015 by

T.Q. Nguyen. C.T. Pham, C.V. Hoang, H.N. Ngo, M.D.

Le, H.T. An (hereafter TQN et al.) and in Tan Lac District

in June 2016 by C.T. Pham, T.V. Nguyen, N.H. Nguyen

Received: 07.01.2018

Accepted: 22.03.2018

(hereafter CTP et al.). Specimens were collected by hand

or by using a snake hook between 8:00 and 23:00 hrs.

Most specimens were photographed in life. Specimens

were euthanized in a closed vessel with a piece of cotton

wool containing ethyl acetate (Simmons, 2002), fixed in

85% ethanol and subsequently stored in 70% ethanol.

Tissue samples of some species were kept separately in

90% ethanol. Specimens were deposited in the collec-

tions of the Institute of Ecology and Biological Resourc-

es (EBR), Vietnam Academy of Science and Technolo-

gy, Hanoi, Vietnam.

Taxonomic identifications of the specimens were made

based on the following literature: Smith (1943), Taylor

(1965), David et al. (2002), Vogel et al. (2004), Gum-

precht et al. (2004), Orlov et al. (2004, 2011), Kim &

Oh (2006), Stuart & Heatwole (2008), Yang et al. (2008),

Vogel et al. (2009), Hecht et al. (2013), Luu et al. (2013a,

b), Le et al. (2015), Nguyen et al. (2011, 2014), Ziegler

et al. (2007, 2014), Nemes et al. (2013), Nguyen et al.

(2016), Vassilieva et al. (2016), and Pham et al. (2017).

For common names, we followed Nguyen et al. (2009)

and Uetz et al. (2018). Abbreviations used for morphom-

etry are as follows: SVL (snout-vent length): from tip of

snout to anterior margin of cloaca; TaL (tail length): from

posterior margin of cloaca to tip of tail.

Identification of sex was performed by dissection (in-

spection of gonads and inspection of presence of hemi-

Corresponding editor: W. Bohme

16 Truong Quang Nguyen et al.

East Sea

(South China Sea)

Elevation (m)

en ‘y 3

. Po

—< . : i ee ws

Gulf of sonepitlnds ae

7 anc

0 5 Thailand

j a ks

Kilometers 150 300 Kilometers

(TS Tia

Fig. 1. Map showing the survey site (black square) in Hoa Binh Province, Vietnam.

Bonn zoological Bulletin 67 (1): 15-24 ©ZFMK

New records of snakes from Hoa Binh Province bz

penes). Measurements were taken after preservation

with a measuring tape. The number of ventral scales was

counted according to Dowling (1951). The numbers of

dorsal scale rows are given at one head length behind

head, at midbody, and at one head length before vent,

respectively. Scalation was studied by using a binocular.

Bilateral scale counts were given as left/right.

RESULTS

Taxonomic accounts

Family Colubridae

Dryocalamus davisonii (Blanford, 1878)

Bridle Snake / Ran dé (Fig. 2a)

Specimen examined (n = 1): IEBR 4058 (adult female)

collected by TON et al. on 22 May 2014, in Ngoc

Son-Ngo Luong NR, Lac Son District (20°24.909’N,

105°19.102’E; elevation 350 m above sea level [hereaf-

ter asl.]).

Description: Morphological characters of the specimen

from Hoa Binh Province agreed well with the descrip-

tions of Smith (1943), Taylor (1965), Das (2010), Orlov

et al. (2011), and Vassilieva et al. (2015). SVL 587.7 mm,

TaL 191.4 mm. Head distinct from neck; nasal undivid-

ed; loreal 1/1, touching the eye; preocular absent; postoc-

ulars 2/2; anterior temporal 1/1; posterior temporals 2/2;

supralabials 7/7, third and fourth entering orbit; infralabi-

als 8/8: dorsal scale rows 13-13-13, all smooth; ventrals

239 (+ 1 preventral); cloacal undivided; subcaudals 109,

divided.

Coloration in life: Dorsum black with 45 white cross-

bars and 26 other bars on tail; white cross-bar narrow-

er at posterior part of body; head dark brown with pale

cream-colored elongated spots on each side, from pari-

etal shield to supraorbital; venter cream anteriorly, grey

posteriorly.

Ecological notes: The specimen was found at 22:00 on

the ground. The surrounding habitat was secondary forest

composed of medium and small hardwoods and shrub.

Distribution: In Vietnam, this species has been record-

ed from Thanh Hoa Province southwards to Kien Giang

Province. This is the first record of D. davisonii from

Ngoc Son-Ngo Luong NR as well as from Hoa Binh

Province and the Northwest of Vietnam. Elsewhere, the

species has been reported from Myanmar, Laos, Thailand

and Cambodia (Nguyen et al. 2009, Orlov et al. 2011).

Euprepiophis mandarinus (Cantor, 1842)

Mandarin Ratsnake / Ran soc quan (Fig. 2b)

Specimen examined (n = 1): Photographic record only,

by T.Q. Nguyen on 12 April 2014, in Hang Kia-Pa Co

Bonn zoological Bulletin 67 (1): 15—24

NR, Mai Chau District (20°43.445’N, 104°53.310’E; ele-

vation 1381 m asl.).

Description: Morphological characters of the specimen

from Hoa Binh Province agreed well with the descrip-

tions of Smith (1943), Yang & Rao (2008), Das (2010),

and Ziegler et al. (2014). The photographed specimen

resembles Euprepiophis mandarinus in the following

characters: Dorsum brown above, with a series of large

black, diamond shaped marks that enclose oval, rounded

or squarish yellow spots; black marks edged by narrow

yellow margins; head with three black marking-bands

across snout, a crescent V-shaped mark through the eye

and divided into two stripes, and a forward pointing

shaped mark on the neck.

Ecological notes: The specimen was found at 21:30

on the ground, on the banks of a rocky stream. The sur-

rounding habitat was secondary forest composed of me-

dium and small hardwoods and shrub.

Distribution: In Vietnam, this species has been record-

ed from Lai Chau and Ha Giang provinces in the North

southwards to Dak Lak Province. This is the first record

of E. mandarinus from Hang Kia-Pa Co NR as well as

from Hoa Binh Province. Elsewhere, the species has

been reported fom India, China, Tatwan, Myanmar and

Laos (Nguyen et al. 2009, Ziegler et al. 2014).

Lycodon futsingensis (Pope, 1928)

Futsing Wolf Snake / Ran khuyét fut-sing (Fig. 2c)

Specimen examined (n = 1): IEBR 4171 Guvenile) col-

lected by TQN et al. on 14 April 2014, in Hang Kia-Pa

Co NR, Mai Chau District (20°43.667’N, 104°51.823’E;

elevation 901 m asl.).

Description: Morphological characters of the specimen

from Hoa Binh Province agreed well with the descrip-

tions of Vogel et al. (2009), Hecht et al. (2013), Luu et

al. (2013), Nguyen et al. (2014), Nguyen et al. (2016),

and Pham et al. (2017). SVL 265.9 mm, TaL 75.7 mm.

Head distinct from neck; rostral broader than high; in-

ternasal not in contact with loreal; nasal divided; loreal

1/1, small, not touching the eye; preocular 1/1; subocular

absent; postoculars 2/2; anterior temporals 2/2; posteri-

or temporals 2/2; supralabials 8/8, third to fifth entering

orbit; infralabials 10/10; dorsal scale rows 17-17-15, all

smooth; ventrals 202 (+ 1 preventral); cloacal undivided;

subcaudals 90, divided.

Coloration in life: Dorsum pale brownish grey with 27

brown rings on body and 8 rings on the tail; head dark

brown with a large light band, from eye to neck; the rings

wider at base, the first one starting at ventral scale 15, at

its base comprising 6 ventrals in width and dorsally com-

prising 2 dorsal scales; venter cream, with dark marbling,

dark grey posteriorly.

Ecological notes: The specimen was found at 21:00 on

a forest path. The surrounding habitat was secondary for-

est composed of small hardwoods, liane and shrub.

©ZFMK

18 Truong Quang Nguyen et al.

Fig. 2. a) Dryocalamus davisonii (IEBR 4058), b) Euprepiophis mandarinus, ¢) Lycodon futsingensis (IEBR 4171), d) Lycodon

meridionalis (IEBR 4151), e) Sinonatrix aequifasciata (IEBR 4226), and f) Sibynophis collaris (IEBR 4224) from Hoa Binh Proy-

ince, Vietnam.

Distribution. In Vietnam, this species was reported

from Lao Cai Province in the North southwards to Da

Nang City. This is the first record of L. futsingensis from

Hang Kia-Pa Co NR as well as from Hoa Binh Province.

Elsewhere, the species has been reported from China and

Laos (Nguyen et al. 2009, Luu et al. 2013, Nguyen et al.

2016, Pham et al. 2017).

Bonn zoological Bulletin 67 (1): 15-24

Lycodon meridionalis (Bourret, 1935)

Southern Big-tooth Snake / Ran léch dau kim tuyén (Fig.

2d)

Specimens examined (n = 4): IEBR 4050, 4051 (two

adult males), and IEBR 4156 (adult female) collected by

TQN et al. in April 2014, in Hang Kia-Pa Co NR, Mai

Chau District (20°25.072’N, 105°19.102’E; elevation

©ZFMK

New records of snakes from Hoa Binh Province 19

300 m asl.); and IEBR 4154 (adult male) collected by

CTP etal. on 19 June 2016, in Ngoc Son-Ngo Luong NR,

Lac Son District (20°28.076’N, 105°18.216’E; elevation

733 m asl.)

Description: Morphological characters of the spec-

imens from Hoa Binh Province agreed well with the

descriptions of Orlov & Ryabov (2004), Hecht et al.

(2013), Ziegler et al. (2014), and Nguyen et al. (2016).

SVL: 929.5—1341.7 mm in males (n = 3), 873.1 mm in

the single female (n= 1), TaL: 246.7—364.8 mm in males

(n = 3), 258.7 mm in the single female (n = 1). Head

distinct from neck; internasals not in contact with loreal;

nasal divided; loreal 1/1, not touching the eye; preocular

1/1; subocular absent; postoculars 2/2; anterior tempo-

rals 2/2; posterior temporals 3/3; supralabials 8/8, third

to fifth entering orbit; infralabials 10/10; dorsal scale

rows 17-17-15, strongly keeled except 5 outermost rows

smooth, outer dorsal scales enlarged; ventrals 242—257

(+ 2-3 preventrals); cloacal undivided; subcaudals 100—

118, divided.

Coloration in life: Dorsum black with 100-119 narrow

yellow cross-bars on body and 31-41 on tail, bifurcated

on the sides, enclosing dark spots; head black with sym-

metrical light markings, the most conspicuous being the

one running from the eye to the margin of the snout and

another stretching from the hind margin of the parietals:

venter light yellow.

Ecological notes: The specimens were found between

19:00 and 22:30, on the ground or on rocks. The sur-

rounding habitat was secondary forest composed of me-

dium and small hardwoods, liane and shrub.

Distribution: In Vietnam, this species has been report-

ed from Lao Cai and Ha Giang provinces in the North

southwards to Thanh Hoa Province. This is the first re-

cord of L. meridionalis from Hang Kia-Pa Co and Ngoc

Son-Ngo Luong NRs as well as from Hoa Binh Province.

Elsewhere, the species has been reported from China and

Laos (Nguyen et al. 2009, Ziegler et al. 2014, Nguyen et

al. 2016).

Sibynophis collaris (Gray, 1853)

Common Many-tooth Snake / Ran rong c6 den (Fig. 2e)

Specimen examined (n = 1): IEBR 4224 (adult female)

collected by V.Q. Luu in March 2009, in Thuong Tien

NR, Kim Boi District (near 20°36’N, 105°29’E; elevation

608 m asl.).

Description: Morphological characters of the specimen

from Hoa Binh Province agreed well with the descrip-

tions of Smith (1943), Taylor (1965), Kim & Oh (2006),

Nemes et al. (2013), and Vassilieva et al. (2015). SVL

324 mm, TaL 184 mm. Head distinct from neck; inter-

nasal not in contact with loreal; nasal divided; loreal 1/1,

small, not touching the eye; preocular 1/1; subocular

absent; postoculars 2/2; anterior temporal 1/1; posterior

temporals 2/2; supralabials 10/10, fourth to sixth enter-

Bonn zoological Bulletin 67 (1): 15—24

ing orbit; infralabials 9/9; dorsal scale rows 17-17-17,

all smooth; ventrals 168 (+ 1 preventral); cloacal divided:

subcaudals 120, divided.

Coloration in preservative: Dorsum brown, with a ver-

tebral series of small black spots, light dorsolateral lines

mostly present; head black with a black stripe running

from neck to the back of the head; a white stripe along

supralabials to the neck; venter yellow, each ventral with

a lateral dark spot.

Distribution. In Vietnam, this species has been report-

ed from Dien Bien and Son La provinces in the North

southwards to Lam Dong and Dong Nai provinces. This

is the first record of S. collaris from Thuong Tien NR as

well as from Hoa Binh Province. Elsewhere, the species

has been reported from India, Nepal, China, Taiwan, Ko-

rea, Laos, Thailand, Cambodia, and Malaysia (Kim & Oh

2006, Nguyen et al. 2009, Nemes et al. 2013, Vassilieva

et al. 2015).

Sinonatrix aequifasciata (Barbour, 1908)

Asiatic Water Snake / Ran hoa can van d6ém (Fig. 2f)

Specimens examined (n = 2). IEBR 4225 (adult male)

and IEBR 4226 (adult female) collected by TQN et al.

on 17 April 2015, in Ngoc Son-Ngo Luong NR, Lac Son

District (20°26.671’N, 105°16.139’E; elevation 250 m

asl.).

Description: Morphological characters of the spec-

imens from Hoa Binh Province agreed well with the

descriptions of Vogel et al. (2004), Stuart & Heatwole

(2008), Hecht et al. (2013), and Le et al. (2015). IEBR

4225: SVL 666.9 mm, TaL 208.3 mm; IEBR 4226: SVL

729.1 mm, TaL 236.1 mm. Head elongated, indistinct

from neck; nuchal groove distinct; loreal present; preoc-

ulars 1/1 or 2/2; postoculars 3/4 or 3/5; anterior tempo-

rals 2/2, posterior temporals 2/3 or 3/3; supralabials 9/9,

the fifth entering orbit, the seventh largest; infralabials

10/10; dorsal scale rows 19-19-17, strongly keeled; ven-

trals 144-153 (+ 2-3 preventrals); cloacal divided; sub-

caudals 70-74, divided.

Coloration in life: Dorsal surface with 21 black dou-

ble-bands on body, flanks with dark markings, in X-shape

and 10-12 on tail; interspaces with brownish tinge on

each band, narrower than the dark bars; venter cream

with black markings.

Ecological notes: The specimens were found between

19:00 and 21:30, on branches of trees, about 1.0—-1.5 m

above the ground, on the banks of a rocky stream. The

surrounding habitat was secondary forest composed of

medium and small hardwoods, liane and shrub.

Distribution: In Vietnam, this species has been reported

from Lao Cai and Ha Giang provinces in the North south-

wards to Nghe An and Ha Tinh provinces. This is the first

record of S. aequifasciata from Ngoc Son-Ngo Luong

NR as well as from Hoa Binh Province. Elsewhere, the

species has been reported from China and Laos (Stuart

©ZFMK

20 Truong Quang Nguyen et al.

et al. 2008, Nguyen et al. 2009, Hecth et al. 2013, Le et

al. 2015).

Family Pareatidae

Pareas hamptoni (Boulenger, 1905)

Hampton’s Slug Snake / Ran h6 may ham-ton (Fig. 3a)

Specimens examined (n = 2): IEBR 4227, 4228 (adult

males) collected by TQN et al. in October 2014, in Ngoc

Son-Ngo Luong NR, Lac Son District (20026.862’N,

105020.144’E; elevation 553 m asl.)

Description: Morphological characters of the speci-

mens from Hoa Binh Province agreed well with the de-

scriptions of Smith (1943), Taylor (1965), Ziegler et al.

(2007), Nguyen et al. (2011), and Nemes et al. (2013).

SVL 363-524 mm, TaL 108-179 mm (n = 2). Body

strongly compressed; head distinct from neck; nasal un-

divided; loreal 1/1, touching the eye; preocular 1/1; pos-

toculars 1/1; subocular 1, long and slender, separating

the eye from the labials; anterior temporal 1/1; posterior

temporals 2/2; supralabials 7/7, third to fifth below the

eye, seventh very long; infralabials 8/8; mental groove

absent; dorsal scale rows 15-15-15, all smooth except

posterior upper dorsal scales slightly keeled, anterior ver-

tebral scales slightly enlarged: ventrals 199-202 (+ 1 pre-

ventral); cloacal undivided; subcaudals 98—100, divided.

Coloration in life: Light brown dorsally, with dorso-

lateral rows of alternating spots, forming a zigzag line,

spots absent at the margin of the ventrals.

Ecological notes: The specimens were found between

19:00 and 22:30 on branches of trees, about 1.5—2.0 m

above the ground, on forest paths. The surrounding habi-

tat was secondary forest composed of medium and small

hardwoods, liane and shrub

Distribution: In Vietnam, this species has been reported

from Lao Cai and Ha Giang provinces in the North south-

wards to Lam Dong and Dong Nai provinces. This is the

first record of P. hamptoni from Ngoc Son-Ngo Luong

NR as well as from Hoa Binh Province. Elsewhere, the

species has been reported from China, Myanmar, Laos,

and Cambodia (Nguyen et al. 2009).

Family Viperidae

Protobothrops mucrosquamatus (Cantor, 1839)

Brown spotted pitviper / Ran luc cuom (Fig. 3b, c)

Specimens examined (n = 2): IEBR 4230 (adult male)

collected by C.V. Hoang in Ngoc Son-Ngo Luong NR,

Lac Son District (20°25.034’N, 105°23.107’E; elevation

440 m asl.) and IEBR 4231 (adult female) collected by

C.T. Pham et al. on 8 June 2016, in Thanh Hoi Com-

mune, Tan Lac District (20°34.865’N, 105°19.731’E; el-

evation 170 m asl.).

Bonn zoological Bulletin 67 (1): 15-24

Description: Morphological characters of the speci-

mens from Hoa Binh agreed well with the description

of Stuart & Heatwole (2008), Nguyen et al. (2011), Luu

et al. (2013), and Nemes et al. (2013). IEBR 4230: SVL

544.0 mm, TaL 129 mm; IEBR 4231: SVL 605.3 mm,

TaL 134.0 mm. Hemipenes short and thick. Head trian-

gular, clearly distinct from the neck; nasal undivided; in-

ternasals separated from each other by three scales; two

small scales between the nasal and the shield bordering

the anterior region of the loreal pit; postoculars 2/2; su-

pralabials 8/8 or 11/11, the first supralabial completely

separated from the nasal, third supralabial large, in con-

tact with the subocular, fourth and fifth supralabials sep-

arated from the subocular by two scales; temporals small;

infralabials 13/14, the first pair in contact with each oth-

er, the first three pairs in contact with the chin shields;

dorsal scale rows 23(25)—23-17(21), rhomboid, strong-

ly keeled throughout but smooth on the first outer row;

ventrals 203-214 (+ 2 preventrals); cloacal undivided;

subcaudals 88-97, divided.

Coloration in life: Dorsal head brown, paler below;

dorsum greyish brown, with a series of large brown, dark-

edged spots; a dark brown line from the eye to the angle

of the mouth, edged in black; ventral surface brownish

with white blotches; dorsal tail light brown, with a series

of conspicuous black spots.

Ecological notes: The specimens were found between

19:00 and 22:30, on forest paths. The surrounding habi-

tat was secondary forest composed of medium and small

hardwoods, liane and shrub.

Distribution: In Vietnam, this species has been reported

from Lao Cai and Ha Giang provinces in the North south-

wards to Kon Tum and Gia Lai provinces. This is the first

record of P. mucrosquamatus from Ngoc Son-Ngo Luong

NRas well as from Hoa Binh Province. Elsewhere, the

species has been reported from India, Bangladesh, China,

Taiwan, and Myanmar (Nguyen et al. 2009, Luu et al.

2013, Nemes et al. 2013).

Trimeresurus gumprechti David, Vogel, Pauwels &

Vidal, 2002

Gumprecht’s green pitviper / Ran luc gum-p-ret (Fig. 3d)

Specimen examined (n = 1): IEBR 3918 (subadult male)

collected by TQN etal. on 12 April 2014, in Hang Kia-Pa

Co NR, Mai Chau District (20°44.184’N; 104°53.362’E,

elevation 1201 m asl.)

Description: Morphological characters of the specimen

from Hoa Binh agreed well with the description of David

et al. (2002). SVL 441.75 mm, TaL 100 mm. Hemipenes

short and thick with spines. Head triangular, clearly dis-

tinct from the neck; rostral visible from above, triangular;

nasal undivided; internasals separated from each other by

a scale; two small scales between the nasal and the shield

bordering the anterior region of the loreal pit; postocu-

lars 2/2; supralabials 10/10, the first separated from the

©ZFMK

New records of snakes from Hoa Binh Province Di

Fig. 3. a) Pareas hamptoni (IEBR 4227), b) Protobothrops mucrosquamatus (male) (IEBR 4230) and c) female TEBR 4231), and

d) Trimeresurus gumprechti (IEBR 3918) from Hoa Binh Province, Vietnam..

nasal, third large, in contact with subocular, fourth and

fifth separated from subocular by a small scale; tempo-

rals small; infralabials 13/12, the first pair in contact with

each other, the first three pairs in contact with the chin

shields; dorsal scale rows 23—21—15, rhomboid, strongly

keeled throughout but smooth on the outermost row; ven-

trals 160 (+ 4 preventrals); cloacal undivided; subcaudals

70, divided.

Coloration in life: Dorsal and ventral surface green

with a white ventrolateral stripe, edged in red below;

lateral head with a white postocular streak, edged in red

below; tail green with upper part of posterior half rusty

red; eyes red.

Ecological notes: The specimen was found at 21:00 on

tree branches near a small stream, approximately 0.2 m

above the ground. The surrounding habitat was second-

ary forest composed of medium and small hardwoods,

liane and shrub. A tree frog (Kurixalus sp.) and a water

Bonn zoological Bulletin 67 (1): 15-24

skink (Zropidophorus sp.) were found in the stomach of

this specimen.

Distribution. In Vietnam, this species has been report-

ed from Lai Chau and Lao Cai provinces (Nguyen et al.

2009). This is the first record of 7? gumprechti for Hang

Kia-Pa Co NR as well as for Hoa Binh Province. Else-

where, the species has been reported from China, Myan-

mar, Laos, and Thailand (David et al. 2004, Nguyen et

al. 2009).

DISCUSSION

Our new records of nine snake species bring the total

number of snake species in Hoa Binh Province to 43

(Table 1). The most diverse family is Colubridae with

27 recorded species, followed by Elapidae (4 species)

and Viperidae (4 species). The snake fauna of Hoa Binh

©ZFMK

22 Truong Quang Nguyen et al.

Table 1. List of snake species recorded from Hoa Binh Province, Vietnam. Data sources: 1: Nguyen et al. (2009), 2: Nguyen et al.

(2010), 3: Ziegler et al. (2010), 4: Luu (2011), 5: This study. Decree 32 (2006) = Governmental Decree No 32/2006/ND-CP dated

on 30 March 2006 by the Government of Vietnam on the management of endangered wild flora and fauna. Group IB: prohibited

exploitation and use for commercial purpose and Group IIB: limited exploitation and use for commercial purpose; RBVN (2007)

= Vietnam Red Data Book. Part I. Animals. Descriptions of nationally endangered species of wild animals. CR = Critically Endan-

gered, EN = Endangered, VU = Vulnerable; IUCN (2018) = The IUCN Red List of Threatened Species. CR = Critically Endan-

gered, EN = Endangered, VU = Vulnerable, LR/nt = Lower Risk/Near Threatened, * new provincial record.

Species name Previous IUCN RBVN Decree 32

record (2017) (2007) (2006)

Pythonidae

Python bivittatus (Kuhl, 1820) 1 VU CR IB

Xenopeltidae

—

Xenopeltis unicolor Reinwardt, 1827

Colubridae

Ahaetulla prasina (Boie, 1827)

Amphiesmoides ornaticeps (Werner, 1924)

Amphiesma stolatum (Linnaeus, 1758)

Boiga kraepelini Stejneger, 1902

Boiga multomaculata (Boie, 1827)

Calamaria pavimentata Duméril, Bibron & Dumeéril, 1854

Calamaria septentrionalis Boulenger, 1890

Coelognathus radiatus (Boie, 1827)

Cyclophiops multicinctus (Roux, 1907)

Dendrelaphis pictus (Gmelin, 1789)

Dryocalamus davisonii (Blanford, 1878)*

Elaphe moellendorffi (Boettger, 1886)

Euprepiophis mandarinus (Cantor, 1842)*

Lycodon futsingensis (Pope, 1928)*

Lycodon meridionalis (Bourret, 1935)*

Oligodon taeniatus (Gunther, 1861)

Opisthotropis lateralis Boulenger, 1903

Plagiopholis nuchalis (Boulenger, 1893)

Ptyas korros (Schlegel, 1837)

Ptyas mucosa (Linnaeus, 1758)

Rhabdophis chrysargos (Schlegel, 1837)

Rhabdophis subminiatus (Schlegel, 1837)

Sibynophis chinensis (Gunther, 1889)

Sibynophis collaris (Gray, 1853)*

Sinonatrix aequifasciata (Barbour, 1908)*

Sinonatrix percarinata (Boulenger, 1899)

Xenochrophis flavipunctatus (Hallowell, 1860)

aé

EN IIB

mS ee Bh NN NN WN BR OR ROD RN

Elapidae

Bungarus fasciatus (Schneider, 1801)

1 EN IB

Bungarus multicinctus Blyth, 1861 i

IIB

VU EN IIB

VU CR IB

Naja atra Cantor, 1842

Ophiophagus hannah (Cantor, 1836)

Homalopsidae

Hypsiscopus plumbea (Boie, 1827) |

Myrrophis chinensis (Gray, 1842) eae

Lamprophiidae

Psammodynastes pulverulentus (Boie, 1827)

—

Pareatidae

Pareas hamptoni (Boulenger, 1905)* 5

Pareas macularius Theobald, 1868 Le

Pareas margaritophorus (Jan, 1866) ]

Viperidae

Protobothrops mucrosquamatus (Cantor, 1839)*

Trimeresurus albolabris (Gray, 1842)

Trimeresurus gumprechti David, Vogel, Pauwels & Vidal, 2002*

Trimeresurus stejnegeri Schmidt, 1925

= nae Nn

Province also contains several species of conservation concern. Three species are listed in the IUCN Red List

Bonn zoological Bulletin 67 (1): 15-24 ©ZFMK

New records of snakes from Hoa Binh Province 23

(2018): Python bivittatus, Naja atra and Ophiophagus

hannah, nine species are listed in the Red Data Book

of Vietnam (2007): Python bivittatus, Coelognathus ra-

diatus, Elaphe moellendorffi, Euprepiophis mandarinus,

Ptyas korros, P. mucosa, Bungarus fasciatus, Naja atra,

and Ophiophagus hannah, seven species are listed in

the Vietnam Governmental Decree No. 32/2006/ND-CP

(2006): Python bivittatus, Coelognathus radiatus, Ptyas

mucosa, Bungarus fasciatus, B. multicinctus, Naja atra,

and Ophiophagus hannah (see Table 1).

Acknowledgements. We are grateful to the directorates of the

Forest Protection Department of Hoa Binh Province for their

support of our field work and issuing relevant permits. We

thank V.Q. Luu, H.T. An, C.V. Hoang, H.N. Ngo, M.D. Le,

N.H. Nguyen (Hano1) for their assistance in the field. We thank

to E. Sterling (New York) and K. Koy (Berkeley) for providing

the map. For the fruitful cooperation within joint amphibian

projects we cordially thank S.V. Nguyen (IEBR, Hanoi) and T.

Pagel and C. Landsberg (Cologne Zoo). This research is sup-

ported by the National Foundation for Science and Technology

Development (NAFOSTED, Grant No. 106.05-2017.329).

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Bonn zoological Bulletin 67 (1): 25-36

May 2018

Taxonomic reassessment of the Common Indian Wolf Snakes Lycodon aulicus

(Linnaeus, 1758) complex (Squamata: Serpentes: Colubridae)

Sumaithangi Rajagopalan Ganesh' & Gernot Vogel”

‘Chennai Snake Park, Chennai - 600 022, Tamil Nadu, India

Society for Southeast Asian Herpetology, Im Sand 3, D-69115 Heidelberg, Germany

* Corresponding author. E-mail: gernot.vogel@t-online.de

Abstract. We studied the population systematics of the group of the Common Indian wolf snake (Lycodon aulicus s.

lat.) based on a series of specimens from throughout most of their geographic range. Two discrete species-groups could

be discerned based on head dimensions, collar band pattern, hemipenal morphology and frontal-preocular-prefrontal-su-

proacular scale contact configurations (with outliers). The first one contains specimens agreeing with the morphology

of the name-bearing type of Lycodon aulicus, the other includes specimens agreeing with the morphology of Lycodon

anamallensis (so far within the synonymy of L. aulicus), which is here revalidated at species-level. We formally report the

presence of the presumed Sri Lankan endemic Lycodon osmanhilli group taxa in the Indian peninsula and we synonymise

L. osmanhilli with the senior nomen L. anamallensis which is based on an Indian specimen. Our series of specimens show

clear and sometimes non-overlapping geographical variations in ventral, subcaudal scale counts and relative tail lengths

within both species.

Key words. Collar band, head dimension, Indian peninsula, Sri Lanka, Wolf Snakes.

INTRODUCTION

The common Indian wolf snake Lycodon aulicus (Lin-

naeus, 1758) is anon-venomous, mainly nocturnal, ovip-

arous colubrid snake found in the Indian subcontinent

(Whitaker & Captain 2004). It is one of the commonest

and most ‘well-known’ snakes in tropical Asia. It was

described by Carolus Linnaeus in his Systema Natu-

rae. Subsequently, Patrick Russell, the “father of Indian

Ophiology”, included this species in his treatise (Russell

1796). It is understood to be widespread, human-com-

mensally and common in almost all herpetological litera-

ture (e.g., Whitaker & Captain 2004). But yet, as often the

case with such ‘well-known’ South Asian snakes like the

Rock Python Python molurus (see Wulf & O’ Shea 2010),

the Spectacled Cobra Naja naja (see Wister 1998a), the

Russell’s Viper Daboia russelii (see Wister 1998 b), the

Bronzeback Tree Snake Dendrelaphis tristis (see Vogel

& Van Rooijen 2009) and the Keelbacks Xenochrophis

piscator and Amphiesma stolatum (see Vogel & David

2012; Guo et al. 2014), the taxonomy of Lycodon aulicus

is still far from being resolved and the variation is not

well known.

This species was originally described based on the ho-

lotype NHR Lin-21 (formerly MAFR), a 250 mm long

specimen (Mus. Drottn.) supposed to come from “Amer-

ica”. The type specimen is still extant in the Royal Mu-

seum of Stockholm, formerly the Museum Adolphi Frid-

erici. The type locality was later proved to be in error

and was corrected to ‘India’ by Kramer (1977). Laurenti

(1768) allocated this species to the genus Natrix, as Na-

Received: 21.12.2017

Accepted: 22.03.2018

trix aulica. Dumeril et al. (1854) transferred Natrix auli-

ca to the genus Lycodon, as Lycodon aulicum.

Cantor (1839) described Lycodon subfuscus based on

a single specimen from Bengal, in north-eastern India.

Cantor (1839) also described Lycodon atropurpureus

based on a single specimen from “Mergut’, now Myeik,

in southern Myanmar. Gunther (1864) described Lycodon

anamallensis based on a single specimen from the Ana-

mallay Hills of the Western Ghats, in peninsular India.

Later Wall (1909) described the subspecies Lycodon au-

licus oligozonatus based on specimens from Cannanore

(in Malabar Coast) and Bellary (in Deccan plateau), in

southern India. All these four nomina were synonymised

by Smith (1943) with Lycodon aulicus (Linnaeus, 1758).

Taylor (1950) described Lycodon osmanhilli based on

two specimens (a holotype and a paratype) from Colom-

bo in Sri Lanka. Gunther (1864) and Boulenger (1893)

listed several ‘varieties’ of L. aulicus. As can be seen

from the list of synonyms which were based on speci-

mens from ‘India’, Bengal, Colombo and Mergui and

the generic transfers from across late 18" century to mid

20" century (Wallach et al. 2014), the Lycodon aulicus

complex has had a rather controversial taxonomic and

nomenclatural history.

Except for Lycodon osmanhilli, all of the above-men-

tioned nomina are currently considered to be subjective

junior synonyms of Lycodon aulicus (see, for example,

Whitaker & Captain 2004; Uetz 2016). Wallach et al.

(2014) considered L. osmanhilli a synonym of L. auli-

cus as well, but other authors (Das & De Silva 2005; So-

maweera 2006) considered it a valid species. While the

validity of Lycodon osmanhilli is still being discussed,

Corresponding editor: W. Bohme

26 Sumaithangi Rajagopalan Ganesh & Gernot Vogel

recently Pyron et al. (2013), in their molecular phylog-

eny, showed that Lycodon osmanhilli is distinct from L.

aulicus. Their phylogenetic tree revealed that L. aulicus

is the sister taxon of L. zawi Slowinski, Pawar, Win,

Thin, Gyi, Oo & Tun, 2001 from the Indoburmese re-

gion, while L. osmanhilli is the sister taxon of L. capuc-

inus (Boie, 1827) of Southeast Asia (Pyron et al. 2013).

Siler et al. (2013) even synonymized L. capucinus with

L. aulicus, although they did not examine Indian or Sri

Lankan material, a hypothesis not followed here and by

later workers (Vogel & Harikrishnan 2013; Wallach et al.

2014).

Ganesh & Chandramouli (2011) remarked on two syn-

topic morphotypes of the Lycodon aulicus complex from

Coromandel Coast and enumerated morphological dif-

ferences. They pointed out differences in general body

colouration, head dimensions and band pattern between

the two morphotypes and stated that one of the morphs

resembled the Sri Lankan endemic L. osmanhilli. Our

further examination of a series of preserved specimens

from several localities, including the type specimens, re-

vealed consistent differences, as suggested earlier (Ga-

nesh & Chandramouli 2011). In this work, we reassess

the systematics of Lycodon aulicus sensu auctorum and

provide formal taxonomic and nomenclatural implica-

tions.

MATERIALS & METHODS

For this study we investigated a total of 74 specimens of

the complex of Lycodon aulicus originating from Mauri-

tius and Pakistan in the west, across India on to Myan-

mar in the east, Nepal in the north and Sri Lanka in the

south, thus essentially covering the Indian subcontinent.

Several live examples were also examined. Specimens

were examined for external morphological characters.

Forty-four morphological characters were recorded for

each specimen. Not all of these characters were useful to

distinguish between species 1n this study, but all of them

were compared because they may be useful for further

taxonomic actions. Measurements, except body and tail

lengths, were taken with a slide-caliper to the nearest 0.1

mm; all body measurements were made to the nearest

millimetre. The number of ventral scales was counted ac-

cording to Dowling (1951). Hemipenial morphological

definitions and terminologies follow Dowling & Savage

(1960). Half ventrals were counted as one. The first scale

under the tail meeting its opposite was regarded as the

first subcaudal, the terminal scute was not included in the

number of subcaudals. The dorsal scale rows were count-

ed at one head length behind head, at midbody (..e., at the

level of the ventral plate corresponding to a half of the

total number of ventrals), and at one head length before

vent. We considered infralabials being those shields that

were completely below a supralabial. Values for paired

Bonn zoological Bulletin 67 (1): 25-36

head characters are given in left/right order. Temporal

scales were defined as the scales of which more than half

of the area lies in front of an imaginary line that extends

from the apex of the last supralabial to the posterolateral

corner of the parietal. Ratio of the length of (complete)

tail to the total length of the snake (i.e., from snout tip

to tail tip) is calculated as relative tail length. The pale

bands on the body and tail were counted on one side,

usually the right side. Hardly visible or incomplete bands

were counted as one band; bands that were fused (often

forming an “X”) were counted as two. The collar on the

neck was not counted and bands covering the anal shield

were added to the bands of the body. Sex of preserved

specimens was determined by dissection of the ventral

tail base, while that of live individuals was examined to

the extent possible by gentle anal palpation. Statistical

tests were carried out in MS Office Excel and PAST soft-

ware (Hammer et al. 2000).

Abbreviations. Avg. — average; BMNH: The Natural

History Museum, London, UK. — CAS: California Acad-

emy of Sciences Museum, California, USA. — CSPT/S:

Chennai Snake Park Museum, Chennai, India. — FMNH:

Field Museum of Natural History, Chicago, USA. —

NHMW: Naturhistorisches Museum Wien, Vienna,

Austria. — MHNG: Muséum d’Histoire Naturelle, Ge-

neva, Switzerland. - SMF: Naturmuseum Senckenberg,

Frankfurt Am Main, Germany. — UPZM: Univ. of Pera-

deniya Zoology Museum, Sri Lanka. - ZFMK: Zoolo-

gisches Forschungsmuseum Alexander Koenig, Bonn,

Germany. — ZMB: Zoologisches Museum Berlin, Ger-

many.

SYSTEMATICS

Lycodon aulicus (Linnaeus, 1758)

Coluber aulicus Linnaeus, 1758

Natrix aulica — Laurenti, 1768

Lycodon subfuscus Cantor, 1839

Lycodon atropurpureus Cantor, 1839

Lycodon aulicum — Dumeril, Bibron & Dumeril, 1854

Lycodon aulicus oligozonatus Wall, 1909

Ophites aulicus — Wall, 1921

Lycodon aulicus — Smith, 1943; Daniel, 2002;

Whitaker & Captain, 2004; Goonawardene et al. 2006

Lycodon travancoricus (not of Beddome, 1871) —

Rao et al. (2005)

Lycodon aulicus morph! — Ganesh & Chandramouli,

2011

Lycodon aulicus — (in part.) Wallach et al. (2014)

Material examined. Males (n=25): Myanmar: NHMW

21699.1 Bhamo; CAS 215387 Sagaing; Nepal: FMNH

62427, Tansing; BMNH 1936.7.2.2 Mae Dist, Doons;

BMNH 80.11.10.138 Nepal; India: BMNH 1908.5.23.15

©ZFMK

Taxonomy of Lycodon aulicus 27

Diburgash, Assam; FMNH 165108 Junganathpur, West

Bengal; FMNH 8650 Central province near Chanda;

FMNH 60647 Central province, Balaghat dist; BMNH

82.8.26.22 Kinelly (=Kimdey) hills, [Andhra Pradesh];

BMNH 74.4.29.958 Wynads, [Kerala] India; ZMB 1790

Bengal; BMNH_ 1904.10.18.5 Cannannore, Malabar:

NHMW 37406:1 Ahmednagar, Maharashtra, NHMW

37406:2 Ahmednagar, Maharashtra; Sri Lanka: FMNH

123906 Ceylon; ZFMK 52137 Kitulgala; ZFMK 52511

Kitulgala, NHMW 21689:5-—7 Sri Lanka; NHMW

14487:2—3 Sri Lanka; Indian Ocean Islands: ZFMK

29976 Mauritius; ZMB 8158 Isla Bourbon?; NHMW

21699.5 Ainoi islands in Hawaii.

Females (n=34): Myanmar: CAS 205000 DNA test-

ed, Rakhin; CAS 245960 Tanintharyi; CAS 219800

Ayeyarwadi,; NHMW 14483 Myanmar; ZMB 11625

Myanmar; NHMW 21702.2 Pegu, ; ZMB 10258 Min-

hla; BMNH 1928.1.4.1 Rangun; Pakistan: SMF 64484

Lahore, W-Pakistan; Nepal: BM 1984.1216 Royal Chit-

wan; FMNH 83090 Kathmandu; India: CAS-SU 12263

Bisrampur, Madhya Pradesh; FMNH 165107 West Ben-

gal, Howrah Dist.;, FMNH 161469 West Bengal, Barni-

junoh; NHMW 14487.1 ‘Alakan’; ZMB 1791 Bengal;

ZMB 9956 Ajmere, Rhajasthan; ZMB 1806 Calcutta;

NHMW 14488 Kolkata, BMNH 1921.6.15.3 Banga-

lore, Karnataka; SMF 32463 Agra; ZMB 1791 Bengal;

BMNH 1955.1.3.11 Mysore, 3500 ft, Karnataka; BMNH

1936.1.3.4 Namakal, Tamil Nadu; BMNH 1924.10.13.9

Punakanaat, 700 ft, Travancore, Kerala) BMNH

69.8.28.94 Matheran, Maharashtra; Sri Lanka: FMNH

123907 Ceylon, Trincomalee; ZFMK 52510 Sri Lanka;

NHMW 21689: 1-3 Sri Lanka; NHMW 14487:1 Sri Lan-

ka; Indian Ocean Islands: ZFMK 21766 Mascarenes,

Reunion, Manapany; ZFMK 29977 Mauritius.

Diagnosis (redefined herein). A species of Lycodon

inhabiting the Indian subcontinent, characterised by (1)

a wide and large head, (2) a distinct creamy white col-

lar-mark on head across parietal scales converging to-

wards snout-tip, (3) a dark blackish-brown body with

creamy white cross bars in life, (4) preocular usually

contacting frontal, (5) supraocular usually not contacting

prefrontal, (6) a fairly elongate hemipenis with smaller

flounces and spines, (7) supralabials white, usually 9 on

each side, (8) divided anal scale, (9) scale rows 17:17:15,

(10) ventrals: 180—215 and subcaudals: 57—78 pairs, (11)

relative tail length 0.15-0.20.

Description and variation (Fig. 1). A medium-sized

(avg. 500 mm total length, our longest specimen was a

female with 719 mm [BM 1924.10.13.9 from Punaka-

naat, 700 ft, Travacore, S India]) snake with heavy thick-

set, stoutly built head and rather robust cylindrical trunk.

Rostral scale scarcely visible from above; nasals small,

sutured, in contact with 1% and 2™ supralabials; interna-

Bonn zoological Bulletin 67 (1): 25-36

sals large, higher than broad; prefrontals vertically ob-

long, as large as frontal, in broad contact with loreal and

preocular; anterior end of prefrontal not half as wide as

posterior end, but only slightly smaller; frontal triangular,

slightly larger than supraocular, usually in clear contact

with preocular; anterior end of frontal not twice as wide

as posterior end; supraocular not in contact with prefron-

tal; postoculars 2, small; temporals usually 2+3+3; su-

pralabials usually 9, 3" to 5" touching eye; parietals very

large, subequal in length to its distance from internasals;

infralabials horizontally elongate, 10-11; 1 to 5" touch-

ing genials; anterior genials larger than posterior genials;

body scales smooth and glossy, imbricate, with mild

apical pits; dorsal scales in 17:17:15 rows around body;

preventrals usually 1-3; ventrals 180—205 (avg. 191.3) in

males and 186—208 (avg. 199.4) in females, angulate lat-

erally; anal scale divided; subcaudals 61—78 pairs (avg.

68.8) in males and 57—74 pairs (avg. 64.5) in females;

relative tail length on average 0.186 in males (0.172—

0.204) and 0.168 in females (0.146—0.191). Hemipenis

fairly thin, cylindrical and short, extending up to 10"

subcaudal scale, mildly forked near tip; pedicle slightly

narrower than hemipenial lobe head; hemipenial head not

quite bilobed; sulcal lips broader and ornamented with

thick pointed spiny flounces, visible heavily on asulcate

side and mildly on sulcate side. Sri Lankan specimens,

in both the sexes, have shorter tails, and a lower number

of ventral and subcaudals scales compared to peninsular

Indian specimens and Indoburmese (here understood as

the region from Northeast India upto Burmese peninsula)

specimens that had the longest tails (see Table 1).

Colouration in life. Dorsum blackish-brown or dark

brown, never without tinge of black; a series of about

8—40 creamy white cross bars either wholly complete

across the dorsum, or broad on the vertebral row then

diverging or disintegrating into two arms laterally; band

width covers 2-4 dorsal body scales; interband distance

typically covers 10—15 dorsal body scales; bands more

thick and prominent on forebody, obscure or absent on

hindbody, rarely completely absent, except for traces of

white collar mark; underside, upper lip, lower lip, throat

and chin pure white, slightly pinkish in juvenile speci-

mens; tongue rosy pink; iris totally black, pupil not vis-

ible.

Remarks. Linnaeus (1758) is his original description

of Coluber aulicus, mentioned ‘vertex albus’ meaning

‘white crown of head’ in Latin. The subsequent taxa

described by Cantor (1839) were based on specimens

in conformity with Linnaeus’ (1758) description. As

explained in Ganesh & Chandramouli (2011), Smith

(1943), Daniel (2002), Whitaker & Captain (2004) and

Goonawardene et al. (2006), correctly described Lyco-

don aulicus sensu stricto in their accounts of Lycodon

aulicus although they did not recognize the two differ-

©ZFMK

28 Sumaithangi Rajagopalan Ganesh & Gernot Vogel

eee ee

Fig. 1. Lycodon aulicus adult in life (a) from Sri Lanka, Photo: Dushantha Kandambi; (b) entire — dorsal view, dark morph; (c)

entire — dorso-lateral view, light morph; (d) entire — dorsal view, (e) head — dorsal view, (f) head — lateral view, (g) mid-body profile

view, all live adult specimens from Mayiladuthurai, India. Photos: S. R. Ganesh & S. R. Chandramouli (h) hemipenis of preserved

specimen CAS (California Academy of Sciences) 215387 from Sagiang Divsn., Myanmar. Photo: Gernot Vogel.

Bonn zoological Bulletin 67 (1): 25-36 ©ZFMK

Taxonomy of Lycodon aulicus 29

Table 1. Geographical variation within the Lycodon aulicus and L. anamallensis groups. Min-max ranges and mean values (within

parenthesis) are provided.

Characters Lycodon aulicus s. str. Lycodon anamallensis

Regions Indoburma Peninsular India Sri Lanka Peninsular India Sri Lanka

(sample sizes of sexes) m=5, f=11 m=11, f=15 m=9, f=6 m=3, f=5 m=2, f=5

Ventrals (males) 182-205 180-206 180-186 174-186 184-186

(193.0) (196.1) (182.9) (180.7) (186.0)

Subcaudals (males) 68-72 65-74 61-70 63-64 71-73

(70.0) (69.8) (65.9) (63.5) (72-5)

Rel. tail length (males) 0.182—0.204 0.175—0.187 0.172-0.188 0.195-0.197 0.185—0.200

(0.195) (0.183) (0.181) (0.196) (0.192)

Ventrals (females) 186-207 191-215 190-202 186-197 195-204

(197.5) (201.5) (195.5) (193.6) (200.4)

Subcaudals (females) 56-74 57-73 57-67 60-74 63-71

(64.0) (65.7) (60.8) (67.5) (67.4)

Rel. tail length (females) 0.154—0.189 0.146-0.191 0.149-0.159 0.176—0.185 0.172-0.180

(0.175) (0.167) (0.155) (0.181) (0.175)

ent morphotypes. Goonawardene et al. (2006) also dealt

with L. osmanhilli. Wall (1909) misunderstood Linnaeus’

definition of this species and went on to name this same

morphotype as his new subspecies Lycodon aulicus oli-

gozonatus and remarked it to be rare in southern India.

Lycodon aulicus is found throughout the Indian subcon-

tinent including Nepal, Pakistan, Bhutan, Bangladesh,

Myanmar, Sri Lanka and the mainland India (but not the

Andaman and Nicobar Islands). It is also found in Mau-

ritius and the Hawaiian Islands.

Lycodon anamallensis Giinther, 1864

Lycodon aulicus ‘typica’— Wall, 1909

Ophites anamallensis — Wall, 1923

Lycodon osmanhilli Taylor, 1950 syn. nov.

Lycodon aulicus (not of Linnaeus, 1758) —

Whitaker, 1978; Das, 2002; Das & De’ Silva, 2005;

Rao et al. (2005)

Lycodon cf. aulicus morph2 —

Ganesh & Chandramouli, 2011

Lycodon aulicus (in part.) Wallach et al. (2014)

Material examined. Males (n=5): India: BMNH

1904.10.18.2 Cannanore, Malabar, south India; BMNH

1904.10.18.4; Cannanore, Malabar, South India; CSPT/

S-28b Madras, India; Sri Lanka: FMNH 25927 Ceylon:

Colombo; MHNG 1198.70 Sri Lanka.

Females (n=10): India: BMNH 1946.1.14.92 Holo-

type of Lycodon anamallensis Anamallays; BMNH

1904.10.18.3 Cannanore, Malabar, Kerala; BMNH no

number Madras; BMNH 1924.10.13.7 Mundakayan,

Bonn zoological Bulletin 67 (1): 25-36

Trawancore, Kerala; CSPT/S-28a Madras; Sri Lanka:

ZFMK 32253 Sri Lanka; UPZM-17a&b Peradentya,

Kandy; MHNG 744.7 Ceylon; NHMW 21689.4 Ceylon.

Diagnosis (See also Taylor, 1950). A species of Lycodon

presently known from peninsular India and Sri Lanka,

characterised by (1) a thin and small head, (2) absence of

collar-mark on head; but the first ‘band’ passing across

neck > 7-10 scales away from parietals, and converging

towards tail, (3) a reddish-brown body with yellow or

cream (never quite white) cross bars in life, (4) preocu-

lar usually not contacting frontal, (5) supraocular usually

contacting prefrontal, (6) a shorter hemipenis with nu-

merous long flounces and spines, (7) supralabials creamy

with a median brown spot, usually 9 on each side, (8)

bifid anal scales, (9) scale rows 17:17:15, (10) ventrals:

174-204 and subcaudals: 60-73 pairs, (11) relative tail

length 0.14—0.20.

Description and variation (Figs. 2—4). A small to me-

dium-sized (avg. 400 mm) snake with a thin head and

neck, trunk and tail subcylindrical to slightly depressed.

Rostral scale scarcely visible from above, nasals more

or less pierced by nostril, partly sutured, in contact with

1*' supralabial; internasals distinctly larger than nasals,

in broad contact with preocular and loreal: loreal one on

each side (two on each side in the nominotypical holo-

type); prefrontals longer than wide, each prefrontal as

large as frontal, usually in contact with supraocular; an-

terior end of prefrontal distinctly half as wide as posteri-

or end; frontal pentagonal, produced posteriorly, slightly

larger than supraocular; anterior end of frontal distinctly

©ZFMK

30 Sumaithangi Rajagopalan Ganesh & Gernot Vogel

Fig. 2. Holotype of Lycodon osmanhilli KUMNH (Kansas University Museum of Natural History) 24141; (a) entire — dorsal view;

(b) entire — ventral view; (c) jar label. Photos: Peter Uetz.

twice as wide as posterior end; frontal usually not in con-

tact with preocular; parietals long, but distinctly smaller

than its distance from internasals; postoculars 2; preocu-

lar 1, half as long as loreal; temporals usually 2+3+4; su-

pralabials 9, 3" to 5“ contacting eye; infralabials usually

10-11, horizontally elongate, usually 1* to 5" touching

anterior genials; anterior genials larger than posterior ge-

nials; preventrals 1-3; ventrals 174-188 (avg. 186.0) in

males and 186—204 (avg. 197.0) in females, angulate lat-

erally; anal scale divided; subcaudals 63-73 pairs (avg.

67.8) in males and 60-72 pairs (avg. 66.6) in females;

average of relative tail length 0.194 in males and 0.172 in

females. Hemipenis short and stout, reaching only 7-8"

subcaudal scale; mildly forked near tip; pedicel barely

visible through elongate spines, hemipenial lobe head

greatly broader than pedicel, unilobed, flattened, circu-

lar and disc-like; sulcal lips ornamented with heavy and

elongate spines, many as long as two subcaudal scales;

sulcus spermaticus duct barely visible on both sulcate

and asulcate sides, being obscured by the spiny flounc-

es. Sri Lankan specimens, in both the sexes have higher

ventral and in males higher subcaudal scale counts than

peninsular Indian specimens (see Table 1).

Colouration in life. Dorsum fawn brown or red-

dish-brown, never with a tinge of black; a series of about

15-32 yellowish-white or cream coloured cross bars,

either wholly complete across the dorsum, or broad on

the vertebral row and diverging or disintegrating into

Bonn zoological Bulletin 67 (1): 25-36

two arms laterally, bands sometimes speckled inside

with background colour; bands more thick and evident

on forebody, obscure or absent on hindbody, rarely alto-

gether absent; upper lip, lower lip, throat and chin pale

pinkish white dotted with brown, venter uniformly white,

pinkish-cream in juvenile specimens; tongue rosy pink;

iris totally black, pupil mildly or not visible.

Remarks. Gunther (1864) described Lycodon anamal-

lensis based on a single specimen from the Anamallay

Hills, Western Ghats, peninsular India, deposited in Col.

R. H. Beddome’s collection. Our re-examination of the

holotype and additional preserved and living examples

from India and Sri Lanka revealed that all specimens ex-

cept the holotype have only one loreal scale on each side

of head and divided anal scales. Therefore, we concur

with Smith’s (1943) remarks that the presence of dou-

ble loreals on each side of head and undivided anal scale

recorded in both Indian and Sri Lankan specimens are

outliers and part of intraspecific variation (see also Wall,

1923). Taylor (1950) described Lycodon osmanhilli, dis-

cerning it from L. aulicus principally based on the char-

acter of the preocular separated from frontal. Our exam-

ination reveals that the holotype of L. anamallensis has

its frontal separated from preocular on one side but con-

tacting the preocular on the other side. However, our ex-

amination of both living and preserved specimens reveals

that the holotype of L. anamallensis might be termed as

a partial outlier in this character too. Nonetheless, preoc-

©ZFMK

Taxonomy of Lycodon aulicus 31

Fig. 3. Lycodon anamallensis in life (a) adult from Nuwalapitiya, Sri Lanka, Photo: Gernot Vogel; (b) adult from Madras, India;

(c) adult, band-less morph from Mayialduthurai, India; (d) adult with eggs; (e) a neonate from Madras; (f) head — lateral view; (g)

head — dorsal view, of specimen from Mayiladuthurai, India. Photos: S. R. Ganesh & S. R. Chandramoul1.

Bonn zoological Bulletin 67 (1): 25-36 ©ZFMK

52 Sumaithangi Rajagopalan Ganesh & Gernot Vogel

Fig. 4. Lycodon anamallensis (a-e) Holotype, BMNH 1946.1.14.92, from Anamallays, India; (a) entire — dorsal; (b) entire — ventral;

(c) head — lateral; (d) head — dorsal; (e) head — ventral views Photos: Gernot Vogel; (f) Non type CSPT/S-28a hemipenis of speci-

men from Madras, India Photo: S.R. Ganesh

Bonn zoological Bulletin 67 (1): 25-36 ©ZFMK

Taxonomy of Lycodon aulicus 33

Component 2

Component 1

Fig. 5. Principal Component Analysis plot showing rather mild separation of the taxa L. anamallensis (green cross — females, red

plus — males) and L. aulicus (pink circles — females and blue squares — males).

y = 0.541x - 0.538 te

R? = 0.875 a

Head Width (in mm)

8 apt y = 0.373x + 1.751

& 2 . R? = 0.899

10 12 14 16 18 20 22 24 26

Head Length (in mm)

Fig. 6. Regression biplot depicting differences in head length vs. head width ratios for L. aulicus s. str. (dotted line) and L. ana-

mallensis (normal line).

Bonn zoological Bulletin 67 (1): 25-36 ©ZFMK

34 Sumaithangi Rajagopalan Ganesh & Gernot Vogel

Table 2. Comparison of diagnostic characters of peninsular Indian and Sri Lankan Lycodon species (except the strongly keeled-

scaled L. carinatus, endemic to Sri Lanka). Data for other taxa sourced from Smith (1943), Whitaker & Captain (2004), and

Mukherjee & Bhupathy (2007).

Characters Lycodon aulicus L. anamallensis L. striatus L. flavicollis L. flavomaculatus L. travancoricus

Collar Present Absent Present Present Present (spot) Obscure

Dorsal Brown (rarely black- brown Blackish brown Brown Blackish brown Blackish brown

ground colour ish)

Colour of Creamy white Yellowish white White (with yel- Yellow Yellow Yellow (rarely

bands low mid-spot) yellowish white)

Supralabials 9 (very rarely 8 or 10)! 9? 8 (rarely 9) 9 9 9

Anal 2 2 : 2 1

Ventrals 180-215 174-204 154-166 210-224 165-183 176-206

Subcaudals 56-78 60-73 35-50 65-72 53-63 64-76

Hemipenis Reaching 10" subcau- Reaching 8" Reaching 10" — Cylindrical, not Reaching 15" Reaching 12" sub-

dal scale; not quite subcaudal scale; subcaudal forked at tip, subcaudal scale; caudal scale; forked

forked; smaller spiny mildly forked; scale; mildly lacks spines _—_ forked at tip, mildly _at tip; lobe head /

flounces broad spiny lobe forked; distal spinose pocket smooth

head 1/3" flounced,

spinose at tip

Distribution Indian subcontinent Peninsular Indian Western Northern Western Hills of

India & Sri subcontinent Ghats (low Ghats peninsular India

Lanka Nilgiris)

' 8 in 1 out of 120 cases, 10 in 3 out of 120 cases with two of these occurring on a specimen from Reunion

? In the holotype there are 10 supralabials on the left side, but it can easily be seen that one supralabial is split

ular-frontal separation is still a very typical character for

L. anamallensis and is diagnostic when used in conjunc-

tion with other characters mentioned above.

As explained in Ganesh & Chandramouli (2011), the

accounts of Lycodon aulicus sensu auctorum by Whita-

ker (1978), Das (2002), Das & De’Silva (2005) and Rao

et al. (2005) refer to Lycodon anamallensis. Das & De

Silva (2005) did so in their book covering Sri Lankan

snakes, despite recognizing L. osmanhilli. Earlier, Wall

(1909) had apparently mistaken the original concept

of L. aulicus [s. str.] and after having discerned this L.

anamallensis morphotype, called it Lycodon aulicus typ-

ica (contra Linnaeus, 1758) and remarked it to be quite

common in southern India, commoner than L. aulicus as

redefined herein. As currently understood, L. anamallen-

sis is known to occur in peninsular India (roughly as far

north as 21°N) and Sri Lanka. Based on our specimen ex-

aminations, L. anamallensis is absent in the Indian Ocean

Islands like Mauritius, Reunion and the Hawaii.

Our Principal Component Analysis (Fig. 5) run based

on 15 characters from 63 specimens, including 13 L. ana-

mallensis specimens (7 m, 6 f) and 50 L. aulicus spec-

imens (29 m, 21 f) revealed a mild separation of these

forms with outliers, indicating their degree of crypsis to

a certain extent. Of the 15 variables analysed, 7 had ei-

Bonn zoological Bulletin 67 (1): 25-36

genvalues > 1, ranging from 1.03 in Component 7 to up

to 2.61 in Component 1. They explained a variance of

99.18%. Component | was loaded on relative tail length,

ventral and subcaudal scale counts, while Component 2

was loaded on head-scale configuration, collar and band

patterns and supralabial markings. As can be seen by the

PCA plot, L. anamallensis (green cross — females, red

plus — males) and L. aulicus (pink circles — females and

blue squares — males) appear fairly separated. There is a

zone of overlap between the females of LZ. anamallensis

and the males of L. aulicus. This explains the crypsis in

this complex well. However, when analysed for head di-

mension, 1.e., head length vs. head width ratio, there is a

clear separation of these two taxa (Fig. 6). The regression

analyses indicate substantial differences in head dimen-

sions: for L. aulicus s. str. y = 0.541x - 0.538 R? = 0.875;

for L. anamallensis y = 0.759x - 4.645 R? = 0.933.

DISCUSSION

These two species, L. aulicus (Linnaeus, 1758) and L.

anamallensis Gunther, 1864, principally differ in head

dimensions, collar and band patterns, sex-specific rela-

tive tail lengths, hemipenal morphology (see Tables 1, 2)

©ZFMK

Taxonomy of Lycodon aulicus 35

as well as in genetics (Pyron et al. 2013). These two spe-

cies are largely sympatric over their geographic ranges

in the Indian peninsula (Whitaker & Captain 2004; Das

2002; this work), leaving little doubt to their specific dis-

tinction.

Our examination of voucher specimens also revealed

distinct and sometimes non-overlapping geographical

variation in relative tail lengths, and ventral and subcau-

dal scale counts between populations within L. aulicus

and L. anamallensis (see Table 1). However, it has to be

noted that each of such geographically-concordant vari-

ant populations do fall within the corresponding morpho-

types outlined above. Additionally, our material did not

reveal these geographically-correlated clinal differences

within populations of L. aulicus s. str. and L. anamal-

lensis to be large enough to warrant any further splitting

within these two species for the moment. Therefore, we

for now refrain from naming each of these individual

populations of L. aulicus from Nepal, Myanmar, penin-

sular India and Sri Lanka, and of L. anamallensis from

peninsular India and Sri Lanka. We conservatively main-

tain this stance for future investigations.

Additionally, we think that the dilution of these clearly

discernible morphotypes representing two distinct spe-

cies under the name L. aulicus, mainly in the Indian pen-

insula, has caused confusion about the identification and

distribution of this group of snakes. In Sri Lanka, how-

ever, the situation is different since the thin-headed form

had been called as LZ. osmanhilli and was usually regard-

ed as non-conspecific with L. aulicus (see Somaweera

2006). But recently Wallach et al. (2014) disregarded

these variations and included ZL. osmanhilli in the synon-

ymy of L. aulicus and worsened the situation. Pyron et al.

(2013), in their genetic analysis, showed that L. aulicus

and L. osmanhilli (i.e., from now on, a synonym of L.

anamallensis) are not only genetically distinct, but actu-

ally belong to different clades, with L. aulicus clustering

with L. zawi and L. osmanhilli clustering with the east-

ern species L. capucinus. As shown these two species are

easily diagnosable by the characters mentioned above.

KEY TO SOUTH ASIAN LYCODON

la. Body scales strongly keeled................... L. carinatus

ib. Bodyscales-fiot stronely-kedleds2%) 2.4.5... 2m oR, 2

Da A MASCa Ove MING weet x AE big ley L. travancoricus

Db Anal Seales: Cvadedh 04. 8s), of, scant Mean sic 1 eo. 3

2a. Ventrals < 200, body more black than brown ......... 4

3b. Ventrals > 200, body more brown than black .......... 5

4a. Usually 8 supralabials, reticulations white or with

VEMOWAMNIGKS POTS in. !u.icceewcvnsle des anise vets L. striatus

Ab. Usually 9 supralabials, reticulations always yellow

Fes Meet cee ek aC L. flavomaculatus

5a. Yellow collar always present, no other pattern,

ventrals not angulate laterally .............. L. flavicollis

Bonn zoological Bulletin 67 (1): 25-36

5b. Collar present or absent, body uniform or banded,

ventralscaneulate laterally. wu 8. wee. oer ees 6

6a. Collar present, touching the parietals, converging

towards snout tip 2.0.0... ecceeeeeeeees L. aulicus

6b. Collar absent, first band far away from parietals,

converging towards tail .............0.... L. anamallensis

Acknowledgments. We are grateful to our respective organi-

sations for supporting our research activities. At the Chennai

Snake Park, thanks are due to Executive Chairman and Trust-

ees. We thank the following persons, who gave us access to

Specimens in their care: Patrick Campbell (BMNH); Jens Vin-

dum and Alan Leviton (CAS), Alan Resetar and Harold Voris

(FMNH), Heinz Grillitsch, Silke Schweiger and Georg Gassner

(NHMW), Andreas Schmitz (NHMG), Gunther Kohler and

Linda Acker (SMF), Rupika Rajakaruna, Madhava Meega-

skumubura (UPZM), Dennis Rédder and Wolfgang Bohme

(ZFMK), Mark-Oliver Rodel and Frank Tillack (ZMB). We are

thankful to Peter Uetz and Dushantha Kandanmby for provid-

ing photographs, S.R. Chandramouli for providing pictures and

for advising on statistics and Johan Van Rooijen for advising on

statistics. We thank Ruchira Somaweera for providing informa-

tion on Sri Lankan snakes and Sameera Suranjan Karunarathna

for his information on Sri Lankan fauna and for all his help

during our Sri Lankan visits. Also a big thank you for Madha-

va Botejue, Dushantha Kandanmby, Udaya Chanaka (the Joga

master) and Thasun Amarasinghe for their help during the field

trips. Finally, we would like to thank Patrick David and Nis

Nicolaisen for reviewing the manuscript.

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©ZFMK

Bonn zoological Bulletin 67 (1): 37-40

May 2018

Epiplatys bifasciatus (Steindachner, 1881) (Nothobranchiidae) and

Hemichromis fasciatus Peters, 1852 (Cichlidae),

two relict fish species in the Sahara desert

Sébastien Trape

6 rue Bocaud, F-34 000 Montpellier, France

* Corresponding author. E-mail: sebastien.trape@gmail.com

Abstract. Two tropical fish species, Epiplatys bifasciatus and Hemichromis fasciatus, are recorded for the first time from

the Sahara desert, in Lake Boukou and Lake Dyara respectively, two of the seven Ounianga Serir lakes in northern Chad.

The nearest known populations of these two species are located 900 km to the southwest in Lake Chad. The Ounianga

Serir lakes, which resisted the increasing aridity of Sahara since the Holocene by virtue of subsurface inflow of fresh

groundwater from a large fossil aquifer, present the richest relict fish fauna of the Sahara, with at least eight species, in-

cluding also Hemichromis cf. letourneuxi, Sarotherodon galilaeus borkuanus, Coptodon zillii, Astatotilapia tchadensis,

Polypterus senegalus and Poropanchax normani.

Key words. Biogeography, relict fish hotspot, climate change, killifish, cichlids, Ounianga, Chad.

INTRODUCTION

Relict fish populations are known in several perennial

bodies of waters of the Sahara desert, most of them locat-

ed in mountainous massifs of central Sahara: the Adrar

mountains in Mauritania, the Ahaggar, Tassili n’Ajjer

and Mouydir in Algeria, and the Tibesti and Ennedi in

Chad (Lévéque 1990, 2006; Trape 2009). Fish diversity

is low, only two dozen of species have been recorded for

the whole Sahara, and most species are known from a

very low number of permanent water bodies, often from

a single spring, guelta, pound or lake (Trape 2009). The

number of species sharing the same water body usually

ranges from one to three, with a maximum of four species

Fig. 1. Satellite view of the Sahara with location of Ounianga

lakes (Chad).

Received: 21.12.2017

Accepted: 28.03.2018

Fig. 2. Satellite view of Ounianga Serir lakes, with location of

Lake Dyjara and Lake Boukou.

in Molomhar guelta in Mauritania, and seven species in

Totous guelta in Tibesti in Chad (Daget 1959, Lévéque

1990, Monod 1951, Trape 2009).

During zoological surveys in northern Chad in 2013

and 2014, fishes, reptiles, amphibians, and invertebrates

were collected in various areas of Borkou, Ennedi and

Tibesti (Brancelj 2015, Dumont 2014, Trape 2013, 2015,

2016). Among fishes collected in Ounianga Serir Lakes in

2013 and 2014 (Figs 1—2), several specimens belonged to

three species never reported before from northern Chad

nor from other areas of the Sahara desert, namely Poro-

panchax normani (Ahl, 1928) (Poeciliidae), Polypterus

senegalus Cuvier, 1829 (Polypteridae), and the previ-

Corresponding editor: X. Mengual

38 Sébastien Trape

ously undescribed Astatotilapia tchadensis Trape, 2016

(Cichlidae) (Trape 2013, 2016). Other species report-

ed from Ounianga Serir lakes included Sarotherodon

galilaeus borkuanus (Pellegrin, 1919), Hemichromis

cf. letourneuxi Sauvage, 1880 (erroneously reported as

Hemichromis bimaculatus Gill, 1862), and Coptodon

zillii (Gervais, 1848) (Daget 1959, Lévéque 1990, Trape

2013). Here I report the occurrence of two additional spe-

cies collected in these lakes in 2016, both representing

remarkable additions to the known relict fish fauna of the

Sahara.

MATERIAL & METHODS

Measurements, counts and diagnosis were made as de-

scribed in Paugy et al. (2003), Wildekamp & Van der

Zee (2003) and Teugels & Thys Van den Audenaerde

(2003). Measurements were made with a digital calliper,

and counts partly under a dissecting microscope. Spe-

cies keys of Teugels & Thys Van den Audenaerde (2003)

and Wildekamp & Van der Zee (2003) were used for the

species diagnosis of Hemichromis fasciatus and Epiplat-

ys bifasciatus, respectively. Preserved specimens are

deposited in the Museum national d’Histoire naturelle

(MNHN) at Paris.

RESULTS

Hemichromis fasciatus Peters, 1852

(Fig. 3)

Material examined. MNHN 2016.0272, previously

IRD TR.4515, collected in Lake Boukou (18°54’54”N,

20°54’40”E, elev. 363 m) (Fig. 4) on April 4" 2016 in the

evening using a fishing rod by Jacques Robin.

Description. Two lateral lines. Scales cycloid. No pha-

ryngeal hanging pad between gills. Outer jaw teeth

Fig. 3. Hemichromis fasciatus from Lake Boukou in life.

Bonn zoological Bulletin 67 (1): 37-40

Fig. 4. View of Lake Boukou.

monocuspid. Upper profile of snout concave. Premaxilla

extremely protrusible. Lower jaw distinctly prominent.

Dorsal fin with 14 spines and 12 soft rays. Anal fin with

3 spines and 9 soft rays. Number of lateral-line scales:

29. Five dark blotches on sides, the first blotch confluent

with the opercular spot, the fifth on caudal-fin base. Stan-

dard length 180 mm.

Epiplatys bifasciatus (Steindachner, 1881)

(Fig. 5)

Material examined. MNHN 2016.0273, 5 specimens

collected in Lake Dyara (18°55’09”N, 20°53’39”E, elev.

355 m) (Fig. 6) on April 9" 2016 by Jean-Francois Trape.

Specimens were collected at night on the shore of the

lake using a dipnet.

Fig. 5. Two specimens of Epiplatys bifasciatus from Lake Dja-

ra in life.

©ZFMK

Relict fish species in the Sahara 39

“ae Raggi ere —

dhe

maa tem E,

Fig. 6. View of Lake Dyara.

Description. A tubular pre-ocular system with five

pores. A frontal supra-orbital neuromast system consist-

ing of one pit with two neuromasts. Dorsal fin 8—9 rays,

anal fin 14-18 rays. Scales on mid-lateral series 26—27.

Life colours of males (Fig. 5) are typical of this species

(see Wildekamp & Van der Zee 2003) with in particu-

lar oblique red stripes on the opercle, a large number of

oblique red bars on the sides and the back, a wide dark

grey longitudinal band extending from the opercle to the

caudal peduncle, and a number of red spots on the anal,

dorsal and caudal fins. Standard length 37-39 mm.

DISCUSSION

Hemichromis fasciatus is a well-known afrotropical

cichlid, abundant and widespread in most hydrographic

basins of West and Central Africa, in particular in Lake

Chad and the Senegal and Niger rivers basins (Lévéque

et al. 1991; Teugels & Thys van den Audenaerde 2003).

Hemichromis fasciatus has never been reported before

in the Sahara, contrary to its widespread congener He-

michromis bimaculatus, a complex of species with unre-

solved taxonomy which ts probably represented by H. /e-

tourneuxi in Ounianga Serir, Ounianga Kebir, and some

other bodies of water in the Sahara (Fig. 7) (Lévéque

1990, Lévéque et al. 1991, Loiselle 1979, Sauvage 1880,

Trape 2013, Teugels & Van den Audenaerde 2003).

Epiplatys bifasciatus is a killifish with a large distribu-

tion from Senegal in West Africa to Sudan in East Afri-

ca, both in weedy parts of rivers, swamps, small brooks

and rivulets in the Guinean and Sudanese savanna, and

in coastal lagoons on the shore of the Atlantic Ocean

(Wildekamp & Van der Zee 2003). The two other known

populations of killifish that survived the increasing arid-

ity of the Sahara since the Holocene are both located in

northern Chad in a single body of water: Epiplatys spilar-

Bonn zoological Bulletin 67 (1): 37-40

gyreius (Dumeril, 1861) in Tigui pool (Borkou) (Esteve

1952, Lévéque 1990), and Poropanchax normani in Lake

Boukou (Trape 2013). The nearest current populations of

these three species are located 900 km to the southwest in

Lake Chad (Blache 1964).

According to Grenier et al. (2009) the Ounianga Serir

lakes were connected to Megalake Chad in the Holocene.

Despite the current extreme aridity in this area — average

annual rainfall is less than 5 mm and annual evaporation

exceeds 6000 mm — these lakes persist by virtue of sub-

surface inflow of fresh groundwater from a large fossil

aquifer (Kropelin 2007). The remarkable biodiversity of

these lakes has remained poorly investigated and until re-

cently only the two fish species collected during the Thilo

expedition (1912-1917) in northern Chad - Sarotherodon

galilaleus borkuanus and Hemichromis cf. letourneuxi —

were known from Ounianga Serir lakes (Pellegrin 1919).

In fact, with at least eight species, these lakes present the

richest fish fauna of the Sahara desert.

Fig. 7. Hemichromis cf. letourneuxi from Lake Dyara (top) and

Ounianga Kebir (bottom) in life.

Acknowledgements. I thank Jacques Robin and Jean-Francois

Trape who collected these interesting specimens.

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adjacent du Mayo Kebbi. Etude systématique et biologique.

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the Tibesti area (Northern Chad) and a re-description of Pi-

locamptus schroederi (van Douwe, 1915). Zootaxa 3994:

53-555

©ZFMK

40 Sébastien Trape

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Travaux de I’ Institut de Recherches Sahariennes 18: 173-181

Dumont HJ (2014) Odonata from the Tibesti mountains and the

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with the first records of a polypterid and a poeciliid in the

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©ZFMK

Bonn zoological Bulletin 67 (1): 41-57

May 2018

News from the Balkan refugium: Thrace has an endemic mole species

(Mammalia: Talpidae)

Boris KryStufek'’, Nedko Nedyalkov’, Jonas J. Astrin’ & Rainer Hutterer°

‘Slovenian Museum of Natural History, Presernova 20, SI-1000 Ljubljana, Slovenia

?National Museum of Natural History, 1 Tzar Osvoboditel Blvd., BG-1000 Sofia, Bulgaria

3Zoologisches Forschungsmuseum Alexander Koenig, Leibniz-Institut fiir Biodiversitat der Tiere, Adenauerallee 160, D-53113

Bonn, Germany

*Corresponding author. E-mail: bkrystufek@pms-lj. si

Abstract. We utilized 1084 bp sequences of the cytochrome b gene to assess the taxonomic status of small blind moles

from eastern Thrace in Bulgaria and European Turkey. So far, these moles were classified either as Zalpa caeca or as T: le-

vantis. Our study showed them to be genetically closer to T. europaea, T. aquitania, and T. occidentalis, albeit not being

part of any of these species. We describe them as a new species, Za/pa martinorum. n. sp. The new species differs from

T. europaea, another mole occupying Thrace, by having a sealed palpebral fissure and a 1‘ upper molar with no parastyle,

and by being smaller. The contemporary distribution range of 7’ martinorum n. sp. 1s small and restricted to the Black Sea

coast between Burgas (Bulgaria) and Istanbul (Turkey). The species name is an eponym to the married couple Vladimir

and Evgeniya Martino, two early students of Balkan mammals.

Key words. Balkans, cytochrome 5b, cryptic species, species delimitation, 7Zalpa martinorum n. sp.

INTRODUCTION

Moles (family Talpidae, Lipotyphla) are widespread

in temperate Eurasia, North America, and in Oriental

Southeast Asia. The majority of species has adapted to

a subterranean mode of life and displays convergent

morphologies. Interspecific differences are usually slight

and further obscured by morphologic plasticity. The tra-

ditional approach towards mole taxonomy, based solely

on cranial and dental traits, frustrated early students and

posed a long-lasting source of dispute and disagreement

over a number of genera and species. Since 1758, when

Linneaus named the first mole species (Linnaeus 1758),

about 180 species group names accumulated (Hutterer

2005) for over 50 currently valid species. In the past, the

authors disagreed which of these names were valid and

which were to be considered of subspecific value or just

synonyms. For example, Schwarz (1948) synonymized

five currently valid generic names from Asia with Jalpa,

and reduced the number of Eurasian fossorial moles to

merely six species; Ellerman & Morrison-Scott (1951)

further condensed these to four species in two genera.

Despite obvious taxonomic problems with moles, mo-

lecular tools were employed in talpid systematics rela-

tively late (Colangelo et al. 2010; Bannikova et al. 2015;

Feuda et al. 2015; He et al. 2016). It became immediately

clear how incomplete our alpha taxonomic knowledge of

the Eurasian moles was. He et al. (2016) proposed that

talpid species richness is underestimated by nearly one

third. Although the majority of these undescribed species

occupy mountain areas in southern China, several cryp-

Received: 22.01.2018

Accepted: 16.04.2018

tic species were recently found also in the genus 7al/pa

in Europe (Nicolas et al. 2017a) and south-western Asia

(Bannikova et al. 2015).

In this paper we utilized molecular evidence in assess-

ing the taxonomic status of small blind moles along the

south-western Black Sea coast, specifically in Thrace of

Bulgaria and European Turkey. Osborn (1964) was the

first to realize that the region features two distinct mole

species, a larger 7? europaea, which is widespread in Eu-

rope, and a smaller one, which he took for 7! caeca. In

Osborn’s times, 7’ caeca contained small blind moles of

southern Europe, northern Anatolia, and the Caucasus

(Ellerman & Morrison-Scott 1951, Grulich 1972), which

are currently classified as distinct species (7! davidiana,

T. levantis, T: occidentalis; Hutterer 2005). Dogramaci

(1988, 1989c) applied the epithet /evantis (T. caeca le-

vantis) for small moles of Thrace and northern Anato-

lia. Following Pavlinov & Rossolimo (1987), Vohralik

(1991) and Kefelioglu & Gencoglu (1996) accepted 7

levantis as a species on its own right specifically for the

European populations. As a result, four mole species are

reported for south-east Europe (the Balkan Peninsula):

7? europaea in the north and east, 7’ caeca and T° stan-

kovici in the south-west, and 7’ /evantis in the south-east

(Hutterer 2005).

MATERIAL AND METHODS

Specimens. We studied 14 individuals of 7? martino-

rum n. sp. (see below) and comparative material of 244

Corresponding editor: J. Decher

42 Boris Kry8tufek et al.

museum vouchers belonging to 11 species of Talpa (Ap-

pendix 1). Individuals of 7? martinorum n. sp. were col-

lected in Bulgaria in 2011 (one individual) and 2017 (the

remaining 13). External measurements were scored be-

fore skinning to the nearest 0.1 mm (hind foot) or 1 mm

(the rest): length of head and body (from snout tip to

anus), length of tail (from anus to tail tip with exclusion

of terminal hairs) and length of hind foot (without claws).

Body mass was recorded to the nearest 0.5 grams. We

examined the palpebral fissure using a 10x magnifying

glass and photographed each mole captured in 2017. In

part the individuals were immersed in ethanol and the rest

were processed following standard mammalogical proce-

dures (KryStufek & Vohralik 2001). Skins were mounted

on cardboard (carded skins) while skulls and postcranial

skeletons were preserved in 95% ethanol, dried subse-

quently and cleaned by Dermestes beetles. Tissue sam-

ples for DNA analysis were placed in non-denaturated

96% ethanol and subsequently refrigerated. All field pro-

cedures involving handling of animals in this study were

in compliance with guidelines approved by the American

Society of Mammalogists (Sikes et al. 2011).

Molecular analysis. The cytochrome b (Cyt b) gene

was sequenced for eight specimens of the new species.

These sequences were analyzed together with 23 addi-

tional Talpa sequences (within 13 species), and three

outgroup Talpinae sequences (Urotrichus talpoides, Eu-

roscaptor parvidens and Euroscaptor klossi), all down-

loaded from GenBank. Collecting data for the new se-

quences and all GenBank numbers are given in Appendix

For the new specimens, total genomic DNA was ex-

tracted from ethanol-preserved muscle tissue using sil-

ica membrane columns of the Blood and Tissue kit by

Qiagen (Hilden, Germany). DNA extracts are available

from the ZFMK Biobank, Bonn (DNA voucher IDs are

listed in Appendix 2). For PCR amplifications, we used

the Qiagen Multiplex PCR kit, following the manufac-

turer’s specifications and based on 2 ul undiluted DNA

template in 20u1] total reaction volumes. DNA fragments

of 1084 bp were amplified with an Applied Biosystems

GeneAmp PCR System 2700 (Life Technologies), apply-

ing the primers L14724ag (5’-ATGATATGAAAAAC-

CATCGTTG-3’) and H15915ag (5’-TTTCCNTTTCTG-

GTTTACAAGAC-3’) (Guillén-Servent & Francis 2006).

PCR routine followed a ‘touch-down’ protocol: Taq ac-

tivation: 15 min at 95°C; first cycle set (15 repeats): 35 s

denaturation at 94°C, 90 s annealing at 60°C (—1°C per

cycle) and 90 s extension at 72°C. Second cycle set (25

repeats): 35 s denaturation at 94°C, 90 s annealing at

50°C, and 90 s extension at 72°C.

After enzymatic cleanup, all PCR products were

Sanger-sequenced at Macrogen Europe’s commercial

Sanger sequencing service (Amsterdam, NL). Sequences

were assembled, inspected and aligned using Geneious

vers. R7 (Biomatters, Auckland, New Zealand).

Bonn zoological Bulletin 67 (1): 41-57

The Maximum Likelihood (ML) tree was inferred with

RAxML-HPC vers. 8.1.24 (Stamatakis 2014). For the

ML search, a GIR+I° model of sequence evolution was

applied following the program recommendations. The

dataset was partitioned to treat 3rd codon positions sep-

arately from 1‘t and 2™ positions. The analysis used the

“-f'a” option (bootstrap analysis and search for best-scor-

ing ML tree in one program run) and included 100,000

bootstrap replicates. Nodes with a bootstrap support of

50 or below were collapsed. To comparatively assess the

topology delivered by the ML reconstruction, a Bayesian

analysis was run for 5 million generations (assuming a

GTR+I model and unlinking 3" positions) in MrBayes

vers. 3.2 (Ronquist & Huelsenbeck 2003), but results

were very similar and hence not shown in a separate il-

lustration.

Morphological analysis. Our study was based on

visual examination of museum specimens, both mac-

roscopically and under a stereomicroscope at different

magnifications. Museum vouchers (skins, skulls and

occasionally postcranial skeletons) are deposited in the

following collections (acronym in parentheses): Field

Museum of Natural History, Chicago (FMNH), National

Museum of Bosnia and Herzegovina, Sarajevo (ZMBiH),

National Museum of Natural History, Washington D. C.

(NMNH), National Museum of Natural History Sofia

(NMNHS), Natural History Museum London (NHML),

Naturhistorisches Museum Wien, Vienna (NMW), Mu-

seum Nationale d’Histoire Naturelle, Paris (MNHN),

Mammal Collection in the Ondokuz Mayis University,

Samsun, Turkey (OMU), Slovenian Museum of Natural

History, Ljubljana (PMS), Zoologisches Forschungsmu-

seum Alexander Koenig, Bonn (ZFMK), and Zoological

Institute and Zoological Museum, Russian Academy of

Sciences, St. Petersburg (ZIN). Vouchers in ZFMK and

PMS were directly compared to 7’ martinorum n. sp..

Material from the remaining collections was examined

by BK prior to this study and records were compiled to

contain measurement data, drawings and photographs.

Skull morphology was quantified using a set of six cra-

nial variables which were scored using a Vernier caliper

with accuracy to the nearest 0.1 mm (acronyms in pa-

rentheses): condylobasal length of skull (CbL), length of

maxillary tooth row (MxT; canine to 3 molar), breadth

of braincase (BcB), height of braincase (BcH; bullae

excluded), breadth of rostrum over canines (RoC), and

breadth of rostrum over molars (ROM). We also mea-

sured the pelvis (greatest length and greatest anterior

breadth) and the humerus (length and width). Length and

width of 3 upper molar were scored under a dissecting

microscope fitted with an eyepiece graticule. Three mor-

photypes of the pelvis (sensu Grulich 1971, Popov &

Miltchev 2001) were distinguished: (1) the caecoid mor-

photype lacked a bony anastomosis between the os sa-

crum and the os ischii posterior to the 4" foramen sacrale,

(i1) the europaeoid morphotype had a well-developed and

©ZFMK

New mole from Thrace 43

ossified anastomosis and a closed 4" foramen sacrale,

(i11) in the intermediate morphotype, the foramen was

closed, the anastomosis, however, was thin and narrow,

and the ischio-sacrale suture was not ossified.

Heterogeneity between samples was assessed in one-

way analysis of variance (Anova). To characterize the

craniometric variation among species of moles and to

find patterns 1n our high-dimensional data, we used prin-

cipal components analysis (PCA), which was performed

on the correlation matrix of log,,-transformed cranial

variables. The resulting principal components (PCs) are

linearly uncorrelated; therefore, each PC measures dif-

ferent ‘dimensions’ of the original dataset. The first PC

(PC1) is responsible for the largest possible variance and

acts in morphometrics as a size vector (KryS8tufek et al.

2015). Rates of correct classification of a priori defined

species were evaluated by discriminant analysis (DA). To

evaluate the performance of the DA and avoid the risk

of overfitting the data, all analyses were cross-validated

using the jackknife procedure, in which each specimen

is Classified into a group using the discriminant function

derived from all specimens except the specimen being

classified. Statistical tests were run in Statistica 7.0 (Stat-

Soft Inc., OK, USA) and SPSS Statistics 2012 (IBM

Analytics, NY).

RESULTS

Molecular results. Sequencing of the eight samples re-

sulted in a single haplotype, with the exception of one

alignment position (pos. 215) with ambiguous informa-

tion (“N’) in 3 sequences. No stop-codon insertions or

deletions were observed in the alignment.

In the ML tree, our new samples of 7) martinorum n.

sp. clustered with T. europaea, T: aquitania, and T. oc-

cidentalis into a weakly supported clade (Fig. 1). The

Bayesian analysis includes also 7. romana and T. caeca

in this cluster. Above species level, the ML branching pat-

tern was poorly supported (the very similar topology in

MrBayes had higher support values — but see Douady et

al. (2003), among others, on potentially misleading high

posterior probability values). Therefore the tree topology

offers only a meagre idea on phylogenetic relationships

among species. At species level however, support values

are usually much higher: the new species 7. martinorum

is recovered as monophyletic with maximal bootstrap

support. Also the genetic metrics show that 7) martino-

rum n. sp. 1s clearly separated from all other moles: the

closest genetic matches to 7? martinorum n. sp. were T.

europaea and T. occidentalis with p-distances of more

than 9% (1.e., distances well above the typical intraspecif-

ic range, see below). Maximal interspecific distances for

7. martinorum n. sp. were registered towards T. altaica

and T. talyschensis (both ca. 14%). Minimal interspecific

distances among all 7alpa species in the dataset were ca.

Bonn zoological Bulletin 67 (1): 41-57

8%, maximal distances 15%. Intraspecific p-distances in

T. martinorum n. sp. were 0.0 to 0.1% (the 0.1% diver-

gence created through the position mentioned above),

0.4 to 1.9% in T. europaea, 2.6% in T. aquitania, 1.8%

in T. ognevi, and 0.4 to 6.9% in T. stankovici. Conspicu-

ously high distances within 7. stankovici result from the

inclusion of genetically highly divergent Greek popula-

tions (from Gravia and Chelmos; see Tryfonopoulos et

al. 2010). The two specimens representing the two sub-

species of T. /evantis (1.¢., T: levantis levantis and T. le-

vantis minima) were separated by a distance of 7.5%.

Morphometric results. Secondary sexual dimorphism

in size (SSD) is usually obvious in the genus Zalpa

(Niethammer & Krapp 1990). Our sample from Thrace

contained nine males and four females, of which nine

were clean skulls. We therefore tested SSD only on the

external measurements. One-way analysis of variance

detected significant heterogeneity between sexes only

in length of hind foot (F=9.45, p=0.012) with males at-

taining a higher mean (17.21+0.204 mm) than females

(16.13+0.289 mm). Since the remaining external traits

showed no significant SSD (F<2.8, p>0.12), we pooled

the sexes.

All cranial measurements retrieved highly significant

interspecific heterogeneity (F>18, p<0.001) with highest

F-values (F>30) for length of skull and breadth of ros-

trum. To visualize morphometric distances and similar-

ity between species we ran PCA on 187 complete skulls

belonging to 11 species. Zalpa caucasica was excluded

because the available skulls were damaged. The first

two principal components (PC1 and PC2) in concert

explained 87.7% of the variance in the original data set

and were used to display morphometric relationships be-

tween species (Fig. 2). PC1 had high positive loadings

for all measurements and sorted moles according to their

overall size, from the smallest (negative scores on the left

hand side of the PC1 axis) to the largest (positive scores

on the right hand side of the axis). PC2 arranged moles

in respect to breadth of rostrum over molars. Moles with

high PC2 loadings had a broad rostrum and vice versa.

There was considerable overlap in the centre of the plot

between 7 martinorum n. sp., T: levantis, T. talyschen-

sis, T: occidentalis, and the larger individuals of 7. caeca.

Talpa europaea was well characterized by a combination

of large skull size and relatively narrow rostrum; 7: aqui-

tania was less variable than 7’ europaea and on average

had a narrower rostrum. The rostrum was most robust in

T. stankovici and T: davidiana, T. romana was transitional

in this respect towards 7! europaea. Talpa caeca attained

smaller dimensions than any other mole in our analysis.

Noteworthy, all principal components showed significant

heterogeneity between species (F>2.5, p<0.072).

Discriminant analysis performed on the same matrix

of log, ,-transformed cranial variables (Wilks’ A = 0.0248,

F = 14.408, p<0.0001) classified 65.6% (cross-validat-

ed 56.8%) of specimens to the correct species. All pair-

©ZFMK

44 Boris Kry8tufek et al.

7-—— Urotrichus talpoides EU918371

100 Euroscaptor parvidens AB823122

Euroscaptor klossi AB823107

Talpa altaica AB037602

98 Talpa talyschensis KP717370 Azerbaijan

Talpa davidiana KP/17367 Turkey

96 Talpa caucasica KP717353 Russia Nalchik

76 | lalpa ognevi KP717360 Georgia

64 Talpa ognevi KP717356 Turkey

Talpa levantis minima KP717346 Russia Adygea

Talpa levantis KP717343 Turkey Zonguldak

81 Talpa caeca KP717333 Italy

Talpa romana HQ233501

Talpa stankovici FJ688089 Greece Chelmos

Talpa stankovici FJ688098 Greece Gravia

g9 7 [alpa stankovici KP717348 Macedonia

Talpa stankovici KP717347 Macedonia

Talpa stankovici FJ688094 Greece Kosmira

Talpa occidentalis KU189723 Spain

100 += Talpa aquitania KF801510 Spain

Talpa aquitania KU189595 France HT

Talpa europaea KP7/17322 Turkey

‘ Talpa europaea FJ688085 Greece

59 Talpa europaea KF801574 Kosovo

Talpa europaea Y 19192 Sweden

Talpa europaea KU189429 France

Talpa martinorum sp. n. BG10

Talpa martinorum sp. n. BG15

Talpa martinorum sp. n. BG09

Talpa martinorum sp. n. BG01

Talpa martinorum sp. n. BG17

Talpa martinorum sp. n. BG14

TS Talpa martinorum sp. n. BG08

Talpa martinorum sp. n. BG02

100

Fig. 1. Maximum likelihood tree inferred from 1084 bp of the mitochondrial cytochrome b gene for 14 species of the genus 7Jalpa.

Bootstrap values are shown on the nodes. The tree 1s rooted with Urotrichus talpoides, Euroscaptor parvidens and Euroscaptor

klossi.

Bonn zoological Bulletin 67 (1): 41-57 ©ZFMK

New mole from Thrace 45

PC2 (8.0%)

—s

-2

-3

-3 -2 -1

PC1 (79.7%)

1 2 3

Fig. 2. Projection of group centroids (upper case letters) onto the two principal components (PCs) resulting from principal compo-

nents analysis of six log, -transformed cranial variables (percentage of variance explained by each component is in parentheses)

of 10 Talpa species. Ellipses are 95% confidence limits of the dispersion in each species except 7’ martinorum n. sp. (specimens

shown by triangles), 7! altaica (squares) and T: talyschensis (only group centroid is shown). Area inside the ellipses for 7’ aquitania,

T. europaea and T. occidentalis are shaded grey. The character vector diagram illustrates the relative contribution of the original

variables (see text for acronyms) to the principal components. Abbreviations for centroids: A — 7? aquitania, C — T. caeca, D — T-

davidiana, E — T. europaea, L — T: levantis, O — T: occidentalis, R — T: romana, S — T. stankovici.

wise-squared Mahalanobis distances (D7?) were signifi-

cant except two (7. talyschensis against T. occidentalis

and 7 martinorum n. sp., respectively). Pairwise D? dis-

tances with 7? martinorum n. sp. (mean+standard error

= 10.72+5.56) were low in comparison to the remaining

pairwise distances (20.21.87+2.62), the difference how-

ever was not significant (F=3.28, p=0.076).

A new mole species from Thrace

Talpa martinorum n. sp. (Figs. 3, 4, 5a, 6a)

Holotype and type locality. Skin mounted on cardboard,

skull, postcranial skeleton, and tissue sample in ethanol

of a young adult female (ZFMK-MAM-2017.1149; tis-

Bonn zoological Bulletin 67 (1): 41-57

sue: ZFMK-TIS-23035), collected by B. KryStufek and

N. Nedyalkov on 27.04.2017 (field No. BG8) on the edge

of a meadow near Zvezdets, Mt. Strandzha, Bulgaria.

DNA of this specimen has been deposited (ZFMK-DNA-

FC19476469) and the Cytochrome b sequence is avail-

able from GenBank (Accession number MH093593).

Measurements of holotype. Body mass 45 g, head

and body 124 mm, tail 24 mm, hindfoot length 16.4 mm,

condylobasal length of skull 30.4 mm, maxillary tooth

row 11.6 mm, breadth of braincase 15.2 mm, height of

braincase 9.2 mm, breadth of rostrum over canines 4.1

mm, breadth of rostrum over molars 8.3 mm, length of

humerus 14.1 mm, width of humerus 9.7 mm, greatest

length of pelvis 22.1 mm, breadth of pelvis 7.5 mm.

©ZFMK

46 Boris Kry8tufek et al.

Fig. 3. Head (a) in lateral view, tip of nose (b) in dorsal (left) and ventral (right) views, and ventral side of tail in Zalpa martinorum

n. sp. Note that the palpebral fissure is covered by transparent skin (a). Museum vouchers PMS 25631 (a), ZFMK 2017.1152 (b)

and ZFMK 2017.1151 (c). Not to scale.

Diagnosis. A member of the subgenus Zalpa. Me-

dium-sized species with palpebral fissure sealed by a

transparent skin (Fig. 3a). First upper molar (M') lacks

parastyle (Fig. 6a); the mesostyle is indistinctly bifur-

cate (Fig. 6a). Pairwise interspecific p-distances (> 9%)

are within the range observed between other species of

moles (e.g., within Zalpa, interspecific distances average

ca. 12%).

Paratypes. Skins, skulls, postcranial skeletons, tissue

samples in ethanol and isolated DNA; voucher NMNHS

1053 is represented by a skull and a tissue sample, and

vouchers NMNHS 1049-1052 are submerged in ethanol.

Specimens were collected on Mt. Strandzha, Bulgaria: Ve-

lika, 1 female, collected on 23.09.2011 by N. Nedyalkov

(NMNHS 1053), 1 female, collected on 01.05.2017 by

N. Nedyalkov & B. KryStufek (PMS 25631); Gramatiko-

vo, 2 males, collected on 25.04.2017 by N. Nedyalkov &

B. Krystufek (NMNHS 1047, 1048); Vizitsa, 1 male, col-

lected on 27.04.2017 by N. Nedyalkov & B. Kry8tufek

(ZFMK-MAM-2017.1150); Kondolovo, 1 male, col-

lected on 28.04.2017 by N. Nedyalkov & B. Kry8tufek

(ZFMK-MAM-2017.1151); Fazanovo, 1 male, collected

on 30.042017 by N. Nedyalkov & B. Kry8tufek (ZFMK-

MAM-2017.1152); Pismenovo, 1 male, 1 female, col-

lected on 01.05.2017 by N. Nedyalkov & B. Kry8tufek

(PMS 25632, 25633); Chengene skele, Burgas, 1 male,

1 female, collected on 19.11.2017 by Nedko Nedyalkov

(NMNHS 1049, 1050), Uzungeren, Burgas, 2 males,

collected on 19.11.2017 by Nedko Nedyalkov (NMNHS

1051, 1052). Paratype voucher numbers for ethanol-fixed

tissue samples (ZFMK-TIS-23033 to -23037 and ZFMK-

TIS-33755 to -33758), and for extracted DNA are given

in Appendix 2.

Bonn zoological Bulletin 67 (1): 41-57

Measurements of paratypes. Reported are mean+-

standard deviation (minimum—maximum, sample size).

Linear measurements are in mm, body mass in grams.

Body mass 58.31+4.06 (52.5-65, 8), head and body

126.7543.196 (124-134, 8), tail 27.8843.137 (2432, 8),

hind foot 16.91+0.861 (15.0-17.4, 7), condylobasal

length of skull 31.80+0.747 (30.3-33.1, 9), maxillary

tooth row 12.07+40.224 (11.8-12.4, 9), breadth of brain-

case 15.47+40.300 (14.9-15.9, 9), height of braincase

9.028+0.323 (8.6-9.5, 9), breadth of rostrum over ca-

nines 4.41+0.088 (4.3-4.5, 9), breadth of rostrum over

molars 8.50+40.158 (8.3-8.8, 9), greatest length of pelvis

23.98+0.564 (23 .3—24.7, 6), breadth of pelvis 7.82+0.366

(7.4-8.3, 6), length of humerus 13.93+0.186 (13.7—

14.2, 6), width of humerus 10.42+0.132 (10.2—10.6, 6).

Description. Zalpa martinorum n. sp. 1s of about the

same external appearance and body proportions as 7) eu-

ropaea (Miller 1912). The tail is rather short (18-26%

of head and body length) and densely covered by up to

6.5 mm long bristles (Fig. 3c). Pelage is dense and vel-

vety, 7.0—-8.0 mm long on the back, 4.5—6.0 mm ventrally.

Fur is blackish dorsally and slightly lighter and with slate

shades ventrally. A skin PMS 25632 has an irregular buff

strike (21 mm long and up to 3.5 mm wide) on the poste-

rior abdomen. Hairs around the eye are short and lighter,

whitish or buff. The tip of the snout is pink to grey, cov-

ered by short hairs. There is a reverse triangle of bare

skin behind the rhinarium. The tail is usually blackish.

Fore foot is 12.6-14.3 mm broad. The skull (Fig. 4, 5a)

shows no peculiarities and is of average size and shape

overall. Rostrum is moderately robust; width across the

canines accounts for 13.3—14.5% and across the molars

for 25.4—27.7% of condylobasal length. Maxillary tooth

©ZFMK

New mole from Thrace 47

Fig. 4. Skull and mandible of the type specimen of 7alpa martinorum sp. n. ZFMK 2017.1149. Scale bar = 5 mm.

Bonn zoological Bulletin 67 (1): 41-57 ©ZFMK

48 Boris Kry8tufek et al.

Fig. 5. Ventral cranium of species of Zalpa: (a) T. martinorum n. sp. ZFMK 2017.1149 (greatest length of skull = 31.2 mm), (b)

T. europaea ZFMK 2014.748 (35.8 mm), (c) 7) aquitania ZFMK 2005.194 (35.6 mm), (d) 7? occidentalis ZFMK 2005.301 (33.4

mm), (e) 7’ romana ZFMK 66.304 (37.7 mm), (f) 7. caeca ZFMK 2005.268 (31.2 mm), (g) 7. stankovici ZFMK 98.709 (34.1 mm),

(h) 7. levantis PMS 21658 (30.8 mm).

row equals to 37.3-38.9% of condylobasal length. The

braincase is rather deep and the height of neurocranium

makes up 26.9—30.3% of condylobasal length. The pos-

terior margin of palatine is usually anterior to the imagi-

nary line connecting the posterior alveolar margins of 3"

Bonn zoological Bulletin 67 (1): 41-57

upper molars, and the anterior border of the infraorbital

foramen is above the 2"! upper molar (Fig. 4).

All three morphotypes of the pelvis were recorded in

T. martinorum n. sp. The most frequent is the interme-

diate morphotype (n=13; own material combined with

data in Popov & Milchev 2001), followed by the cae-

©ZFMK

New mole from Thrace 49

Fig. 6. Lateral view of left upper molar in species of Zalpa: (a) T. martinorum n. sp. ZFMK 2017.1149, (b) T. europaea ZFMK

2007.006, (c) 7! aquitania ZFMK 2005.160, (d) 7 occidentalis ZFMK 2005.303, (e) 7? romana ZFMK 1972.202, (f) 7: caeca

ZFMK 1966.310, (g) 7) stankovici ZFMK 2005.326, (h) 7? levantis PMS 10650. Anterior is to the left. Not to scale. 1 — parastyle,

2 — paracone, 3 — mesostyle, 4 — protocone. Note that the parastyle is missing in 7’ martinorum sp. n. (1*).

coidal morphotype (n=10) and the europaeoid morpho-

type (n=2). The sezamoidal os falciformis 1s robust in its

proximal part.

Incisors are of decreasing size with 1* incisor being

nearly twice as large as 3™ incisor, which is the smallest.

The molars are robust, and the 3 molar is particularly

large. The 1“ upper molar entirely lacks the parastyle

(Fig. 6a). The 1* lower premolar has a prominent distal

cusp; the 4" lower premolar lacks metaconid, and the 3"

lower molar 1s always without hypoconulid while the en-

tocristid is present only exceptionally. Oligodonties were

present on three skulls out of nine studied: both 1* upper

premolars are missing in ZFMK-MAM-2017.1150, left

1*' upper premolar is missing in PMS 25632, and left 1*

upper and 2™ lower premolars are missing in PMS 25631.

Comparison. Morphologically, 7! altaica (subgenus

Asioscalopus) 1s peculiar by its large size, short tail, slim

skull (Fig. 2) and weak dentition with a reduced 1“ upper

molar (Stroganov 1957; Zaytsev et al. 2014). Differenc-

es between Asioscalopus and the subgenus TJalpa are so

Bonn zoological Bulletin 67 (1): 41-57

obvious that no comparison with 7? martinorum n. sp. 1s

required.

In Thrace, 7’ martinorum n. sp. can be safely differen-

tiated from 7: europaea by (1) a sealed palpebral fissure

(open in 7! europaea), (11) absence of parastyle on 1“ upper

molar (present in 7’ europaea; Fig. 6b), and (111) smaller

size. Ranges for the two species overlap only marginally;

dimensions of 7’ europaea are from Thrace in Bulgaria

and Turkey (Osborn 1964; Dogramac1 1989a,b; Vohralik

1991; Popov & Miltchev 2001): body mass (in grams)

43—65 in T. martinorum n. sp. (own material and data in

Dogramaci 1988) vs. 60-105 in T: europaea; condylobas-

al length (in mm) 29.3-—33.1 in 7. martinorum n. sp. (own

material and data in Osborn 1964; DoSramaci 1988; Pop-

ov & Miltchev 2001) vs. 32.4—37.0 in T: europaea; length

of pelvis (in mm) 21.5—25.0 in 7. martinorum n. sp. (own

material and data in DoSramaci 1989b; Vohralik 1991;

Popov & Miltchev 2001) vs. 24.9-30.1 in T. europaea,

length of humerus (in mm) 12.9-14.9 in 7? martinorum

n. sp. (own material and data in Vohralik 1991; Popov

& Miltchev 2001) vs. 14.5-17.8 in 7! europaea. Molars

©ZFMK

50 Boris Kry8tufek et al.

Bulgaria (1)

rare

O72, . Black Sea

7) |

Turkey

e a #

Esri, DeLorme, GEBCO, NOAA NGDC, and other contributors

“ zi” 28° 29”

Fig. 7. Records of Zalpa martinorum n. sp. in Bulgaria (BG) and Turkey (TR). Localities with known mitochondrial genetic identity

are shown as black circles. Elevation is parenthesized. Legend: 1 — BG, Mt. Strandzha, Zvezdets (295 m); 2 — BG, Mt. Strandzha,

Gramatikovo (210 m); 3 — BG, Mt. Strandzha, Vizitsa (274 m); 4-— BG, Mt. Strandzha, Kondolovo (298 m); 5— BG, Mt. Strandzha,

Fazanovo (106 m); 6 — BG, Mt. Strandzha, Velika (54 m); 7 — BG, Mt. Strandzha, Pismenovo (9 m); 8 — BG, Sozopol (50 m); 9 —

BG, Burgas, Chengene skele (sea level); 10 —- BG, Burgas, Tvarditsa, Uzungeren (4 m); 11 — BG, Valchanovo (300 m); 12 — BG,

Tsarevo (40 m); 13 — BG, Lopushna Reserve (250 m); 14— BG, Sinemorets, Silistar (40 m); 15 —TR, Kirklareli, Derekoy; 16—TR,

Kirklareli, Demirkoy; 17 — TR, Kirklareli, Pinarhisar; 18 — TR, Subasi; 19 — TR, Kagithane Dere; 20 — TR, Rumeli Hisar; 21 — TR,

Bahg¢ekoy. Corresponding references: localities 1-7, 9, 10, 12: own data; 8: Vohralik (1991); 11, 13, 14: Popov & Miltchev (2001);

15-17, 21: Dogramaci (1988); 18 — Collection H. Vierhaus (ZFMK); 19, 20 -NMNH. Localities with no numbers are unspecified

in Popov & Miltchev (2001). Museum vouchers from the localities 18—20 are labelled as 7’ caeca; published records were classified

as T: caeca levantis (pts. 15-17) or T: levantis (the remaining).

are relatively larger in 7. martinorum n. sp. (dimensions

of 3 upper molar: 1.46—-1.61 x 1.96—2.17 mm) than in

T. europaea (1.29-1.59 x 1.77—2.24 mm; specimens

from various parts of Europe). Pelvis in 7: europaea is

either europaeoidal or intermediate but never caecoidal

(Petrov 1971b; Popov & Miltchev 2001).

Talpa martinorum n. sp. is well characterized by the ab-

sence of a parastyle on the 1* upper molar (Fig. 6a). Para-

style is present in the majority of species of 7alpa, spe-

cifically in 7? europaea (Fig. 6b), 7: aquitania (Fig. 6c),

T. occidentalis (Fig. 6d), T: caeca (Fig. 6f), T. stankovici

(Fig. 6g), T. levantis (Fig. 6h), 7’ davidiana, T: talyschen-

sis, and 7! caucasica. Talpa romana, which clearly lacks

Bonn zoological Bulletin 67 (1): 41-57

the parastyle (Fig. 6e), can be reliably differentiated from

T. martinorum n. sp. by the robust rostrum (Fig. 5e) and

large 3" upper molar (1.67—2.20 x 1.87—2.72 mm).

Talpa martinorum n. sp. can be unambiguously set

apart from all the remaining species of the genus Jalpa

by the nucleotide sequence of the Cyt b gene.

Distribution. The distribution range of 7? martinorum

n. sp. extends along the south-western Black Sea coast

from Burgas in Bulgaria to Istanbul in European Tur-

key, covering Mt. Strandzha (also Strandja in Bulgari-

an; Istranca in Turkish) in Bulgaria and Turkey, and the

forested coastal belt between the southern foothills of

Mt. Istranca and the city of Istanbul (Fig. 7). Records

©ZFMK

New mole from Thrace |

tm |

* » 7 ta

. hia te

pi oe a

: ‘alate

» ae

——_

Fig. 8. Vladimir and Evgentya Martino near their home in

Top¢tidersko Brdo (part of the city of Belgrade), Serbia, in early

1930s. Courtesy by Georgy Bakhtadze.

associated with published Cyt 5 sequences are all from

Strandzha in Bulgaria. Zalpa martinorum n. sp. 1s para-

patric or allopatric with respect to 7’ europaea which is

widespread in the lowlands further west. For the distribu-

tion range of 7’ europaea see Osborn (1964), Dogramaci

(1989a), Vohralik (1991), and Popov & Miltchev (2001).

The identity of small blind moles on the Asiatic side

of the Bosporus strait (Osborn 1964) is not known. All

moles from north-western Anatolia sequenced so far have

the molecular profile of 7) /evantis (vicinity of Zongul-

dak; Bannikova et al. 2015) and show a distinct parastyle

on 1* upper molar (vicinity of Bolu and Bursa).

Habitat. 7alpa martinorum n. sp. is a fossorial mole of

similar habitat requirements as other species of the sub-

genus Jalpa (cf. Niethammer & Krapp 1990). Character-

istic mounds of unearthed soil (molehills) are a common

feature in Mt. Strandzha. Specimens were captured in me-

sic meadows, pastures and orchards on deep medium-tex-

tured and well-drained soils (zheltozem and cinnamonic

forest soils; Shishkov & Kolev 2014) in hilly regions, on

the edges of arable land and on abandoned fields, in light

black soil on low alluvial plains, in oak woodland and

forests of river floodplains (own data as well as Osborn

1964 and Popov & Miltchev 2001). Altitudinal range is

from near sea level up to at least 445 m.

Bonn zoological Bulletin 67 (1): 41-57

Etymology. 7a/pa martinorum n. sp. 1s an eponym

to Vladimir Emmanuilovich Martino (Bsagqumup

OmManywosny Mapruno, 1888-1961) and Evgeni-

ya Veniaminovna Martino (Esrenua BenuamuHosHa

MaptuHo, 1894-1979) née Stepanova (CremaHosa),

ethnic Russians who in 1920 escaped the October Rev-

olution by emigrating to the Kingdom of Serbs, Croats

and Slovenes (Kingdom of Yugoslavia since 1929). In

politically insecure and frequently violent Eastern Eu-

rope of the 20" century, the Martinos were refugees for

more than three decades. In 1949 they moved to Bulgaria

and in 1955 returned to Russia (at that time still Soviet

Union). Inspired by G. S. Miller’s (1912) “Catalogue of

the Mammals of Western Europe” they initiated mam-

mal research in south-eastern Europe and named, among

others, two Balkan endemics: Dinaromys bogdanovi (V.

Martino & E. Martino, 1922) (Martino & Martino 1922:

413) and Talpa stankovici V. Martino & E. Martino, 1931

(Martino & Martino 1931: 53) (Fig. 8). While Vladimir

had a degree in Biology from the University in Novoros-

stysk (1913), Evgeniya had no formal academic educa-

tion. Despite this, she attained competency in mammal-

ogy and successfully collaborated with her spouse, both

in the field and cabinet. Although they published several

joint papers (as V. and E. Martino), the contribution by

Evgentya is mainly ignored and she is nearly anonymous

today (cf. Beolens et al. 2009). At least nine subspecif-

ic names for mammals with the epithet martinoi (see

Appendix 3) were proposed by mammalogists between

1935 and 1971, and all are eponyms to Vladimir Marti-

no. With the name martinorum Nn. sp. we stress the equal

share by Evgeniya in the tandem “V. et E. Martino” and

correct the injustice done to her contribution in the past

decades. Along with Dorothea Bate (1878-1951) and Ga-

briele Neuhduser (1911-1998) Evgeniya was one of the

early women who studied the taxonomy of Palaearctic

mammals already between the two great wars. In addi-

tion to their publications, a renowned legacy of Vladimir

and Evgentya Martino is their meticulously prepared and

carefully labelled mammal collection, deposited primar-

ily in the Natural History Museum London and the Zo-

ological Institute and Museum of the Russian Academy

of Sciences in St. Petersburg. For biographies of Vlad-

imir Martino, see Mezentsev (1961), Paspalev (1962),

Pusanov (1962), Zimmermann (1962), Gus’ kov (1965),

Taranenko (1999), and Boreiko (2001).

DISCUSSION

During the Pleistocene climatic changes over the last ca.

2.6 My, temperate species have endured glacial maxima

in geographically restricted refugial areas. A legacy of

the repeated extinctions of northern populations on the

one hand, and the long-term persistence in southern refu-

gia on the other hand is a number of taxa with restricted

©ZFMK

52 Boris KryStufek et al.

distributions in the former glacial refugia (Hewitt 2000).

Contemporary biogeographic pattern of the genus Jalpa

obviously results from the allopatric evolution in such

constrained refugia. Only two mole species (7! altaica

and 7: europaea) have extensive distribution ranges in

temperate and southern boreal regions of Europe and

western Asia, while the remaining ten or so species oc-

cupy small areas along the southern edge of the distribu-

tion range of the genus (Bannikova et al. 2015). The new

species from Thrace, with its small distribution range, is

therefore well nested within a broader biogeographic pat-

tern of the western Palaearctic.

Mammals endemic to the Balkan Peninsula, which in-

clude a mole 7. stankovici and several rodents, are re-

stricted to the topographically rough south-west Balkans

(KryStufek 2004). Talpa martinorum n. sp. therefore

provides the first evidence on speciation of a mammal

in the extreme eastern part of the peninsula. Besides, our

new evidence also corroborates an earlier assumption on

the persistence of multiple independent refugia inside the

topographically complex Balkan refugium (cf. KryS8tufek

et al. 2007).

The new species shows no close phylogenetic links

with the Balkan endemic 7? stankovici nor with the Bal-

kan-Italian 7? caeca. Instead, 7’ martinorum n. sp. forms

a (weakly supported) clade in the Cyt-b tree contain-

ing also the widespread 7: europaea and two endemics

from Western Europe, 7) aquitania and T: occidentalis.

Biogeographically, it is straightforward to interpret this

finding. Inside the Balkan refugium there was no evolu-

tionary divergence of an ancestral mole to 7. martinorum

n. sp. and either 7’ stankovici or T: caeca. Instead, T. mar-

tinorum putatively originated from a small peripheral

fragment population of a taxon ancestral also to 7! eu-

ropaea. The process replicates itself in each of the three

major southern European refugia, namely with 7. aquita-

nia and T: occidentalis in the west, with a deeply diver-

gent phylogeographic lineage of 7. europaea in northern

Italy (Feuda et al. 2015), and finally in the eastern part

of the Balkan refugium with 7 martinorum n. sp. (this

study). The observed pattern fits well the hypothesis by

Bilton et al. (1998) on Mediterranean refugia as areas of

endemism rather than sources of postglacial northward

recolonizations. Based on molecular clock estimates of

divergences between 7? europaea, T. aquitania and T:

occidentalis (1.88-3.71 Mya; Bannikova et al. 2015;

Feuda et al. 2015; Nicolas et al. 2017b), the most recent

common ancestor to 7? martinorum n. sp. and T: euro-

paea must be of Early Pleistocene age or older. Late

Pliocene small moles from Varshets in north-western

Bulgaria, classified as 7. cf. /evantis, closely resemble

T: martinorum n. sp. by their small size and the 3" upper

molar with no parastyle and with an undivided mesostyle

(Popov 2004). Middle Pleistocene strata of Yarimburgaz

Cave near Istanbul, 1.e., inside the contemporary range

of the new species, also yielded a small species of mole

Bonn zoological Bulletin 67 (1): 41-57

tentatively identified as 7: /Jevantis (Santel & Konigswald

1998). Fossil evidence therefore provides strong support

for a long persistence in the eastern Balkan refugium of

a mole most similar to the current 7’ martinorum Nn. sp.

Molecular evidence (Bannikova et al. 2015; this study)

so far did not yield any support for the earlier assumption

that moles colonized southeast Europe via an intermit-

tent Bosporus land bridge during one of the low sea level

phases (Vohralik 1991; KryStufek & Vohralik 2001). The

alternative submergences and emergences of the Bospo-

rus Strait enabled migrations of mammals from Europe

to Asia (e.g., Glis glis; Helvaci et al. 2012), from Asia to

Europe (e.g., Microtus hartingi, KryStufek et al. 2009),

or in both directions (Crocidura leucodon, Dubey et al.

2007). As shown by sedimentological evidence, however,

the Bosporus land bridge emerged as a permeable barrier

only in the Middle Pleistocene (Kerey et al. 2004) and

therefore postdates the postulated split between 7) mar-

tinorum n. sp. and T: europaea (see discussion above).

Therefore, even if moles from Asia would colonize Eu-

rope via the Bosporus land bridge, they would face an

established congeneric species in the new habitats. How-

ever, such speculations should be taken with a grain of

salt. So far, none of the moles in the Bosporus region on

either side of the strait have been screened genetically. A

denser sampling in that particular part of Turkey is there-

fore a necessity for a more holistic understanding of mole

evolution in this biogeographically dynamic region.

Acknowledgements. We thank Jevgeni Shergalin for help with

biographic information on Vladimir and Evgeniya Martino,

and Georgy Bakhtadze for providing Fig. 8. Franc Janzekov-

i¢ ran analyses in SPSS Statistics, Mojca Jernejc Kodrié pro-

cessed the skeletal material of the new species, and Hannah

Petersen performed the wet-lab work. B. K. acknowledges

the curators who made the study of museum vouchers under

their responsibility possible for him: the late William Stanley

(FMNH), the late Svetoslav Obratil (ZMBiH), Linda Gordon

(NMNH), Paula Jenkins (NHML), the late Kurt Bauer and Frie-

derike Spitzenberger (NMW), Haluk KefelioSlu (OMU), and

Nataliya Abramson and Alexandra Davydova (ZIN). Alexandr

Pozdnyakov helped with references, Georgy Popgeorgiev with

Fig. 7, Jorg Adrian provided photographs shown in Figs 4 and

5, and Henning Vierhaus donated owl pellet material from his

own collection. B. K. acknowledges the financial support from

the Slovenian Research Agency (research core funding no.

P1-0255).

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APPENDIX 1

Comparative museum vouchers

Listed are museum vouchers with the appropriate collec-

tion numbers; see text for collection acronyms. Material

is itemized by country and sorted alphabetically accord-

ing to locality.

Talpa altaica Nikol’skiy, 1883 (Nikol’skiy 1883: 165).

Russian Federation — ZFMK-MAM-1987.751 and

1987.752 from Akademgorodok, Novosibirsk Oblast.

Talpa aquitania Nicolas, Martinez-Vargas & Hu-

got, 2017 (Nicolas et al. 2017a: 641). France — ZFMK-

MAM-2005.242 from Arles-sur-Tech, Departement

Roussilon. ZFMK-MAM-2005.156 to 2005.159 from

vicinity of Blaye, Departement Gironde. Spain — ZFMK-

MAM-2005.203 to 2005.2005 from vicinity of Barbadillo

de Pez, Province of Burgos. ZFMK-MAM-2005.201 from

Burguete, Province of Navarra. ZFMK-MAM-2005.202

from Cameros, Sierra de Cebollera, Province of Logro-

no. ZFMK-MAM-2005.189 to 2005.200 from vi-

cinity of Corconte, Province of Santander. ZFMK-

MAM-2005.160 to 2005.166, 2005.169 to 2005.175,

2005.177 to 2005.179, and 2005.181 from Ramales de la

Victoria, Province of Cantabria. ZFMK-MAM-2005.176

and 2005.180 from Lanetosa, Province of Vizcaya.

Talpa caeca Savi, 1822 (Savi 1822: 265). Bosnia and

Herzegovina — PMS 7239 from Cemerno. PMS 7478

from Lake Boratko jezero, Mt. Prenj. ZMS 293 (type

of Zalpa hercegovinensis Bolkay, 1925; Bolkay 1925: 1)

from Stolac. Greece — NHML 31.11.11.24 to 31.11.11.26

and 31.11.11.27 (type of Talpa olympica Chaworth-Mus-

ters, 1932; Chaworth-Musters 1932: 166), 31.11.11.28

to 31.11.11.31 from eastern slope of Mt. Olympus. Ita-

ly —- ZFMK-MAM-2005.260 to 2005.263, 2005.268, and

2005.269 from Abetone Pass, Province of Pistoia. PMS

9813 from Caramanico, Province of Pescara. ZFMK-

MAM-2005.266 and 2005.267 from vicinity of Colle di

Nava, Province Imperia. ZFMK-MAM-2005.264 and

2005.265 from Osiglia, Province of Savona. Kosovo —

PMS 7473 from Pavlov kamen, Mt. Sar planina. Mace-

donia — PMS 7475 from Derven Pass above Prilep. PMS

7474 from Magarevo, Bitola. Montenegro — PMS 7485

from Mt. Kom Vasojevic. ZFMK-MAM-1966.310 from

Mt. Lovéen. PMS 7523 from Moraéa. PMS 8131 from

Bonn zoological Bulletin 67 (1): 41-57

Nik8i¢. Switzerland — ZFMK-MAM-1965.065 from

Bergell, Canton Graubtinden.

Talpa caucasica Satunin, 1908 (Satunin 1908: 5). Rus-

sian Federation - NMW 19943 to 19945 from Psebaj,

Krasnodar krai, north-western Caucasus.

Talpa davidiana (Milne-Edwards, 1884) (Milne-Ed-

wards 1884: 1143). Iran — FMNH 111007 from 1 mile

south of Divandarreh, Kurdistan. FMNH 96421, 96423

and 96424 (type of Zalpa streeti Lay, 1965; Lay 1965:

227) from Hezar Darreh, Kurdistan. Turkey — FMNH

82136 and 82137, PMS 21503 from Bitlis, Tatvan. OMU

231 from Megabuti yaylasi, Hakkari. NMW 20326 and

20327 from Cilo-Sat-Mts., Mergan Zoma, Hakkari.

MNHN 1883.469 (type of Scaptochirus davidianus

Milne-Edwards, 1884) from Meydanekbez, Gaziantep.

OMU 166 and 232 from Otluca koyt, Hakkari. Israel

— NHML M15277 (type of TZalpa chthonia Bate, 1937;

Bate 1937: 399), M16092 from the Upper Pleistocene

layers of Tabun Cave.

Talpa europaea Linnaeus, 1758 (Linnaeus 1758:

52). Austria — ZFMK-MAM-2005.143, 2005.146, and

2005.185 from Kleinalm, Lower Tauern, Styria. Bos-

nia and Herzegovina — PMS 21511 from Gornji Malo-

van. Bulgaria —- ZFMK-MAM-1938.146 from Bansko,

Mt. Pirin. Germany — ZFMK-MAM-2005.342 from

Bonn-Kessenich, NRW. ZFMK-MAM-2005.334 from

Bonn-Lengsdorf, NRW. ZFMK-MAM-2007.006 from

vicinity of Munster, NRW. ZFMK-MAM-2009.021

from Wahner Heide, Niederrheinische Bucht, NRW.

ZFMK-MAM-2014.748 from Welldorf, Dueren, NRW.

ZFMK-MAM-2002.138 from Wesseling-Urfeld, North

Rhine-Westphalia (NRW). ZFMK-MAM-2005.139

from Angeln, Schleswig-Holsteinisches Htgel-

land, Schleswig-Holstein. ZFMK-MAM-2005.153

from Grofhansdorf, Schleswig-Holstein. France —

ZFMK-MAM-2005.084 and 2005.085 from St. Tro-

pez, Pampelonne, Departement Var. Italy — ZFMK-

MAM-2005.086 from vicinity of Albenga, Province of

Savona. ZFMK-MAM-2005.089 from Dego, Province of

Savona. ZFMK-MAM-2005.066, 2005.074, 2005.075,

and 2005.328 from Merano, Province of South Tyrol.

ZFMK-MAM-2005.090 to 2005.093 from Pass Colle

di Nava, Province of Imperia. ZFMK-MAM-2005.087

from Pass Col di Sistriere, the Cottian Alps (Alpi Cozie),

Region of Piedmont. ZFMK-MAM-2005.088 from Pi-

etralunga, Province of Perugia. Montenegro — PMS 393

from Lubnice, Mt. Bjelasica. Serbia — ZIN 33974 (Type

of Zalpa europaea pancici V. Martino, 1930; Martino

1930: 60) from Kraljevo. Slovenia — PMS 25634 from

Mt. Pohorje.

©ZFMK

56 Boris Kry8tufek et al.

Talpa levantis Thomas, 1906 (Thomas 1906: 416). Tur-

key — NHML 25.11.1991 (type of Talpa caeca levan-

tis) from Altindere, Trabzon. NHML 6.5.1.1 to 6.5.1.4

from Cosandere, Trabzon. NHML 6.3.6.6 from Euthey,

Trabzon. NHML 6.3.6.4, NMNH 327252 and 327253

from Meryemana, Trabzon. OMU 233, 234, and 236 to

242 from Bitlis, Tatvan. NMVW 13042 and 13043 from

Bolu. PMS 10299 from Kiurtler, Samsun. PMS 21658

from 10 km east of Savsat, Ardahan. NMW 19858 from

Ulubey, Ordu. NMW 19859 from Yavuz-Kemal, Bicik,

Giresun. PMS 11372 from Sehitlor Gegidi, Mts. Giresun

Daglar1, Tamdere. NMNH 327263 and 327264, PMS

10650 from Mt. Ulu Dag, Bursa.

Talpa occidentalis Cabrera, 1907 (Cabrera 1907: 212).

Spain — ZFMK-MAM-2005.304 from 4 km south-west

of Cantoral, Province of Burgos. ZFMK-MAM-2005.279

and 2005.280 from 1 km north-east of Cazorla, Province

of Jaén. ZFMK-MAM-2005.271 to 2005.275 from Es-

pinama, National Park Picos de Europa, Province of Can-

tabria. ZFMK-MAM-2005.290 to 2005.294, 2005.297,

and 2005.298 from vicinity of Espinosa de los Monteros,

Province of Burgos. ZFMK-MAM-2005.276 to 2005.278

from 7 km south-west of Lanetosa, Puerto de los To-

mos, Province of Santander. ZFMK-MAM-2005.308

and 2005.309 from vicinity of Las Rozas, Province of

Santander. ZFMK-MAM-2005.281 to 2005.286 from

10 km south of Rascafria, Province of Segovia. ZFMK-

MAM-2005.299 to 2005.301 from Llanares de la Reina,

Province of Leon. ZFMK-MAM-2005.301 and 2005.302

from 10 km south of Riano, Province of Leon. ZFMK-

MAM-2005.314 from near Samosierra, Province of

Segovia. ZFMK-MAM-2005.310 to 2005.313 from Sier-

ra de Gredos, Province of Avila. ZFMK-MAM-2005.270

from Paso di Somosierra, Sierra de la Guadarrama, Prov-

ince of Segovia.

Bonn zoological Bulletin 67 (1): 41-57

Talpa romana Thomas, 1902 (Thomas 1902: 516). It-

aly — PMS 9855 from Caramanico, Province of Pes-

cara. ZFMK-MAM-1977.675 from Catanzano, Prov-

ince of Calabria. ZFMK-MAM-1966.291 to 1966.302,

1966.305, 1966.306, and 2005.335 from Monte Garga-

no, Province of Puglia. ZFMK-MAM-1966.303 from the

National Park Abruzzo. ZFMK-MAM-1966.304 from

Pescasseroli, Province of L’ Aquila.

Talpa stankovici V. Martino & E. Martino, 1931

(Martino & Martino 1931: 53). Greece — ZFMK-

MAM-2005.322 to 2005 .326 from Corfu, Ionian Islands.

ZFMK-MAM-205.327 from 7 km south of Ioannina,

Epirus. ZFMK-MAM-1963.680 and 1977.031 from Mt.

Olympus. Macedonia — ZIN 34014 (type of Talpa roma-

na stankovici) from Magarevo, Bitola. PMS 7488 from

Derven Pass above Prilep. PMS 7491 to 7493 and 7502

from Mt. Galicica. ZFMK-MAM-1998.709 and PMS

7497 from Mt. Bistra. PMS 7499 from Mt. Kajmak¢alan.

PMS 7486 and 7487 from Prilep. PMS 7496 and 7503

from Popova Sapka, Mt. Sar planina. PMS 7504, 7505,

and 7507 from Resen. PMS 7495 and 7497 from Stru-

ga. Montenegro — PMS 3202, 3203, 3205 (type of Tal-

pa stankovici montenegrina KryStufek, 1994; KryStufek

1994: 14) and 3207 from Ulcinj.

Talpa talyschensis Vereschchagin, 1945 (Vereschchagin

1945: 67). Iran— Ghilan, FMNH 96416 to 96419 from 12

km west of Chalus.

©ZFMK

New mole from Thrace 57

APPENDIX 2

GenBank accession numbers. Specimens of 7alpa martinorum n. sp. sequenced in this study. List includes collecting

data for the new sequences and GenBank accession numbers. See text for collection acronyms and Fig. 7 for locality

numbers (Loc. No.).

NMS BGI 42.04736 | 27.64744

NMS BG2 42.04697 | 27.64782

ZFMK-MAM- 2017.1150

Collection No. Loc. | Latitude | Longi- Field | DNA voucher

No. tude number | Tissue voucher

ZFMK-MAM- 2017.1149 42.09068 | 27.47637

42.12798 | 27.59862 | B ZFMK-DNA-FC19476382 | MH093594

ZEMK-TIS-23036

GenBank No.

B ZFMK-DNA-FC19476453 | MH093591

B ZFMK-DNA-FC19476461 | MH093592

B ZFMK-DNA-FC19476469 | MH093593

ZFMK-MAM- 2017.1151 4 42.09424 | 27.68921 BG10 | ZFMK-DNA-FC19476390 | MH093595

ZFMK-TIS-23037

ZFMK-MAM- 2017.1152

42.19826 | 27.73069 BG14_ | ZFMK-DNA-FC19476398 | MH093596

ZFMK-TIS-33755

PMS 25631 7 | 42.19739 | 27.79592 | BGI5 | ZFMK-DNA-FC19476406 | MH093597

pene? ee pe ee Lenmcrmanse |

PMS 25633 5 | 42.2287 |27.70733 | BGI7 | ZFMK-DNA-FC19476422

APPENDIX 3

Eponyms to (Vladimir) Martino

Names are arranged chronologically. Species name as is

currently in use (ex Wilson & Reeder 2005) is in square

brackets when appropriate.

Pitymys subterraneus martinoi Ehik, 1935 (Ehik 1935:

60) [Microtus subterraneus]|

Sumeriomys guentheri martinoi Petrov, 1939 (Petrov

1939: 363) [Microtus guentheri|

Rhinolophus ferrumequinum martinoi Petrov, 1940 (Pe-

trov 1940: 59)

Arvicola terrestris martinoi Petrov, 1949 (Petrov 1949:

186) [Arvicola amphibius]

Bonn zoological Bulletin 67 (1): 41-57

Mustela erminea martinoi Ellerman & Morrison-Scott,

1951 (Ellerman & Morrison-Scott 1951: 256)

Citellus citellus martinoi Peshev, 1955 (Peshev 1955:

290) [Spermophilus citellus|

Glis glis martinoi Miri¢c, 1960 (Miri¢ 1960: 36)

Spalax leucodon martinoi Petrov, 1971 (Petrov 1971a:

13P)

Lynx lynx martinoi Miri¢, 1978 (Mirié 1978: 30)

©ZFMK

BHL

Blank Page Digitally Inserted

Bonn zoological Bulletin 67 (1): 59-69

May 2018

Amphibian and reptilian records

from south-central Mali and western Burkina Faso

Wolfgang Béhme' & Jeffrey Heath” *

'Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D-53113 Bonn, Germany

?Department of Linguistics, University Michigan, Ann Arbor, Michigan 48109-1220, USA

* Corresponding author. E-mail: w.boehme@leibniz-zfmk.de

Abstract. Some amphibians and reptiles from southern Mali and southwestern Burkina Faso have been deposited in the

Zoologisches Forschungsmuseum Alexander Koenig (ZFMK) since the year 2000. Supplemented by some photographic

voucher material they contain several remarkable, in some cases even new records for the two respective countries. Malian

specimens were collected mainly in the region between Douentza and Bandiagara in the south-central part of the country,

while those from Burkina Faso are from the Bobo Dioulasso area in the southwestern part of the country. Three amphibian

species are new for Mali’s faunal list (Leptopelis bufonides, Ptychadena trinodis, Pyxicephalus sp. as the first represen-

tative of its genus). Two specimens of a small-sized Agama species cannot yet be assigned to a described species with

certainty but are in any case of zoogeographical interest. A specimen of Chamaeleo gracilis documents the second record

of this species for Burkina Faso, the first one in the western part of the country. The slender blind snake Leptotyphlops

albiventer is recorded for the first time for this country.

Key words. West Africa: Mali, Burkina Faso; herpetofauna, first country recordss.

INTRODUCTION

While the forests of the Upper Guinean region of West

Africa have been the focus of much herpetofaunal atten-

tion in recent decades (e.g., Penner et al. 2011), many

areas of landlocked West Africa remain under-surveyed.

Some work has been done on a number of these countries,

for example, Guinea-Bissau (Aulltya et al. 2012), Senegal

(Joger & Lambert 2002), Guinea (B6hme 2000, Rodel

et al. 2004, Greenbaum & Carr 2005, Hillers et al. 2008,

Bohme et al. 2011); relatively little, however, has been

published on the herpetofauna of Mali and Burkina Faso

when compared with their southern neighbours. For Mali,

Joger & Lambert (1996, 1997) provided an annotated

checklist and analysis of the diversity and biogeography

of the herpetofauna, while Bohme et al. (1996) provided

some additional records for both countries. R6del (2000)

described the West African frog fauna and listed coun-

try records for each species, including Mali and Burkina

Faso. Chirio (2009) gave records for the geographic tri-

angle formed by Niger, Benin and Burkina Faso where

the border-crossing “W Transfrontier Biosphere Reserve”

(“Réserve de Biosphere Transfrontaliere du W”, RBTW)

is situated. More recently, Trape & Mané (2006) and

Trape et al. (2012) provided updated distribution maps

at a one-degree scale for snakes and other reptiles respec-

tively, for all West African countries. Mediannikov et al.

(2012) revised the genus Agama all over West Africa with

an integrative approach which is, however, partly in con-

flict with the results of Wagner et al. (2009) and Leaché

Received: 05.02.2018

Accepted: 27.06.2018

et al. (2014). Finally, Trape & Mané (2017) published a

summarising paper on the snake fauna of Mali.

Here, we report on some herpetological voucher mate-

rial from Mali and Burkina Faso (Fig. 1) that has entered

the Zoologisches Forschungsmuseum Alexander Koenig

(ZFMK) in Bonn after 1996, since the publication of a

first note on this topic happened 22 years ago (Bohme et

al. 1996).

Our Malian records were made by Jeffrey Heath and

two native collaborators (Seydou Moro and Oumar Per-

gourou, see below) in September 2009, July 2010 and

on various occasions in 2011 and 2012 in the Dogon

Province, southern Mali, at Sévaré near Mopti (14°32’N,

04°06’W) and in the following villages of the area be-

tween Douentza and Bandiagara:

— Anda near Douentza (14°49’N, 03°01’W) is nested at

the base of a small rocky hill (inselberg). Between the

inselbergs are sandy fields, but there are also some creeks

and waterholes so that there are several microenviron-

ments (Fig. 2).

— between Douentza (15°00’N, 01°22’W) and Boni

(15°07’°N, 01°22’W) on pasture ground with wetlands;

— Pergué (14°85’N, 03°02’W) near Douentza (Fig. 3)

which is on a rocky shelf forming part of an inselberg. In

the outskirts of the village, to the south of the inselberg,

there are sandy plains.

— Kikara (15°12’N, 02°44’W) near Douentza, on the

northern slope of Gandamia inselberg (750 m a.s.l.),

mountain top.

— Koporo-Pen (14°08’N, 03°11’W), sandy plains east of

Bandiagara.

Corresponding editor: P. Wagner

60 Wolfgang Bohme & Jeffrey Heath

py, Ouagadougou

fo ( @%

BURKINA \

ere FA 50.

BURKINA FASO

Ouagadougou

* COT

* D'IVOIRE

Fig. 1. Map of Mali and Burkina Faso. Inset: Study area. Localities (Mali): 1. Mopti; 2. Sévaré; 3. Bandiagara; 4. Koporo Pén; 5.

Pergué village; 6. Anda village; 7. Douentza; 8. Kikara village; 9. Boni; (Burkina Faso); 10. Bobo Dioulasso; 11. 10 km SE Tiéfora;

12. Dérégoué; 13. Karankasso Vigué. Drawing: Morris Flecks.

Fig. 2. Anda village, Mali.

The area between Douentza and Bandiagara was visit-

ed several times, in 2009 (September), 2010 (July) and in

2011/2012, in the course of Dogon linguistic studies car-

ried out by Jeffrey Heath in the Dogon Province. Some

amphibians and reptiles were seen, photographed and

— by focal sampling — collected. These voucher speci-

mens are deposited in ZFMK’s herpetological collection,

as are the photographs in ZFMK’s herpetological photo

archive.

In 2015, J. Heath also visited Burkina Faso and was

able to gather some photographic vouchers of herpetolog-

Bonn zoological Bulletin 67 (1): 59-69

Fig. 3. Pergué village, Mali.

ical specimens which stemmed from Karankasso-Vigué

(10°61’N, 03°54’W), southeast of Bobo Dioulasso,

and from Dérégoué (locally also spelled Dérégboue;

10°45°N, 04°05’W, 295 m a.s.l.), 50 km east of Bobo

Dioulasso.

From the latter locality (4 km NE) we also received

some specimens collected in 1985 by Harald Schreiber

and co-workers, who was earlier the responsible curator

of the herpetological collection of the University of Saa-

rbricken (Paul Muller collection). Harald Schreiber and

co-workers collected alsol0 km SE of Tiéfora (10°58’N,

©ZFMK

Amphibian and reptilian records from south-central Mali and western Burkina Faso 61

04°33’W), in Bobo Dioulasso and in Ouagadougou it-

self. In 2010, the entire Paul Muller collection which had

initially been transported to the University of Trier, was

finally deposited in Bonn where it is now integrated in

the ZFMK herpetological collection (B6hme 2014). This

small part of the Paul Muller collection together with the

records made by J. Heath complements ZFMK’s pre-

vious holdings from Burkina Faso and Mali which had

been collected by Ulrich Joger, Harald Meier and Holger

Meinig. Their materials which contained also interesting

records including new ones for these two countries have

been published earlier (Joger 1979, 1981, Bohme et al.

1996). The present paper is an update of these earlier

contributions.

COMMENTED SPECIES LIST

AMPHIBIANS

Xenopus (Silurana) tropicalis (Gray, 1864)

Burkina Faso: ZFMK 93959965, 4 km NE Dérégoue,

gallery forest at Koba River.

The first record of this West African forest-dwelling

pipid species (Loumont 1984, Rédel 2000) from Burki-

na Faso was published by Bohme et al. (1996) based

on ZFMK specimens collected 1983 from Bobo Diou-

lasso by Harald Meier. Our newly acquired series from

Dérégoué corroborates the distribution of this zoogeo-

graphical outlier far from the forested coast in the arid

savannas of interior West Africa. However, the gallery

forest enables its survival in this environment, as it has

also been found in Senegambia (Bohme 1979) and north-

ern Ivory Coast (Comoé National Park: Rodel 2000).

Leptopelis bufonides Schiotz, 1967

Mali: ZFMK 93766—767, from between Douentza and

Boni, pasture ground with swamps.

Known from few localities from the open, relatively

dry savannas of West Africa, and seemingly patchily dis-

tributed (Hillers et al 2008). Joger & Lambert (1996) in-

dicated that it is likely to occur in Mali, since it is known

from Senegal and Burkina Faso, see also Rodel 2000).

However, our records listed here (Fig. 4) are apparently

the first confirmed ones for Mali.

Sclerophrys regularis (Reuss, 1834)

Burkina Faso: ZFMK 93968-969, 10 km SE Tiéfo-

ra, gallery forest at Sinlo River; ZFMK 101292-293,

Dérégoue.

A comment to be made on this widely distributed and

common species refers to the generic nomenclature. After

the partition of the collective genus Bufo, the Afrotropi-

Bonn zoological Bulletin 67 (1): 59-69

Fig. 4. Leptopelis bufonides from between Douentza and Boni,

Mali.

cal species were accommodated in an own genus Ami-

etophrynus (Frost et al., 2006), a name that has been in

use since then and has been established in numerous pa-

pers which means that nomenclatural stability was again

achieved. Recently, however, Ohler & Dubois (2016)

found an old specimen described as Sclerophrys capensis

Tschudi, 1838 to be assignable to Bufo (or respectively

Amietophrynus) rangeri, and synonymized the latter ge-

neric nomen with Sclerophrys. They were right in stating

that, despite only few citations, Sclerophrys was not a

nomen oblitum, so that this name has currently to be used

as the oldest available one for these toads.

Phrynobatrachus cf. latifrons Ahl, 1924

Burkina Faso: ZFMK 93966, 4 km NE Deérégoué, gal-

lery forest at Koba River.

The taxonomy of these little puddle frogs 1s complicat-

ed since a sibling species from the more forested regions

(P. accraensis) 1s involved. In addition, there are numer-

ous synonyms to be evaluated. These frogs are in need of

revision, so that our assignment above 1s tentative.

©ZFMK

62 Wolfgang Bohme & Jeffrey Heath

Fig. 5. Ptychadena trinodis from between Douentza and Boni, Mali. Left: ZFMK voucher; right: specimen not collected.

Fig. 6. Plate from Boettger (1881) showing the (presumably

lost) type specimens of his Rana (currently Ptychadena) trino-

dis (lower left) and Maltzania (currently Pyxicephalus) bufonia

(upper).

Bonn zoological Bulletin 67 (1): 59-69

Hoplobatrachus occipitalis (Gunther, 1859)

Burkina Faso: ZFMK 93958, 4 km NE Dérégoué, gal-

lery forest at Koba River; ZFMK 93967, 10 km SE Tié-

fora, gallery forest at Sinlo River.

A very common species in sub-Saharan Africa, enter-

ing both desert habitats in the north and forest habitats

in the south. Represented by voucher material from sev-

eral localities in the ZFMK collection (see Bohme et al.

1996).

Ptychadena trinodis (Boettger, 1881)

Mali: ZFMK 93770-771 from between Douentza and

Boni, pasture ground with swamps.

According to Frost (1985) distributed from Senegal to

the Democratic Republic of Congo, Mali being listed as

one of the countries with published records (Rodel 2000).

However, M.-O. Rédel (pers. comm.) kindly verified the

identification of our specimens (Fig. 5) and informed us

that they should actually be the first proven voucher spec-

imens for Mali. The single holotype of this species (Fig.

6), collected by Hermann and Agnes von Maltzan for the

Senckenberg Museum in Frankfurt am Main, seems to be

lost as it is not included in the type list of this collection

by Mertens (1967). For some details about the collectors

see below under Pyxicephalus.

©ZFMK

Amphibian and reptilian records from south-central Mali and western Burkina Faso 63

Fig. 7. Four specimens of Pyxicephalus sp. from between Mopti and Sévaré, Mali, to show the variability in color pattern. The

specimen on lower left is a juvenile (not on scale).

Pyxicephalus sp.

Mali: ZFMK 93763, 93772-773, Sévaré near Moptt.

Our new records are the first ones of this genus for

Mali and bridge a large geographical gap between Sen-

egal (Rufisque: Boettger, 1881; south of Rosso: Bohme

et al. 2001), Gambia (Frost 2017) and Mauritania (Abdul

Behru and Chlim: ZFMK vouchers: 76773 and 76781—

782 respectively) on the one hand, and Benin and Nigeria

(Rodel 2000, Nago et al. 2006) on the other. West African

Pyxicephalus are in high need of revision. Currently, the

West African populations of this frog are still assigned

to P. edulis Peters, 1854 (type locality Mozambique! )

(R6ddel 2000, Bohme et al. 2001, Nago et al. 2006, Frost

2015), but Rodel (2000) who listed as westernmost oc-

currence only Nigeria, regarded it as not unlikely that the

West African Pyxicephalus might not be edulis but a dis-

tinct species, citing an older personal communication by

one of us (WB). Boettger (1881) described a Senegalese

specimen as the sole member of his new genus Maltza-

nia from Rufisque in westernmost Senegal (Fig. 7) and

Bonn zoological Bulletin 67 (1): 59-69

assigned it to his new species M. bufonia. This is the

oldest and geographically closest name applicable to the

West African members of Pyxicephalus. Also Monard’s

(1951) name reiensis (from Rey Bouba, northern Camer-

oon) must be taken into consideration once a systematic

revision of Central and West African Pyxicephalus will

yield taxonomic differences between these two regions

(for the East African forms see Scott et al. 2013). But if

the Central and West African populations proved to be

identical, Boettger’s name would have high priority over

that of Monard (1951). Figure 7 documents the variable

colour pattern in the Malian population.

Boettger (1881) himself already regarded his often

overlooked new genus Maltzania as closest to Pyxiceph-

alus. He dedicated this name to Baron Hermann von

Maltzan (1843-1892) and his wife, Baroness Agnes von

Maltzan, who both had collected in Senegambia for the

Senckenberg Museum in Frankfurt am Main. The single

holotype of MZ bufonia (Fig. 6) evidently shared the fate

of the other new frog described by Boettger (1881) in

the same paper, viz. Ptychadena trinodis (see above),

©ZFMK

64 Wolfgang Bohme & Jeffrey Heath

and seems to be lost, since it is not included in Mertens’

(1967) type catalogue. Also, von Maltzan’s frogs can-

not be traced in the natural history museum of Waren/

Mecklenburg, the so-called Maltzaneum (today called

Muritzeum) which had been founded by him in 1866

(Gebhardt 1964, Hauff 2016).

Hildebrandtia ornata Nieden, 1907

Mali: ZFMK 93768-069, from between Douentza and

Boni, pasture ground with swamps.

Despite its large distribution area in the sub-Saharan

savanna belt, this species “is rarely encountered” (Rodel

2000), so our record is of faunistic interest.

Amnirana galamensis (Dumeéril & Bibron, 1841)

Mali: ZFMK 90468, between Douentza and Bandiagara.

A. galamensis is widely distributed in sub-Saharan

Africa, the typical form (type locality Lake Galam in

Senegal) being confined to West and Central Africa and

replaced by A. g. bravoana in eastern Africa und further

south. A photographic voucher from Burkina Faso is

mentioned in the appendix (see below).

REPTILES

Pelusios castaneus (Schweigger, 1812)

Burkina Faso: ZFMK 93957, 4 km NE Deérégoué, H.

Schreiber and B. Basten, I-IV. 1985.

The map sketch of Pelusios castaneus in Branch

(2008) leaves Burkina Faso outside the distribution range

of this species, while the text says only “from Senegal to

northwestern Angola, and inland to Central African Re-

public”. However, the latter country (CAR) is also far

outside the drawn range so that the small map sketches

by Branch (2008) are not reliable. Apart from a record

from the Burkina Faso part of RBTW (Chirio 2009), P.

castaneus is also registered in the grid map by Trape et al.

(2012) in the southwest of Burkina Faso, 1.e., in the area

of our voucher specimen.

Agama agama (Linnaeus, 1758)

Mali: ZFMK 91052, male, Douentza, J. Heath, VII.2010;

Burkina Faso: ZFMK 93950—955, 2 males, 1 female,

3 subadults, 4 km NE Déerégoué, near gallery forest of the

Koba River; ZFMK 93970, Ouagadougou.

The taxonomy of this widespread and anthropophilous

lizard is complicated, since it represents a species com-

plex of closely related forms. Moreover, its Linnean type

series, composed of three different species, has been dif-

ferently interpreted by Wagner et al. (2009) and by Medi-

annikov et al. (2012). In the light of the results by Leaché

et al. (2014) we follow the concept of the former authors.

Bonn zoological Bulletin 67 (1): 59-69

Fig. 8. Agama sankaranica from NE of Dérégoué (left) and

from Ouagadougou (right), Burkina Faso. Note the absence of

a light vertebral line in the right specimen.

Agama sankaranica Chabanaud, 1918

Burkina Faso: ZFMK 93956, 4 km NE of Dérégoué,

near gallery forest at the Koba river.

A former voucher specimen from Burkina Faso (ZFMK

39032) was the first country record of this species (B6hme

et al. 1996). The two voucher specimens (Fig. 8) differ in

that one (ZFMK 93956) has a white middorsal line as de-

scribed to be typical for this species (Trape et al. 2012),

the other belongs to the less common morphotype with-

out such a stripe. The specimens bridge a distributional

gap between two records in Mali and one each in NE

Ivory Coast and NW Ghana. From Burkina Faso, there is

only one earlier record on the map by Trape et al. (2012)

in the centre of the country. It may be noted that the east-

ernmost records of A. sankaranica in this map are situ-

ated in central Nigeria (Jos Plateau), ignoring a record

from Minim, Adamaoua Plateau, Cameroon (Bohme &

Schneider 1987) which extends the distribution range of

this species for more than 500 km further towards the

©ZFMK

Amphibian and reptilian records from south-central Mali and western Burkina Faso 65

Fig. 9. Agama sp. from Dérégoué, Burkina Faso. Left: male,

and right: female.

southeast. Recorded also for the Burkina Faso part of the

RBTW in the east of the country (Chirio 2009).

Agama sp.

Burkina Faso: ZFMK 93948-949, male and female,

4 km NE of Dérégoueé, near gallery forest at the Koba

River.

The two specimens (Fig. 9) seem to be interesting be-

cause they are small and nonetheless seemingly adult,

having a head-body length of only 69 and 65 mm respec-

tively. They are not yet assignable to a described species.

Since they also belong to the A. agama species complex,

their identity can only be cleared by extensive interspe-

cific comparisons for which the present faunistic study 1s

not the right place.

Chamaeleo gracilis Hallowell, 1842

Burkina Faso: ZFMK 101235, 4 km NE Dérégoué, gal-

lery forest at the Koba River.

Bonn zoological Bulletin 67 (1): 59-69

Although stated to be widely distributed in Africa, from

Senegal to Ethiopia and Tanzania (Trape et al. 2012),

there 1s no locality record for this chameleon in Burkina

Faso on the grid map of these authors, although it was

recorded from Burkina Fasoan part of the RBTW in the

easternmost part of the country and only one in southern

Mahi. From C. senegalensis which is known from several

localities in both countries, our C. gracilis female is dis-

tinguished by the shape of the casque and the rudimenta-

ry occipital flaps which are virtually absent in the former

species. It seems that our specimen is the second docu-

mented voucher specimen from Burkina Faso, next to a

record given by Chirio (2009) for the easternmost edge

of the country, just at the border to Benin which is not

included in the map by Trape et al. (2012). It documents

thus a much wider potential distribution in this country

than assumed before.

Chalcides ocellatus (Forskal, 1775)

Mali: ZFMK 90467, 91055—056) from Anda village be-

tween Douentza and Bandiagara.

The three voucher specimens of this widely distributed

species are the second record of this species from Mali.

Before, this skink was only once reported from Mali by

Greenbaum et al. (2006) from two localities, viz. Tom-

bouctou and between Kona and Sévaré (Joger & Lambert

1997, see also Trape et al. 2012).

Chalcides delislei (Lataste & Rochebrune, 1876)

Mali: ZFMK 93764, Pergué, sand dunes outside the vil-

lage.

A Saharan element with several records in south-cen-

tral Mali where also our voucher specimen comes from.

Leptotyphlops albiventer Hallermann & Rodel, 1995

Burkina Faso: ZFMK 97767, 4 km NE Deérégoué, near

gallery forest at the Koba River.

This species was described from the Comoé National

Park in northern Ivory Coast (Hallermann & Rodel 1995)

and subsequently recorded from Guinea Bissau and Mali

(Trape & Mané 2006, 2017). In Trape & Mané (2017) the

authors used the new generic name Myriopholis Hedges,

Adalsteinsson & Branch, 2009, created in the course of

a family-wide new molecular phylogeny by Adalsteins-

son et al. (2009). Myriopholis albiventer was, however,

placed in this genus without molecular data, and a mi-

cro-CT scan of ZFMK 97767 showed that its skull char-

acters do not fit the pattern of the other Myriopholis spe-

cies but rather resemble the situation in Leptotyphlops in

the strict sense (Claudia Koch, unpubl. data), so that we

maintain here its original genus name. Our specimen has

a total length of 120 mm, which is near the maximum

for this species, and the relations of its total length to tail

©ZFMK

66 Wolfgang Bohme & Jeffrey Heath

Fig. 10. Leptotyphlops albiventer from Dérégoué, Burkina

Faso.

length as well as to its body diameter, the midbody scale

count, the number of subcaudals and the color pattern of

a brownish dorsal and a whitish ventral side fit the char-

acteristics of this species as described by Hallermann &

Rodel (1995) and Trape & Mané (2006). ZFMK 97767

(Fig. 10) documents thus the fourth record from a fourth

country, but the first for Burkina Faso.

Lamprophis fuliginosus (Boie, 1827)

Burkina Faso: ZFMK 93971, Bobo Dioulasso.

A widespread, partly anthropophilous snake (vernacu-

lar name “house snake’) distributed in sub-Saharan West

Africa from the forest belt up to the Sahelian savanna

(Trape & Mané 2006).

Lamprophis lineatus (Duméril, Bibron & Dumeril, 1854)

Burkina Faso: ZFMK 101291, Bobo Dioulasso.

This close relative of the former species is known to be

broadly sympatric in West Africa including Burkina Faso

(Trape & Mané 2006).

Naja nigricollis Reinhardt, 1843

Burkina Faso: ZFMK 101547, Dérégoué (a mutilated

adult specimen).

Widely distributed in in the sahelian, sudanian and

guinean savanna types of West and Central Africa. In

both Mali and Burkina Faso N. nigricollis is represented

in most parts of these countries (see the map in Trape &

Mané 2006). According to these authors, West African

specimens are entirely black above and below, except

some light ventral crossbands below the neck and the an-

terior part of the body. ZFMK 101547 corresponds to this

characterization.

Bonn zoological Bulletin 67 (1): 59-69

Bitis arietans (Merrem, 1820)

Burkina Faso: ZFMK 101290, 10 km SE Tiéfora.

As the former species widely distributed in West Africa

including Burkina Faso (Trape & Mané 2006).

Echis leucogaster Roman, 1972

Mali: ZFMK 91055 (head and forepart only) 91056 (mu-

tilated subadult), Douentza.

This Sahelian species is common and widely distributed

both in Mali and in Burkina Faso. Our two specimens are

badly damaged and thus not suitable for providing scale

counts which would be necessary to corroborate their

identification against the sympatric congeners E. jogeri

and E. ocellatus (Trape & Mané 2006, 2017). However,

head and neck size of ZFMK 91055 and the unspotted

white underside in both specimens argue for E. /eucogas-

ter, aS does the locality Douentza which is situated in a

center of Malian records (Trape & Mané 2006).

APPENDIX: PHOTOGRAPHIC VOUCHERS

Some locality records which are based on photographs

only have been separated from the species list above

since the physical specimens are absent. However, in all

cases except Jarentola cf. ephippiata, their specific iden-

tification is indubitable.

Hemisus marmoratus (Peters, 1854)

Burkina Faso: Photo voucher (Fig. 11), Karankas-

so- Vigué, pasture ground with swamps.

The first record of this burrowing frog in Burkina Faso

was made by M.-O. Rodel in 1992 (Bohme et al. 1996)

from halfway between Ouagadougou and Bobo Dioulas-

so. This photographic record extends the range south-

westwards towards Ivory Coast where Rodel (2000) re-

corded it from the Comoé National Park. It has not yet

been listed for Mali so far.

Ptychadena schubotzi (Sternfeld, 1917)

Burkina Faso: Photo voucher from Karankasso-Vigué.

Ptychadena schubotzi is the savanna sibling of the

forest-dwelling P. /ongirostris with which it has often

been mixed up in the past (Perret 1981, R6del 2000). It

is widely distributed over the sub-Saharan savanna belt

countries and was first discovered in Mali by Schatti

(1986) and in Burkina Faso by Bohme et al. (1996).

Amnirana galamensis (Dumeéril & Bibron, 1841)

Burkina Faso: Photo voucher, Karankasso- Vigué.

Not to be mistaken and thus a reliable locality record.

©ZFMK

Amphibian and reptilian records from south-central Mali and western Burkina Faso 67

Fig. 11. Hemisus marmoratus from Karankasso-Vigué, Burkina

Faso.

Tarentola cf. ephippiata O’ Shaugnessy, 1857

Mali: Photo voucher from Douentza.

Tarentola ephippiata turned out to be a species com-

plex, and some forms described as subspecies have been

lifted to specific rank (Trape et al. 2012). According to

these authors, the Bandiagara-Douentza area has two

forms of this complex (7! ephippiata sensu stricto and 7:

hoggarensis) occurring in sympatry, thus corroborating

their specific status. The identification of these two close

relatives after a photograph only ts difficult, so the above

assignment remains tentative.

Tropiocolotes tripolitanus Peters, 1880

Mali: Photo voucher from Kikara near Douentza, near

mountain top, under stones.

This Saharan faunal element has a disjunct distribution

on the southwestern and northeastern margins of the Sa-

hara. In the southwestern part of its range, it is distributed

in Mauritania and Mali (Trape et al. 2012). The identifi-

cation of the photographed specimen as T° tripolitanus 1s

unproblematic since only two well distinguishable spe-

cies are possible to occur in the area.

Agama boueti Chabanaud, 1917

Mali: Photo vouchers from Douentza, Tupéré near Boni,

and Koporo Pén.

This ground-dwelling Agama was described after two

specimens from Gao on the Niger River in Mali and con-

sidered to be so rare that even its validity was questioned

(Wermuth 1967). But six decades later it was found to

be very common close to Dakar, Senegal (Bohme 1979),

and subsequently also recorded from the Air Mountains,

Republic of Niger (Joger 1981). Today it is known to be

continuously distributed in the Sahelian belt from Sene-

gal to Niger (Mediannikov et al. 2012, Trape et al. 2012),

Bonn zoological Bulletin 67 (1): 59-69

Fig. 12. Agama boueti from Douentza, Mali.

and our records from the Douentza area (Fig. 12) in Mali

fit well into this pattern.

Latastia longicaudata (Reuss, 1834)

Mali: Photo voucher, Douentza.

This long-tailed lacertid lizard has a wide sub-Saha-

ran distribution across the Sahelian savanna belt. For

West Africa, the grid map in Trape et al. (2012) shows

a concentration of records in western Senegal and north-

ern Cameroon, connected by scattered grid cells in Mali,

©ZFMK

68 Wolfgang Bohme & Jeffrey Heath

Fig. 13. Latastia longicaudata from Douentza, Mali.

Bonn zoological Bulletin 67 (1): 59-69

Niger and northern Nigeria. For Mali, there are only two

erid cells given (Trape et al. 2012) which evidently refer

to Goundam and south of Gao (see Joger and Lambert

1996) and their distance from each other is bridged by

our record (Fig. 13) from Douentza.

Acknowledgements. We thank Mark-Oliver Rodel and Mi-

chael Barej, (Museum fiir Naturkunde Berlin: ZMB), for com-

menting on frog specimens, Claudia Koch (ZFMK), for shar-

ing unpublished information on leptotyphlopid osteology, and

Morris Flecks (ZFMK) for valuable help with the figures. One

of us (JH) wants to thank Seydou Moro from Anda/Douent-

za, and Omar Pergourou from Pergué/Douentza for their hard

work to collect some voucher specimens. We are also grateful

for constructive reviewer’s comments.

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