Bonn zoological Bulletin 64 (2): 59-76

March 2016

Neglected Trichalophus (Coleoptera: Curculionidae):

DNA barcode and phylogeography of high-altitude flightless weevils

rediscovered in Southwest China

Vasily V. Grebennikov

Canadian Food Inspection Agency, K.W. Neatby Bldg., 960 Carling Ave., Ottawa, ON K1 A 0Y9, Canada

vasily.grebennikov@inspection.gc.ca; v_grebennikov@mail.ru

Abstract. Trichalophus LeConte weevils are rediscovered in Southwest China (Yunnan and Sichuan) after last being col-

lected in 1915. Populations are all found at high altitudes (3704-4158 m) and are attributed to three species: T. caudic-

ulatus (Fairmaire, 1886) (= compressicauda Fairmaire, 1887, syn. n.), T. scylla sp. n. and T. tibetanus (Suvorov, 1915).

Type specimens of all four species-group names are illustrated. A DNA barcode library of five Trichalophus species (29

sequences) is presented at doi: dx.doi.org/10.5883/DS-TRICHAL. All examined species of Trichalophus are flightless.

Phylogenetic relationships of Southwest China Trichalophus based on Maximum Likelihood and Maximum Parsimony

analyses suggest their monophyletic origin and monophyly of each species. Results of the temporal analysis are consis-

tent with the basic Quaternary expansion-contraction model of altitudinal range change. The warm period following the

Last Glacial Maximum (26,000-19,000 years before present) is linked to the present day high altitude Trichalophus refugium

in Southwest China, but not for the lineage diversifications, which are much older (8.08-5. 17 Mya). An illustrated overview

of ten extant Alophini genera is provided.

Key words. New species, new synonym, Yunnan, Sichuan, COl, Entiminae, Alophini, Father Delavay

INTRODUCTION

The exclusively Holarctic weevil genus Trichalophus

LeConte, 1876 consists of 51 valid species (Yunakov

2013). Adults of Trichalophus are believed to be flight-

less and brachypterous (Anderson 1997; Fig. 1). Loss of

flight ability is commonly observed in a number of unre-

lated edaphic, alpine, high latitude, subterranean, island,

or litter- inhabiting insects. Examples from weevils include

Macaronesian Laparocerus Schonherr, 1 834 (Machado et

al. 2008), VoXynQsmnRhyncogonus Sharp, 1885 (Macha-

do 2007), predominantly Oriental Trigonopterus Fauvel,

1862 (Riedel 2011), Middle American Theognete Cham-

pion, 1902 (Anderson 2010), an assemblage of likely un-

related Old World genera historically placed in Molytini

(Grebennikov 2014b) and the Western Palaearctic genera

of former “Cryptorhynchinae (Lyal 2014) traditionally

linked to Acalles Schoenherr, 1825 (Astrin & Stiiben

2008). The reduced dispersal capacity predisposes such

evolutionary lineages to become hostages of their habi-

tats and, therefore, subject to more frequent bottleneck ef-

fects. These conditions favour allopatric speciation (Grant

& Grant 2006; Ikeda et al. 2012; Vogler & Timmermans

2012) seemingly accompanied by an accelerated rate of

DNA evolution (Bromham 2008), as compared to their

flight-capable relatives (Mitterboeck et al. 2013). Such bi-

ological characteristics make flightless weevils a model

group for phylogeographical studies on such dynamic ter-

Received: 03.02.2015

Accepted: 08.09.2015

rains as oceanic islands (Tanzler et al. 2014; Toussaint et

al. 2015) or mountaintops (Grebennikov 2014a), if it were

not for their often highly inadequate taxonomy. The lat-

ter, if not updated, either lacks names for newly detected

evolutionary lineages (= new species), or has too many

names for the same clade (= synonyms), or a combina-

tion of both. In such situations the historical burden of Lin-

naean names impedes, rather than advances evolutionary

studies (Riedel et al. 2013a, b).

The genus Trichalophus has a trans-Beringian distribu-

tion range, with species found on the Pacific sides of both

Asia and North America, evoking the Bering land bridge

hypothesis (Berman et al. 2011). All but one species are

restricted to either Asia or North America; the exception

being the Nearctic T hylobinus (LeConte, 1876) recent-

ly reported from North Korea (Yunakov 2013). Trans-

Beringian distributions are commonly observed in a num-

ber of weevil genera: Alaocybites Gilbert, 1956 (Greben-

nikov 2010), Thalasselephas Egorov & Korotyaev, 1977,

Emphyastes Mannerheim, 1852, Lepyrus Germar, 1817,

Lobosoma Zimmermann, 1964, or Lepidophorus Kirby,

1837 (Egorov et al. 1996; Bousquet et al. 2013). Indeed,

all but one (Yunakov et al. 2012) Palaearctic records of

Trichalophus pertain to Siberia, Mongolia, Central Asia

and the northern part of the Pacific Asia including Japan,

while the Nearctic species are predominantly found in

Alaska, the western USA, and Canada west of Ontario

(Anderson 2002; Bousquet et al. 2013). Yunakov (2013)

Corresponding editor: D. Ahrens

Vasily V. Grebennikov

listed 43 Trichalophus species and one non-nominal sub-

species {T. vittatoides striola Reitter, 1913) for the

Palaearctic region, while Anderson (2002) mentioned eight

North American congeners. Since then the Nearctic diver-

sity of Trichalophus has lost two species names due to syn-

onymy (T. seriatus Mannerheim, 1843 and T. brunneus

Van Dyke, 1927) and has gained two others through the

recently synonymized genus Acmaegenius LeConte,

1 876 (Bright & Bouchard 2008). As a result, eight Nearc-

tic species are currently recognized (T. arcuatus Fall, 1907,

T. hylobinusTQConiQ, 1876, T. planirostrisTQCQniQ, 1876,

T. seminudus Van Dyke, 1938, T. granicollis Van Dyke,

1927, T. didymus LeConte, 1854, T. simplex LeConte,

1 876, and T. alternatus Say, 1 832), the latter four being

recorded from Canada (Bright & Bouchard 2008).

Despite the relatively large size of these beetles and their

occasional abundance in suitable habitats, biological da-

ta on Trichalophus are remarkably scarce. Adult beetles

appear highly polyphagous (Anderson 2002) being found

on a number of shrubs and herbs (personal observation).

North American T didymus was mentioned as an occa-

sional pest of strawberries (Fragaria sp.) in Washington

State (see references in Bright & Bouchard 2008). Imma-

ture stages and larval host plants are adequately known

only for a widely distributed Siberian species T leucon

Gebler, 1841. Larvae of this species feed externally on the

roots of Ribes L. (Grossulariaceae) as well as on a few oth-

er shrubs and take two years to complete development

(Krivets & Burlak 1986; Krivets 2006). The host plant

record is of potential economic significance, since the host

genus includes cultivated currants and a number of orna-

mental plants. The genus Ribes also includes alternate

hosts for the White Pine Blister Rust (Cronartium ribico-

la J.C. Fischer, Cronartiaceae), a fungus accidentally in-

troduced to North America about 1900 from Europe or

Asia, which causes significant damage to American white

pines (Pinus spp.; Maloy 2001).

Nothing is known about the evolutionary history of

Trichalophus and the phylogenetic relationships of this

taxon. Flightlessness is not unique for Trichalophus, but

it is found in some other genera of Alophini (Bright &

Bouchard 2008), including the West Palaearctic Graptus

Schoenherr, 1 823 (Davidian & Arzanov 2004). The latter

genus has long been known under its synonymous name

Alophus Schoenherr, 1 826 and was widely used in origi-

nal combinations for Palaearctic Trichalophus prior to Re-

itter ’s generic revision (1913). No members of either

Trichalophus or any other Alophini were subjected to a

phylogenetic analysis so far. The taxonomic recognition

of either the genus or the tribe, along with a few diagnos-

tic characters used in the keys (i.e. Anderson 2002) are,

therefore, the only hints of their possible monophyly. Since

the genus Trichalophus was historically linked to Grap-

tus, they both might fomi a clade, even if paraphyletic with

respect to the North American Plinthodes LeConte, 1 876

(Bright & Bouchard 2008) and perhaps other oligotypic

Holarctic genera of Alophini {sensu Alonso-Zarazaga &

Lyal 1999, with subsequent modifications of Bright &

Bouchard 2008 and Alonso-Zarazaga et al. 2010; see al-

so below). DNA data for Trichalophus are exceptionally

scarce, with only four public partial COl sequences (>400

bp) of T alternatus presently available from either Bar-

code of Life Database (BOLD) or GenBank.

LeConte (1876) established the genus for six nominal

Nearctic species known to him, four of them having been

described earlier as Alophus (A. didymus, A. constrictus

LeConte, 1857, A. alternatus, A. seriatus) plus two new-

ly described (A. simplex, A. planirostris). Six other cur-

rently valid Palaearctic species (Yunakov 2013) described

prior to 1 876 were added later; one of them was original-

ly described as Hypsonotus Germar {H. boeberi Schoen-

herr, 1826) and five others as Alophus (A. albonotatus

Motschulsky, 1869, A. humeralis GqHqy, 1834, A. linea-

Gebler, 1841, A. quadrigiittatus GqHqx, 1829, A. rud-

is Boheman, 1 842). After 1 876, the Palaearctic part of the

genus grew quickly in size. By the year 1915 Trichalo-

phus included all but six of its 44 currently valid Palaearc-

tic species-group taxa (Yunakov 2013). This notable in-

crease was mainly due to the efforts of Johannes K.E.

Faust and Edmund Reitter who, together with a few oth-

ers, introduced 30 currently valid species-group names de-

scribed from specimens collected on the Asian frontiers

of the rapidly growing Russian Empire (Siberia, Russian

Far East, Turkmenistan, Tajikistan, Uzbekistan, Kyrgyzs-

tan, Kazakhstan, Mongolia, Xinjiang Uyghur Autonomous

Region of China; Pierce 1960; Bajtenov 1974). Oddly

enough, two wq^n Alophus species, A. caudiculatus and A.

compressicauda, were reported by Fairmaire (1886,

1887, respectively) from geographically distant Yunnan

situated on the extreme southwest of China. The latter

records, if indeed belonging to Trichalophus, would ex-

tend the genus’ range for over 1,000 kilometers south-

wards. In 1913 Reitter described 13 species-group taxa in

a key to all Palaearctic species known to him. Until now

Reitter’s revision has remained the most comprehensive

single publication on Trichalophus weevils.

For the following hundred years the genus was neglect-

ed and only six new species-group names were introduced.

Among them are both species recorded from Japan: T

rubripes Zherikhin & Nazarov, 1990 and T nutakkanus

Kono, 1936; the former also found on the continent, while

the latter is endemic to Hokkaido. Two other species were

named from the mountains of the former Soviet Central

Asia {T. lixomorphus Bajtenov, 1974 and T krauseanus

Bajtenov, 1975), while one {T. korotyaevi Zherikhin &

Nazarov, 1990) was described from Sakha Republic

(=Yakutia). Additionally, Suvorov (1915) established a

monotypic genus Pseudalophus for his new species P. ti-

betanus described from an unknown number of syntypes

collected during Pyotr K. Kozlov’s (1863-1935) Mongol-

Bonn zoological Bulletin 64 (2): 59-76

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Neglected Trichalophus: DNA barcode and phylogeography of high-altitude flightless weevils in Southwest China 61

#601 a

Tibetan Plateau

Sichuan

Basin

lyanmar

Mongolia

Russian

Far East

India

approximate southern edge of known

continuous Trichalophus range in Asia

weevils Is Southwest China:

1 . "Kundur-Tschu rivef : T. f/iwlanus, type locality

2. Mt. Gonnga; T. tibetanus

3. Gang Shan: J. caudicuiatus

4. Mt. Haba; T. oaudicutatus

5- Songpan: T. sp, n. & 7

?. not localized in "Yunnan*: T. compfassicauda

Kazakhstan

Kyrgyzstan

stub of

hind wing

Gap of at least 1,000 km

Fig. 1. Known geographic distribution of Trichalophus is Southwest China. Inserted image of female specimen #60 1 8 from Kaza-

khstan shows hind wing brachyptery in Trichalophus.

Tibetan expeditions 1899-1901. The published temporal

and geographical data of this species are identical with

those of Notaris kozlovi Korotyaev, 1979 (Grebennikov

& Kolov, unpublished) and the type locality is in the pres-

ent day extreme northwest of Sichuan (Fig. 1). The four-

line description of the new genus hinged on two charac-

ters distinguishing it from ''Alophus'\ namely (a.) elytra

strongly compressed “internally” (i.e. as if a force was ap-

plied in the horizontal plane therefore flattening elytra in

the vertical plane) and (b.) elytral surface “naked” (= lack-

ing vestiture). The former character is known to occur in

Palaearctic Trichalophus and was used by Fairmaire

(1887) to derive the name compressicauda for a species

from nearby Yunnan, while the latter character might per-

haps be attributed to abrading. Suvorov’s original descrip-

tion was suggestive of a Trichalophus species, and, not

surprisingly, Yunakov (2013) synonymized both generic

names.

This project began on May 19, 2010, when the first two

Trichalophus specimens (#0713 & #0714, Figs 3, 5),

among those reported below were found under stones in

the alpine zone of the Gang Shan Mountain Range in Yun-

nan (Fig. 1). The find was most inspiring and seemingly

supportive of Fairmaire’s historical claim that the genus

was present so far south. During the next two years addi-

tional specimens were recovered in the same and three oth-

er high altitude localities in Yunnan (Mount Haba) and

Sichuan (Mount Gongga and Songpan, the latter seeming-

ly supporting two sympatric species). Phenetic similari-

ties and subsequent analysis of DNA barcodes suggested

that those were indeed species of Trichalophus. The wide

gap seemingly separating these Trichalophus of Southwest

China from their congeners in the north (Fig. 1) became

partly bridged when the former ''Pseudalophus” ti-

betanus was transferred to Trichalophus (Yunakov 2013).

At that stage it became evident that Trichalophus was in-

deed present in Yunnan and Sichuan. The discovery of the

high-altitude and the extreme southern representatives of

a widely distributed trans-Beringian genus suggests a refu-

gial distribution since the last glacial period (Darwin,

1 859: 373-382). The goals of the present paper, therefore,

are (1.) to attempt unfolding the evolutionary past of the

newly sampled Trichalophus specimens from Southwest

China using mainly mtDNA data and (2.) to report these

findings in the framework of ranked Linnaean classifica-

tion. Additionally, an attempt is made to provide an illus-

trated overview of all ten extant genera of Alophini and,

therefore, to bring attention to this poorly defined, and tax-

onomically disorganized weevil tribe.

MATERIAL AND METHODS

Museum abbreviations, followed by the curator’s name:

CNC Canadian National Collection of Insects, Arach-

nids and Nematodes, Ottawa, Canada (P.

Bouchard)

IZCAS Institute of Zoology, Chinese Academy of Sci-

ence, Beijing, PR. China (R. Zhang)

MNHN Museum National d’Histoire Naturelle, Paris,

France (H. Perrin, A. Mantilleri)

Bonn zoological Bulletin 64 (2): 59-76

©ZFMK

Vasily V. Grebennikov

Fig. 2. Habitats of Trichalophus spp. in Southwest China. A-C; T. scylla sp. n., Songpan, Sichuan; D-F: T. tibetanus, Songpan,

Sichuan; G-I: T. caudiculatus. Gang Shan, Yunnan; J-L: T. caudiculatus, Mt. Haba, Yunnan.

Bonn zoological Bulletin 64 (2): 59-76

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Neglected Trichalophus: DNA barcode and phylogeography of high-altitude flightless weevils in Southwest China 63

scyits sp. n.

#4437 V

#4438 M

#4439 M

#4440 F

#4441 M

#5385 F

#5386 F

#5387 M

#5388 F

Ffidwifop/tus (jiwlBinius

Songpan

o-Sb^,7a

#2742 F

#2743 M

#2744 F

#2745 M

'Songpien

#4418 M

#4419 IVt

#4436 HHOLOTYPE

FnohfffctphfJS Sp. r-

#6713 M

#0714 F

#2767 F

#2763 F

ICsrg Shafi

#4624 M

1 \ #4625 M

FxSbaitipfMrs esMfcuffltus

Ww1/0 ^

Haba'

0,81 ro, 64

#4620 F

#4627 M

#5464 M

#5405

#5466 F

#2960

T aKemaSus

Tfictmkipttiji csiiiiiajtBliiS

vMble 4

(-slemite 8)

venble 5

(=s1emite 7)

#5383 F

#0714 F

#5406F

Fig. 3. Maximum Likelihood inference phylogram of Trichalophus weevils from Southwest China using the 658 bp of the mtD-

NA barcoding COl gene fragment. The tree is rooted on Graptus circassicus (Entiminae: Alophini; not shown); two extraterrito-

rial outgroup Trichalophus (#2960 & #2968) are in grey. Digits at intemodes are ML/MP bootstrap values. Six geographical evo-

lutionary groups of Trichalophus are marked as clades A-F in white squares; note that Mount Haba and Songpan each harbours

two evolutionary groups. Black long arrows link respective terminal clades with an image of their representative. Specimen num-

bers in bold are those of imaged males (M, to the left of the tree) or females (F, in the insert showing ventral view). Black and

white short arrows indicate morphological characters and, after a slash, their states (Table 1).

Bonn zoological Bulletin 64 (2): 59-76

©ZFMK

Vasily V. Grebennikov

MTD Senckenberg Naturhistorische Sammlungen,

Dresden, Germany (K.-D. Klass, O. Jager)

ZIN Zoological Institute, Russian Academy of Sci-

ences, St. Petersburg, Russia (B.A. Korotyaev)

The length of the body was measured in dorsal aspect from

the elytral apex to the anterior edge of the pronotum. Dis-

tribution map (Fig. 1) is generated using the on-line Sim-

pleMappr tool (Shorthouse 2010). The chronostrati-

graphic timing follows Cohen et al. (2013). Nomenclature

of male genitalia follows that of Wanat (2007). The term

“Southwest China” is delimited to two Chinese provinces,

Yunnan and Sichuan. The term “base pair” is abbreviat-

ed as bp when referring to sequence length; abbreviations

“syn. n” and “sp. n.” denote new synonym and new

species, respectively.

Specimen sampling, handling and

gathering DNA data

Except for two specimens #4418 and #4419 sifted from

Rhododendron L. leaf litter, all newly collected Trichalo-

phus were handpicked from under stones (Figs 2A-L) in

the alpine zone (Fig. 2J), or on a glade in the upper for-

est zone (Fig. 2D) in Southwest China. In total 38 adult

Trichalophus beetles were collected in the following four

localities (Fig. 1 ; in brackets are the total number of spec-

imens followed after a slash by the number of those suc-

cessfully sequenced for DNA barcode >400 bp): the Cang

Shan Mountain Range (4/4), Mount Haba (12/7), Mount

Gongga (4/4), and the vicinity of Songpan township

(1 8/12). A leg was removed from a specimen for DNA ex-

traction. All specimens used for DNA barcoding have at

least one unique identifier label with the code CNC-

COLVGOOOOXXXX; this format is shortened to the last

four digits #XXXX when a specimen is referred to (Figs

3,4). Specimen images, geographical data, primers, orig-

inal electropherograms and other relevant data pertaining

to all 35 matrix-forming sequences can be seen online in

the publicly accessible dataset ''Trichalophus 35 [DS-

TRICHAL]” on the Barcode of Life Database portal (doi:

dx.doi.org/10.5883/DS-TRICHAL). Genitalia of six males

each representing a terminal cluster (=evolutionary group)

as detected in the phylogenetic analyses (see below) were

dissected, imaged (Fig. 3) and stored in microvials with

glycerol pinned with the specimens.

DNA analyses and matrix constrnction

Three separate DNA analyses were performed. The Max-

imnm Likelihood (ML) and the Maximnm Parsimony

(MP) analyses attempted to place the diversity of

Table 1. Discrete morphological characters for diagnostics of Trichalophus weevils in Southwest China (Fig. 3)

1. Elytral shoulders and elytral sides in basal 2/3, dorsal view; shoulders rounded, sides evenly widening posterad (0); shoulders

angular, sides subparallel (1).

2. Elytral dorsal and posterior profile (=dechvity), lateral view; evenly and gently rounded throughout (0); flattened dorsally and

abmptly curved (1).

3. Female, ventrite 5, bumps and depressions on surface, ventral view; absent (0); present (1).

4. Female, ventrite 4, two sharp points at posterior edge, ventral view; absent (0); present (1).

5. Female, posterior projections of elytral apices, ventral view: absent (0); present (1).

6. Male genitalia, long, thin, and curved apical lamella of aedeagus (lateral view); absent (0); present (1).

7. Male genitalia, aedeagus, dorsal view: symmetrical (0); asymmetrical (1).

8. Male genitalia, aedeagus, notch in lateral outline, dorsal view; absent (0); present (1).

9. Male genitalia, sternal apodeme 9, dorsal view; thin (0); thick (1).

clade

species

Locality

tibetanus

Songpan

tibetanus

Mt. Gongga

scylla sp. n.

Songpan

caudiculatus

Cang Shan

caudiculatus

Mt. Haba

caudiculatus

Mt. Haba

Bonn zoological Bulletin 64 (2): 59-76

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Neglected Trichalophus: DNA barcode and phylogeography of high-altitude flightless weevils in Southwest China 65

Trichalophus from Southwest China into phylogenetic and

geographical perspective. Both ML and MP used the same

matrix of 35 DNA barcodes with a minimum and maxi-

mum length of 400 bp and 658 bp, respectively. The in-

group consisted of 27 Trichalophus specimens from

Southwest China, while the outgroup included two extra-

territorial Trichalophus specimens representing T rubripes

from the Russian Far East and T alternatus from Cana-

da (Fig. 3). The rest of the outgroup was formed by six

specimens of Graptus circassicus Solari, 1945 from Geor-

gia: Abkhazia. Both ML and MP analyses were imple-

mented using MEGA 6 (Tamura et al. 2013), including (a.)

topology building, (b.) statistical support test by using

1000 repetitions of bootstrapping and (c.) search for the

best substitution model for ML analysis (T92+G). The root

was consistently placed between Graptus and Trichalo-

phus. The GenBank accessions for these 35 sequences are

KM538655-86, KJ445708, KJ445709, KJ445712; all of

them are new, except for the three latter Graptus se-

quences.

The third DNA analysis was performed to date the in-

group branching events and to re-test the phytogenies sug-

gested by the ML and MP analysis. The original matrix

of 35 sequences was reduced to include only 23 full-length

DNA barcodes (658 nt; except for the sequence of T al-

ternatus with 609 bp). The second analysis was performed

in BEAST vl.8.0 (Drummond et al. 2012) utilizing the

Bayesian inference (BI) approach with no a priori group-

ing, all default priors and options, GRT+G+I nucleotide

substitution model (instead of the T92+G+I not offered in

the software; the latter model was detected in a separate

model- searching analysis in MEGA 6 as having the best

fit), strict linear molecular clock and nucleotide substitu-

tion rate of 0.018 (Papadopoulou et al. 2010). Tracer 1.6

(Rambaut et al., 2014) was used to graphically determine

stationarity and to check convergence of runs. The “burn

in” option was implemented eliminating the first 2500 of

the 10000 obtained trees. The resulting topologies from

each of three analyses (ML, MP, BI) were visualized in

FigTree vl.4 (Rambaut et al. 2014).

Contribution from morphology

Morphological data are not expected to contribute deci-

sively in DNA-dominated phylogenetic analysis, partic-

ularly in shallow branches of the tree of life (Ward 2011)

conventionally called “species” in ranked classification

(Hey 2001). Consequently, no effort was made to merge

the DNA matrix with a few morphological characters

scored for the ingroup (Table 1). Instead, DNA-determined

Trichalophus clusters (= evolutionary groups by Hey 2001

or cladesA-F on Fig. 1) were a posteriori scrutinized in

search of diagnostic morphological characters (Maddison

2014), not necessarily synapomorphic (Ward 2011). The

easily observable dorsal color pattern, being either too

variable or subject to abrasion, was judged as unreliable

for diagnostic purposes in Trichalophus. An additional ef-

fort was made to explore structures of male genitalia by

dissecting a single male per each of six clades detected

on the phylogenetic tree (Fig. 3).

Integrating molecular phylogenetic results

into taxonomy

In biodiversity studies taxa are normally named first and

then their phylogeny and boundaries are analysed, if ever.

This approach, although logically awkward, has strong his-

torical roots from the times when (a.) phylogenetic theo-

ry was not practiced by taxonomists, and (b.) researchers

lacked adequate data to perform sufficiently detailed

analyses when naming their new species. Advent of Hen-

nigian principles coupled with availability of DNA se-

quences challenges this classical and logically deficient

approach (Ward 201 1). A modern student of biodiversity

is expected to (A.) delimit evolutionary groups through a

formal analysis, then (B.) make a balanced, responsible

and subjective judgement using all available evidence

sources as to which clades need names (Hey 2001) and

then (C.) conservatively apply formal names, either pre-

existing or newly proposed. In other words whenever pos-

sible, taxa naming should not be done before but after the

analysis and discussion, not to abuse logic by putting the

cart before the horse. Through most of the present paper

all six tree-delimited clades representing candidate species

(Fig. 3) are referred to by using informal non-taxonomic

names (clades A-F, italicized). Therefore, the taxonomic

part of this paper using three valid species-group names

(two previously used and one new) and synonymizing one

name follows the Results section and most of the Discus-

sion.

RESULTS

The Maximum Likelihood (ML) analysis produced the

best tree (Fig. 3) with the highest log likelihood of

-3265.92. All Trichalophus specimens from Southwest

China formed a weakly supported clade with highly re-

solved internal structure consisting of six clades A-F. The

Maximum Parsimony (MP) analysis resulted in seven best

trees (length: 558, consistency index: 0.64, retention in-

dex: 0.90; topologies are not shown) also recovering the

same six clades A-F (as in Fig. 3). In both analyses the

ingroup was recovered as a clade. The only backbone

topological difference of the MP strict consensus tree (as

compared to the ML topology. Fig. 3) was that the clade

C was recovered as the sister to the rest of Trichalophus

from Southwest China.

Bonn zoological Bulletin 64 (2): 59-76

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Vasily V. Grebennikov

Pliocene Pleistocene

T rubhpes

10 6 0

Fig. 4. Ultrametric tree dating evolutionary events of Trichalophus beetles in Southwest China. Digits at nodes and on the scale

below are million years before present. Node bars represent 95% confidence intervals of the age estimate.

Temporal analysis in BEAST (Fig. 4) recovered all

Trichalophus specimens from Southwest China in a clade

with the same internal backbone topology as in the ML

analysis (Fig. 3, although some specimens and the entire

clade E were not represented in the BEAST analysis due

to inadequate sequence length). The inferred timing of the

origin of the Southwest China Trichalophus is 1 1 .65 MY,

while the clade’s diversification leading to the five clades

A-D and clade F range between 8.08 MY and 5.17 MY

(Fig. 4).

DISCUSSION

mtDNA phylogeny and phylogeography

of Trichalophus in Sonthwest China

Recovery of a monophyletic Trichalophus radiation in

Southwest China (Figs 3, 4) should be treated with cau-

tion, since limitations in the number of the in- and out-

group representatives did not permit a rigorous test. Lit-

tle other evidence is available to challenge this hypothe-

sis. The entire clade A-F has a compact range allopatric

to that of the rest of the genus (Fig. 1), although the dis-

junct distribution might be plausibly attributed to the lack

of adequate sampling effort to bridge the gap. Morpho-

logically Trichalophus beetles from Southwest China are

seemingly large-bodied compared to other two species

(Fig. 3), although adding more Trichalophus in the analy-

sis might challenge this pattern. Nearly all of the ingroup

morphological characters (Table 1) cannot be adequately

matched with those for the rest of the genus due to the lack

of comparative data. Summing up, compact distribution

(Fig. 1) of monophyletic Trichalophus of Southwest Chi-

na (Figs 3, 4) is a weakly supported hypothesis to be even-

tually retested. Internal structure of six ingroup clades (Fig.

3) is mainly consistent with limited geographical (Fig. 1)

and morphological (Table 1) data and is further discussed

when Linnaean species are delimited (see below).

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Syntypa of

Syn^M of coimressvcaLic

Vi totl .

IbiJiia Firli

:»u.

It. C(f*rtliiiT

^ l«2

rOHiln

(X&pytj 1

Vfi. $tt^O-rQV duL

Fig. 5. Type specimens and original labels of three historical Trichalophus species names from Southwest China {caudiculatus,

compressicauda, tibetanus currently assigned to two valid species T. caudiculatus and T. tibetanus), together with three T. caudic-

ulatus females (#0714, #2767, #27678) sequenced for DNA barcode (Fig. 1). Black and white arrows indicate morphological char-

acters and their states (Table 1, separated by a dash). Images of caudiculatus and compressicauda syntypes and their labels: An-

toine Mantilleri, © MNHN.

Unlike at least some of its more northern congeners,

Trichalophus in Southwest China inhabit high altitudes

(3704-4158 m). Such a characteristic of the southern-most

representatives of a temperate northern hemisphere clade

of low-dispersing organisms is consistent with the basic

Quaternary expansion-contraction model of latitudinal

range change (Qiu et al. 2011). The latter stipulates ex-

tensive latitudinal range shifts in the form of southward

movement during glacials followed by rapid expansions

northwards during interglacials (Qiu et al. 2011). If so, dis-

tribution of Trichalophus in Southwest China (Fig. 1) is

a direct result of the last warming following the Last Gla-

cial Maximum (26,000-19,000 ybp). More specifically,

the observed data are consistent with at least three sub-

hypotheses (numbering after Qiu et al. 2011): (iii) long-

term isolation and survival in multiple localized refugia

(cladesA-D and clade E+F in Fig. 3), (ii) population iso-

lation and endemism due to river course dynamics {clade

D versus clade E+F) and (iv) glacial in situ survival of

some hardy alpine species on the Tibetan plateau itself

(population of T tibetanus represented by the type spec-

imens, Fig. 5). Like the hypothesis on monophyly of

Trichalophus in Southwest China, all phylogeographical

inferences are highly preliminary due to material and da-

ta limitations.

With no suitable fossils to calibrate a Trichalophus mo-

lecular clock, the temporal aspect of Trichalophus evolu-

tion in Southwest China (Fig. 4) relies on the a priori mtD-

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Vasily V. Grebennikov

NA substitution rate of distantly related Tenebrionidae (Pa-

padopoulou et al. 2010). The obtained dates of the

Trichalophus cladogenesis (Fig. 4) are comparable to those

of the sympatric and similarly high-altitude and flightless

weevil genus Niphadomimus Zherikhin, 1987 (Greben-

nikov 2014a). Both agree that the lineage divergence took

place well before the onset of the Pleistocene climate fluc-

tuations. Both time estimations were based, however, on

identical methods and substitution rates, which might have

biased them both. Exact substitution rates may significant-

ly vary depending on population size, founder effects, and

a number of other less well-understood factors and, there-

fore, markedly differ from the assumed 0.018 substitutions

per site per million years. In other words application of

the user-friendly BEAST software and the obtained clear-

cut dates (Fig. 4) should be treated carefully, since we are

far from understanding the molecular clock (Eanfear et al.

2010), even for such relatively simplified and recent sce-

nario as that of Trichalophus weevils in Southwest Chi-

na.

Delimitation of Linnaean species for Trichalophus in

Southwest China

The practical task of assigning Einnaean species names

to the newly discovered Trichalophus from Southwest

China, even with the help of a tree (Fig. 3), is far from

being trivial. It involves at least one theoretical and two

practical difficulties. First, imposing ranked classification

on the continuum of the tree of life cannot be objectivised

(Hey 2001 ; Ward 2011) and, therefore, involves arbitrary

decisions (Sites & Marshall 2004). Second, despite sam-

pling and analytical efforts, relatively little data on

Trichalophus are available. For example, the relatively

well-resolved DNA tree (Fig. 3) is that of a very short frag-

ment of a fast-evolving mitochondrial maternally inher-

ited gene and, therefore, only a proxy to organismal evo-

lutionary history. Third difficulty is the existence of three

available historical names, which have nomenclatorial pri-

ority and have to be interpreted and used, if considered

as valid. These three issues have to be considered before

the freshly sampled Trichalophus in Southwest China

might be incorporated into the existing taxonomic scheme

(Yunakov 2013). The first difficulty, concerning the issue

of what a Trichalophus “species” is, will be resolved in

this passage, while both practical issues are considered fur-

ther below.

A “species” as a taxonomic category is purely and sole-

ly a label routinely and often inconsistently assigned to

the shallow branches of the tree of life ever since Einnaeus.

As such, “species” is no more real than other taxonomic

categories like “genus”, “family” or “phylum” (Hey 2001).

Eike every other taxonomic category, a “species” is a mere

convenience required by the human mind to categorize and

count biological diversity. Acceptance of this basic philo-

sophical and methodological principle denies “species” re-

ality in the same sense at it denies reality “genera” or oth-

er “higher” taxonomic categories. In practical terms a

species is nothing more than a morphologically (or oth-

erwise) diagnosable group of organisms preferably form-

ing a clade and, most importantly, considered practically

worthy of being called a species (Hey 2001). If to follow

such an approach, decision on species boundaries almost

fully rests with the revising author, which, in turn, results

in the splitters versus lumpers issues, particularly in re-

gard to a clade of allopatric populations. The authors nor-

mally feel free to either split them into as many species

as possible, or lump them into a single one. The first ap-

proach is an example of unnecessary taxonomic inflation

(Isaac et al. 2004) adding nothing but unnecessary names.

The alternative lumping approach using a single species

name and a geographic reference would label every al-

lopatric evolutionary group equally well, while avoiding

unnecessary additions to the already heavy nomenclato-

rial burden. Moreover, a scramble to call a “species” an

allopatric evolutionary group, even if accompanied by cor-

relating morphological and other differences, can be mis-

leading when linked to the phenomenon of Sisyphean evo-

lution (McKay & Zink 2014). Summing up, introduction

of new species names should be done only when all al-

ternative options have been shown as inadequate. Such a

careful and reserved approach would not have created the

multitude of meaningless and cryptic taxa (Riedel et al.

2013a, b; Vences et al. 2013).

Two among six terminal clades representing Trichalo-

phus evolutionary groups (Fig. 3) are allopatric to all oth-

er ingroup clades {clade B and clade D from Mt. Gong-

ga and Gang Shan, respectively). They should, therefore,

be first assessed if each of them can be merged together

with its strongly supported sister-group into a more inclu-

sive clade to merit a species name. Indeed, clade B is

strongly linked to the allopatric clade A (Fig. 3), so the

clade A+B might itself be considered a candidate species.

Clade C is recovered in the ME analysis as a sister to clade

A+B (Fig. 3), but in MP analysis the sister group of clade

A+B was the clade D+E+F. Remarkably, specimens of

both clade A and clade C occur in sympatry in Songpan

(or at least in parapatry; their two geographically closest

samples were taken a few hundred meters apart). More-

over, while males and females of both clade A and clade

B share all eight morphological character states (Table 1),

their males differ in three genitalia characters from those

of clade C. Such considerations strongly suggest that

clades A+B and clade C (Fig. 3) should be treated as two

separate species, respectively, and the former one as con-

sisting of at least two geographically and morphological-

ly unique evolutionary groups {clade A and clade B, Fig.

3). It is possible that in the future each clade A and clade

B might be considered as separate species, but not until

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the current nomenclatorial lumping arrangement is refut-

ed as impractical.

The clade D+E’+F emerges as the third and the last one

to be designated as a Linnaean species among those rep-

resented in the ingroup (Fig. 3). Remarkably, clade E and

clade F from Mount Haba, although genetically distinct,

are formed by morphologically and geographically indis-

tinguishable specimens (Fig. 3). This notable genetic di-

morphism accompanied by full sympatry and morpholog-

ical similarity is perhaps linked to incomplete lineage sort-

ing (Funk & Omland 2003). The clade E+F is strongly

supported as a sister to the geographically closest clade

D and both share all but one morphological character

(Table 1). Like the clade A+B, the clade D+E+F 3)

might also later require two Linnaean species, but not be-

fore the present conservative nomenclatorial decision is

shown as inadequate.

Matching historical names with the tree-delimited

Trichalophus Linnaean species

The most significant practical constraint is how to link

three clades delimited for designation as Linnaean species

(clade A+B, clade C and clade D+E+F, Fig. 3) with three

available historical names (T. caudiculatus, T. compres-

sicauda and T. tibetanus). The type specimens of the lat-

ter are well preserved and available for study (Fig. 5).

Matching the type specimens of three historical names

with three evolutionary groups in Fig. 1 can be done us-

ing three sources of evidence: (a.) similarity in body shape

and in male genitalia, (b.) geographical proximity and (c.)

biological characteristics expressed through the altitude

of the type locality. No attempt was made to extract DNA

from the type specimens, mainly because they were judged

too old to warrant an attempt.

Of the three historical names, only the type specimen

of T. tibetanus has information available from all three

sources. Its type locality can be traced precisely (Figs 1,

5), while habitus and shape of male genitalia of a syntype

(Fig. 5) match most closely those of the clade A+B (Fig.

3). The name T. tibetanus is, therefore, used to designate

the clade A+B (Fig. 1). Such matching is far less straight-

forward with the both Fairmaire’s names.

The most significant uncertainty with both names cau-

diculatus and compressicauda is that their type localities

are imprecise, originally given as “Yunnan”. The years

when the types were collected are also unknown. Only the

younger of these two names, compressicauda, has the col-

lector’s name stated: Pere Jean Marie Delavay (=Father

Delavay). It seems, however, plausible that Father

Delavay also collected the type series of caudiculatus.

Even though Fairmaire cited Armand David (= Father

David) as the type specimen source for species described

together with caudiculatus in the 1886 paper. Father

Delavay was likely the original collector of at least some

of them, since Fairmaire in the same paper also named Ci-

cindela delavayi Fairmaire, 1886 in his honour. Indeed,

the years before both species were described (1 886, 1 887)

correspond with Delavay’s second stay in China

(1882-1891) (Anonymous 2014). This trip took place af-

ter Father Delavay’s meeting with Father David in 1881,

the latter convincing the former to collect specimens for

the Museum national d’histoire naturelle (Anonymous

2014). During his second stay in China, Delavay was

mainly based in a Yunnan village somewhere between

Lake Erhai and Eijiang township given as “Dapingzi”

(Anonymous 2014). Delavay had two favourite climbing

spots nearby: “Mount Heishanmen” (or “Ma’an Shan”,

west of “Dapingzi”, see Handel-Mazzetti 2014; not def-

initely located, but distinct from the Cang Shan Mountain

Range) and the Cang Shan Mountain Range along the

western shore of lake Erhai (Eancaster 1993, Anonymous

2014). Besides these two high mountain localities. Father

Delavay is definitely loiown to have collected in the alpine

zone around Deqin in northwestern Yunnan, from where

numerous alpine Carabus were sampled (T. Deuve, per-

sonal communication). It is highly probable that Father

Delavay visited many other alpine localities in Yunnan,

however those three were apparently most frequently vis-

ited and/or sampled for high altitude beetles.

Both caudiculatus and compressicauda were described

from an unknown number of syntypes. Curatorial search-

es in MNHN in 2014 revealed a single syntype for each

for these names (Antoine Mantilleri, personal communi-

cation). Their syntype status is corroborated by the fact

that both specimens bear original identification labels and,

moreover, fully agree with relatively detailed descriptions

of both nominal species (Antoine Mantilleri, personal

communication). Both syntypes, although not dissected,

appear to be females by possessing posterior projections

on elytral apices (character 5/1, Fig. 5) and posterior pro-

jections of ventrites 4 (character 4/1, Fig. 5). These two

syntype characters match with those of the female spec-

imens from clade D (Fig. 3); the other being angular ely-

tral shoulders and subparallel elytra in their basal 2/3

(Table 1). Specimens from clade D inhabit the Cang Shan

Mountain Range, which was visited on many occasions

by Father Delavay, including altitudes above 4,000m (Ean-

caster 1993). Both Fairmaire’s names, therefore, represent

the best fit for the Einnaean species represented on Fig.

3 by the clade D+E+F, which, in turn, suggests their syn-

onymy proposed below.

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Vasily V. Grebennikov

Taxonomic overview of Trichalophus

in Sonthwest China

Trichalophus LeConte, 1876

Type species: Alophus didymus LeConte, 1854, fixed by

subsequent designation (Bright & Bouchard 2008: 57).

Trichalophus caudiculatus (Fairmaire, 1886)

(Figs 2, 3, 5)

caudiculatus YdLixmmxQ 1886: 353 {Alophus)

Type locality. Yunnan.

Type specimens. Syntype (MNHN, Fig. 5), likely female,

examined from images in Fig. 5, labels in Fig. 5. De-

scribed from unknown number of syntypes.

= compressicauda Fairmaire, 1887: 129 {Alophus) syn. n.

Type locality. Yunnan.

Type specimens. Syntype (MNHN, Fig. 5), likely female,

examined from images in Fig. 5, labels in Fig. 5. De-

scribed from unknown number of syntypes.

Diagnosis. This species is recognized by presence of api-

cal elytral projections in females (character 5/1, Fig. 3).

Intraspecific variation. GenBank accessions:

KM538662, KM538665, KM538667-68, KM538670-71,

KM538676-77, KM538681-82, KM538684. Length:

12.3-14.5 mm (Gang Shan), 11.1-11.8 mm (Mount Ha-

ba, clade E) and 11.4-14.5 mm (Mount Haba, clade F).

Specimens from both sampled localities (Fig. 1) slightly

differ in dorsal coloration, shape of elytral shoulders and

arrangement of posterior projection on ventrite 4 of fe-

males (Fig. 3). Aedeagus of a single male known from the

Gang Shan Mountain Range is less curved and more elon-

gate, as compared to those of two males dissected from

Mount Haba (Fig. 3), while female posterior projections

on elytral apices of the Gang Shan specimens are notice-

ably longer than those from Mount Haba (Fig. 3).

Additional material examined. 16 exx in total: 2 exx

#0713-0714 (GNG) “P.R. GHINA, Yunnan, E slope Gang-

shan at Dali, N25°39’54.7” E100°06’04.5”, 19.V.2010,

3815m, turn rock, V. Grebennikov”; 2 exx #2767-2768

(GNG) “RR. GHINA, Yunnan, Gang Shan at Dali,

N25°39’51” E100°06’05”, 04.vii.2011, 3815m, under

stone, V. Grebennikov”; 12 exx #4623^628, #5403-5406,

#6207-6208 (GNG) “GHINA, Yunnan, Haba Shan,

N27°20’51” E100°05’33”, 27.vi.2012, 4158m, under rock,

V. Grebennikov”.

Distribution. This species is known from the Gang Shan

Mountain range and nearby Mount Haba, both in Yunnan

(Fig. 1). Elevation: 3815^158 m.

Trichalophus scylla sp. n.

(Figs 2, 3)

Diagnosis. Specimens of this species are unique among

known congeners in Southwest Ghina by two characters:

they are the smallest and possess asymmetrical aedeagus

in dorsal view (character 7/1, Fig. 3).

Description. Holotype, male (Fig. 3). GenBank accession:

KM538655. Eength: 9.4 mm. Gombination of other mor-

phological characters as in Table 1 .

Intraspecific variation. GenBank accessions:

KM538656, KM538679. Eength: 9.0-9.4 mm.

Material examined. Holotype (Fig. 3) male (IZGAS):

#4436: “GHINA, Sichuan, 23km E Songpan, N32°38’07”

E103°49’10”, 24.V.2012, 3704m, under rock, V. Greben-

nikov”. Paratypes (GNG): 2 males #4418 and #4419 “GHI-

NA, Sichuan, 23km E Songpan, N32°37’38”

E103°50’03”, 26.V.2012, 3791m, sifting 09, V. Greben-

nikov”.

Distribution. This species is known only from the type

locality some 20 km E of Songpan, Sichuan (Fig. 1),

where it is found sympatrically with T. tibetanus. Eleva-

tion: 3704-3791 m.

Etymology. The species epithet is the Eatinized Greek

mythical name of Scylla, one of the Nereids, transformed

by Girce into a six-headed monster and who, together with

its counterpart Gharybdis, threatened Odysseus’ crew on

their return voyage from Troy to Ithaca; noun in apposi-

tion.

Trichalophus tibetanus (Suvorov, 1915)

(Figs 2, 3, 5)

tibetanus Suvorov 1915: 338 {Pseudalophus)

Type locality. Ghina, basin of the Blue river (=the

Yangtze), the Kundur-Tschu river, 13200’.

Type specimens. Syntype (ZIN, currently on loan in

MTD, Fig. 5), male, dissected by Rudiger Krause, labels

as in Fig. 5. Described from unlmown number of syntypes.

Twenty other similar specimens collected together with the

imaged syntype are also likely part of the type series; of

them four specimens each have a golden circle as in Fig.

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Diagnosis. This species is best distinguished by the pres-

ence of elongate and curved apical labella of aedeagus

(character 6/1, Fig. 3) and relatively thick apodeme of

male sternite 9 (character 9/0, Fig. 3).

Intraspecific variation. GenBank accessions:

KM538657, KM538658, KM538659, KM538661,

KM538663-64, KM538672-75, KM538680,

KM538685-86. Length: 11.5-13.3 mm (Songpan) and

10.3-12.1 mm (Mount Gongga). Each elytron with sin-

gle indistinct apical spot (Fig. 5), two indistinct spots, or

two distinct spots (Fig. 3). Specimens from Mount Gong-

ga have a long white longitudinal stripe laterally on each

elytron (Fig. 3).

Additional material examined. 19 exx in total: 4 exx

#2742-2745 (CNC) “RR. CHINA, Sichuan, NE slope

Gongga Shan, N25°53’53” E102°0r49”, 8.vi.2011,

4085m, under stone, V. Grebennikov”; 15 exx

#4437-4441, #5385-5388 and six not numbered speci-

mens in ethanol (CNC) “CHINA, Sichuan, 23km E Song-

pan, N32°38’07” E103°49’10”, 24.V.2012, 3704m, under

rock, V. Grebennikov”.

Distribution. Besides the type locality in the extreme

north-western Sichuan, this species is also known from

Mount Gongga and from vicinities of Songpan, both in

Sichuan (Fig. 1); in the latter locality this species is found

sympatrically with T. scylla sp. n. Elevation: 3704^085

Generic overview of the tribe Alophini

The proposed amalgamation of Alophini (as delimited be-

low) with the tribe Tropiphorini {sensu Alonso-Zarazaga

& Eyal 1999) by Zherikhin & Egorov (1991), for which

a synonymous name Eeptopiini was used by Marvaldi et

al. (2014), is not followed here. Neither taxonomic

arrangement was phylogenetically tested, therefore none

is better than the other. Additionally, dissolving the com-

pact and predominantly Holarctic Alophini in the much

larger cosmopolitan Tropiphorini would discourage any

practical effort to shed light on the genera involved, as at-

tempted below.

The tribe Alophini was first proposed by EeConte (1876:

115) to incorporate the Palaearctic species grouped then

in Alophus Schoenherr and species belonging to sixNearc-

tic genera, five of them newly established: Triglyphus

EeConte, Plinthodes EeConte, Acmaegenius EeConte,

Trichalophus EeConte, Lophalophus EeConte and Lepi-

dophorus Kirby, 1837. By the end of the millennium the

tribe consisted of 15 valid genera, including three de-

scribed from the Oligocene of the USA (Centron Scud-

der, 1893, Ger alophus Scudder, 1893 and Limalophus

Schudder, 1893, see Alonso-Zarazaga & Eyal 1999). Since

then Bright & Bouchard (2008) synonymised the genus

Acmaegenius under Trichalophus and reviewed the

Alophini of Canada and Alaska. Alonso-Zarazaga et al.

(2010) demonstrated that the sole known specimen of the

monotypic genus Ctenolobus Desbrochers des Eoges,

1 892 from Morocco is conspecific with the type species

of the otherwise strictly South American genus Stran-

galiodes Schoenherr, 1842 (Tropiphorini). As a result,

Strangaliodes was transferred to the otherwise strictly

Palaearctic Alophini and was keyed out against three oth-

er Mediterranean genera: Graptus, Rhytideres and Seidl-

itzia (Alonso-Zarazaga et al. 2010). Finally, Yunakov

(2013) synonymized Pseudalophus under Trichalophus.

At present the following 10 extant genera constitute the

tribe Alophini:

Graptus Schoenherr, 1823 (Figs 6A, B) with 37

species-group taxa is distributed in Western Palaearctic

(Yunakov 2013). Davidian & Arzanov (2004) revised and

keyed 10 Graptus species from Russia and adjacent lands,

including two newly described ones, and mentioned that

many poorly known nominal species have been reported

from the West Palaearctic.

Lepidophorus Kirby, 1837 (Figs 6C, D) consists of 11

brachypterous species found in western North America

(Anderson 1997, 2012; Bright & Bouchard 2008), two of

which, Z. inquinatusMmnQxhQim, 1852 andZ. lineaticol-

lis Kirby, 1837, are also found on the Asian side of the

Bering Strait (Yunakov 2013). Another extant North

American species, Z. thulius Kissinger, 1974, is known

from numerous subfossil records on both side of the

Bering Strait and, therefore, extant populations might per-

haps be discovered in the northern Pacific Asia (as Vitat-

itiis Kissinger, 1974 in Egorov et al. 1996 and in Ander-

son 1997). Anderson (2002) mentions that the genus can-

not be reliably distinguished from Dirotognathus Horn,

1876 (Tropiphorini Alonso-Zarazaga & Eyal 1999),

giving support to a notion to synonymize both tribes

(Zherikhin & Egorov 1991; Marvaldi et al. 2014).

Plinthodes LeConte, 1876 (Figs 6E, F) consists of two

North American species, P.foveirostris Chittenden, 1925

from Ohio, North Carolina, Tennessee and Virginia and

P taeniatus EeConte, 1857 from British Columbia, Wash-

ington and Oregon (Anderson 2002). Bright & Bouchard

(2008) questioned the distinctness of this genus from

Trichalophus.

Pseudobarynotus Desbrochers des Loges, 1891 (Fig.

8) contains a single mysterious species P. laticeps (Des-

brochers des Eoges, 1875) known only from the type se-

ries and described from “Pyrenes”. The type series could

have been mislabelled, while Kazakhstan was suggested

as its true origin (Alonso-Zarazaga et Eyal 1999). The lat-

er possibility is not unlikely, since the depicted syntype

(Fig. 8) resembles a species of Trichalophus and might

perhaps be later demonstrated as such.

Bonn zoological Bulletin 64 (2): 59-76

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Vasily V. Grebennikov

Fig. 6. Type species of the Alophini genera. A-B; Graptus triguttatus (Fabricius, 1775), Austria, Wien, date and collector un-

known, CNC; C-D: Lepidophorus lineaticollis Kirby, 1 837, USA, Alaska, Wasilla, 1 .viii. 1988, J.Pilny, CNC; E-F ; Plinthodes tae-

niatus LeConte, 1857, Canada, British Columbia, Victoria, 6.vii.l962, B.Carr, CNC; G-H: Rhytideres plicatus (Oliver, 1790), no

collecting data, CNC. Scale; 2 mm.

Rhytideres Schoenherr, 1823 (Figs 6G, H) includes

three species widely distributed around the Mediterranean

Sea (Yunakov 2013).

Seidlitzia Desbrochers des Loges, 1891 (Figs 7A, B)

consists of two species and one non-nominal subspecies

from Spain and Morocco (Yunakov 2013).

Strangaliodes Schoenherr, 1842 (habitus image in

Alonso-Zarazaga et al. 2010, figs lA, B) includes nine

species from the South American Cordillera, all of them

found in Chile and a few in neighbouring countries (Wib-

mer & O’Brien 1986); one of them also questionably

recorded from Morocco (Alonso-Zarazaga et al. 2010).

This is the only non-Palaearctic member of Alophini.

Trichalophus LeConte, 1876 (Figs 7C, D) consists of

51 species and one non-nominal subspecies distributed on

both sides of the Bering Strait; for more details see the

current paper.

Triglyphulus Cockerell, 1906 (Figs 7E, F) consists of

two species, T. ater LeConte, 1 876 and T. nevadensis Van

Dyke, 1938 from the western part of the USA (Anderson

2002 ).

Xeralophus Korotyaev, 1992 (Figs 7G, H) was estab-

lished to accommodate Alophus cretaceus Reitter, 1 894,

described from present day Ulan Bator, Mongolia. Besides

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Neglected Trichalophus: DNA barcode and phylogeography of high-altitude flightless weevils in Southwest China 73

Fig. 7. Type species of the Alophini genera. A-B: Seidlitzia maroccana (Fairmaire, 1 868), Morocco, date and collector unknown,

MNFIN, image: Antoine Mantilleri, © MNHN, original image showing right lateral view was digitally flipped horizontally to ap-

pear as left; C-D; Trichalophus didymus (LeConte, 1854), Canada, British Columbia, Kitsumkalum Lake, 16. vi. 1960, B.S.Hem-

ing, CNC; E-F; Triglyphulus ater (LeConte, 1 876), USA, Idaho, Bear, 26.vii. 1977, B.Carr, CNC; G-H: Xeralophus cretaceus (Re-

itter, 1894), Russia, Tyva, Kyzyl, 6.V.1974, B.A.Korotyaev, ZIN, image: Audrey Frolov. Scale: 2 mm.

the type series, four more specimens were later reported,

all found dead in sandy steppe of the neighbouring Tyva

Republic of Russia (Korotyaev 1992). Korotyaev (1992)

hypothesised that this xerophilic taxon is phylogenetical-

ly nested within the predominantly mesophilic '"Alophus”

(= Graptus). This hypothesis, if corroborated, would ren-

der the name Xeralophus a junior subjective synonym of

Graptus.

CONCLUDING REMARKS

It seems worthy of reiterating some important and perhaps

not too obvious generalities emerging from this study.

First, if the genus Trichalophus does exist in the phylo-

genetic sense, then its presence has been reconfirmed in

Southwest China for the first time since Suvorov (1915).

Second, the last glacial retreat is likely responsible for the

Bonn zoological Bulletin 64 (2): 59-76

©ZFMK

Vasily V. Grebennikov

Fig. 8. Pseudobarynotus laticeps (Desbrochers, 1874), syntype, MNHN, image: Antoine Mantilleri, © MNHN.

present day high altitude presence of Trichalophus in the

high mountains of Yunnan and Sichuan, but not for the

diversification of the lineages leading to the extant pop-

ulations. Third, in spite of the large body size and rela-

tive ease of sampling, Trichalophus is among the least un-

derstood genera of the Holarctic weevils. Such neglect is

partly due to the abundance of ambiguous historical

species names, particularly in Central Asia, which creates

a nomenclatorial impediment and hinders further research.

Fourth, relationships of Trichalophus in Alophini, and the

overall phylogenetic validity of this tribe and most of its

genera (particularly Plinthodes, Pseudobarynotus mdXer-

alophus) remain untested. When adequately studied, how-

ever, the genus Trichalophus is expected to be of signif-

icant biogeographic potential, similarly to other clades of

low-dispersing organisms most suitable to reveal the ge-

ographical component on their evolutionary past (Muri-

enne et al. 2014; Tanzler et al. 2014; Toussaint et al. 2015).

Acknowledgements. Curators of the collections mentioned

above variously helped in accessing specimens under their care.

Antoine Mantilleri (Paris, France) (1.) took syntype habitus and

label images of both Fairmaire’s names (Fig. 5); (2.) critically

verified the match between both original descriptions and the im-

aged syntypes (Fig. 5) to help establish their name-bearing sta-

tus and (3.) took images of Barynotus laticeps syntype (present-

ly Pseudobarynotus laticeps) and of Seidlitzia marrocana syn-

types (Figs 8 and 7A, B, respectively). Genrikh Ed. Davidian

and Boris A. Korotyaev searched on my request for the ZIN spec-

imen ofXeralophus cretaceus, which was imaged (Figs 7G, H.)

by Andrey V. Frolov (all St. Petersburg, Russia). Thierry Deuve

Bonn zoological Bulletin 64 (2): 59-76

(Paris, France) advised on collecting sites of Father Delavay in

Yunnan. David J. Clarke (Chicago, USA) collected and made

available the sequenced specimen of Trichalophus alternatus

(#2968). Ignacio Ribera (Barcelona, Spain) advised on imple-

mentation of DNA analytical techniques. Christian Schmidt, Ed-

uard Jendek and Bruce D. Gill (all Ottawa, Canada) reviewed

early versions of this paper.

REFERENCES

Alonso-Zarazaga M, Eyal CMC (1999) A world catalogue of fam-

ilies and genera of Curculionoidea (Insecta: Coleoptera) (ex-

cepting Scolytidae and Platypodidae). Entomopraxis,

Barcelona

Alonso-Zarazaga MA, Perrin H, Sanchez-Ruiz M (2010) Sur f i-

dentite du genre Ctenolobus Desbrochers des Eoges, 1892

(Coleoptera, Curculionidae). Bulletin de la Societe Ento-

mologique de France 115; 479-483

Anderson RS (1997) Weevils (Coleoptera; Curculionoidea, ex-

cluding Scolytinae and Platypodinae) of the Yukon. Pp.

523-562 in: banks HV & Downes JA (eds.) Insects of the

Yukon. Biological Survey of Canada (terrestrial arthropods),

Ottawa

Anderson RS (2002) Chapter 131. Curculionidae Eatreille 1802.

Pp. 722-815 in: Arnett RH Jr, Thomas MC & Skelley P (eds.)

American Beetles, Volume 2. CRC Press, Boca Raton

Anderson RS (2010) A taxonomic monograph of the Middle

American leaf-litter inhabiting weevil genus Theognete

Champion (Coleoptera: Curculionidae; Molytinae; Eymanti-

ni). Zootaxa 2458: 1-127

Anonymous (2014) Pierre Jean Marie Delavay. Online at;

http://fr.wikipedia.org/wiki/Pierre_Jean_Marie_Delavay last

accessed on September 16, 2014

©ZFMK

Neglected Trichalophus: DNA barcode and phylogeography of high-altitude flightless weevils in Southwest China 75

Astrin JJ, Stilben PE (2008) Phylogeny in cryptic weevils; mol-

ecules, morphology and new genera of western Palaearctic

Cryptorhynchinae (Coleoptera; Curculionidae). Invertebrate

Systematics 22: 503-522

Bajtenov MS (1974) Zhuki-dolgonosiki (Coleoptera: Attelabidae,

Cucrulionidae) Srednej Azii i Kazakhstana Illjustrirovannyj

opredeliteP rodov i katalog vidov. Izdatel’stvo Nauka Kaza-

khskoy SSR, Alma-Ata

Berman D, Alfimov A, Kuzmina S (2011) Invertebrates of the

relict steppe ecosystems of Beringia, and the reconstruction

of Pleistocene landscapes. Quaternary Science Reviews 30:

2200-2219

Bousquet Y, Bouchard P, Davies AE, Sikes DS (2013) Check-

list of beetles (Coleoptera) of Canada and Alaska. Second edi-

tion. Pensoft Publishers, Sofia

Bright DE, Bouchard P (2008) Coleoptera, Curculionidae, En-

timinae. Weevils of Canada and Alaska; Volume 2. The insects

and arachnids of Canada; part 25. NRC Research Press, Ot-

tawa

Bromham E (2008) Reading the Story in DNA: a Beginner’s

Guide to Molecular Evolution. Oxford University Press, Ox-

ford

Cohen KM, Finney S, Gibbard PE (2013) International Chronos-

tratigraphic Chart, International Commission on Stratigraphy.

Online at http://www.stratigraphy.org/ICSchart/Chronos-

tratChart2013-01.pdf last accessed on March 10, 2014

Darwin C (1859) On the origin of species by means of natural

selection, or the preservation of favoured races in the strug-

gle for existence. John Murray, Eondon

Davidian GE, Arzanov YuG (2004) A review of the weevil genus

Graptus Schoenherr (Coleoptera, Curculionidae) of the fau-

na of Russia and adjacent countries. Entomological Review

84; 998-1118

Desbrochers des Eoges J (1875) [new taxa]. Annales de la So-

ciete Entomologique de France (5)4 [1874]: ccxxvi-ccxxvii

Drummond AJ, Suchard MA, Xie D, Rambaut A (2012)

Bayesian phylogenetics with BEAUti and the BEAST 1.7.

Molecular Biology and Evolution 29: 1969-1973

Egorov AB, Zherikhin VV, Korotyaev BA (1996) 112b. Cur-

culionidae - Dolgonosiki. Pp. 341-522 in: EerPA(ed.) Opre-

delitel’ Nasekomykh Dal’nego Vostoka Rossii. Tom 3. Zh-

estkokrylye, ili Zhuki. Chasf 3. Dal’nauka, Vladivostiok (in

Russian)

Fairmaire E (1886) Descriptions de coleopteres de I’interieur de

la Chine (2e partie). Annales de la Societe Entomologique de

France 6; 303-356

Fairmaire E (1887) Coleopteres de Tinterieur de la China. An-

nales de la Societe Entomologique de Belgique 31; 87-136

Funk DJ, Omland KE (2003) Species-level paraphyly and poly-

phyly; frequency, causes, and consequences, with insight from

animal mitochondrial DNA. Annual Review of Ecology, Evo-

lution and Systematics 34: 397-423

Grant PR, Grant BR (2006) Evolution of character displacement

in Darwin’s finches. Science 313; 224-226

Grebennikov VV (2010) First Alaocybites weevil (Insecta:

Coleoptera; Curculionoidea) from the Eastern Palaearctic; a

new microphthalmic species and generic relationships.

Arthropod Systematics and Phylogeny 68: 331-365

Grebennikov VV (2014a) DNA barcode and phylogeography of

six new high altitude wingless Niphadomimus (Coleoptera:

Curculionidae; Molytinae) from Southwest China. Zootaxa

3838; 151-173

Grebennikov VV (2014b) Morimotodes, a new genus for two

minute wingless litter species from southwest China and Tai-

wan with an illustrated overview of Molytina and Plinthina

Bonn zoological Bulletin 64 (2): 59-76

genera (Coleoptera; Curculionidae: Molytini). Bonn Zoolog-

ical Bulletin 63: 123-147

Handel-Mazzetti H (2014) A botanical Pioneer in South West

China. Experience and Impressions of an Austrian Botanist

during the First World War. Online at

http;//www.paeo.de/hl/hand_maz/pioneer/buch.html last ac-

cessed September 16, 2014

Hey J (2001) Genes, categories, and species: the evolution and

cognitive causes of the species problem. Oxford University

Press, New York

Ikeda H, Nishikawa M, Sota T (2012) Eoss of flight promotes

beetle diversification. Nature Communications 3, 648; 1-7

Isaac NJB, Mallet J, Mace GM (2004) Taxonomic inflation: its

influence on macroecology and conservation. Trends in Ecol-

ogy and Evolution 19: 464-469

Korotyaev BA (1992) New and little known Palaearctic weevils

(Coleoptera; Apionidae, Curculionidae). Entomologicheskoe

Obozrenie 70(1991): 875-902, 951 (in Russian)

Krivets SA (2006) The biology of a weevil, Trichalophus leu-

con (Gebl.) (Coleoptera: Curculionidae), in Western Siberia.

Proceedings of the Russian Entomological Society 77:

189-191

Krivets SA, Burlak VA (1986) To the knowledge of morpholo-

gy of the weevil larvae (Coleoptera, Curculionidae). 1. Ento-

mologicheskoe Obozrenie 65 ; 592-603 (in Russian)

Eancaster R (1993) Plantsman’s paradise: travel in China. Gar-

den Art Press/Antique Collectors’ Club, Woodbridge

Eanfear R, Welch JJ, Bromham E (2010) Watching the clock:

Studying variation in rates of molecular evolution between

species. Trends in Ecology and Evolution 25; 495-503

EeConte JE (1876) Pp. 1-12, 112-445 in: EeConte JE & Horn

GH (eds.): The Rhynchophora of America North of Mexico.

Proceedings of the American Philosophical Society held at

Philadelphia for promoting useful knowledge 15, i+xvi, 1^55

Eyal CH (2014) 3.7.7 Molytinae Schoenherr, 1823. Pp. 424-451

in: Eeschen RAB & Beutel RG (eds.) Handbook of Zoology,

Arthropoda: Insecta: Coleoptera. Volume 3: Morphology and

Systematics (Phytophaga). Walter de Gruyter, Berlin

Machado A (2007) Rhyncogonus and Eaparocerus (Coleoptera,

Curculionidae, Entiminae), a parallel case of success in island

evolution. Report of a study trip to Moorea, Tahiti and Ruru-

tu. Vieraea 35; 61-76

Machado A, Eopez M, Almeida T, Hernandez M (2008) Mito-

chondrial DNA phylogenetic analysis of the genus Eaparo-

cerus (Coleoptera, Curculionidae, Entiminae). I. The Madeiran

clade. Zoologica Scripta 37: 415-427

Maddison DR (2014) An unexpected clade of South American

ground beetles (Coleoptera, Carabidae, Bembidion). ZooKeys

416: 113-155

Maloy OC (2001) White pine blister rust. Plant Health Progress

2001. Online at doi:10.1094/PHP-2001-0924-01-HM last ac-

cessed September 16, 2014

Marvaldi AE, Eanteri AA, Guadalupe del Rio M, Oberprieler RG

(2014) 3.7.5 Entiminae Schoenherr, 1823. Pp. 503-522 in:

Eeschen RAB & Beutel RG (eds.) Handbook of Zoology,

Arthropoda; Insecta: Coleoptera. Volume 3: Morphology and

Systematics (Phytophaga). Walter de Gruyter, Berlin

McKay BD, Zink RM (2014) Sisyphean evolution in Darwin’s

finches. Biological Review 90; 689-698

Mitterboeck TF, Adamowicz SJ (2013) Flight loss linked to faster

molecular evolution in insects. Proceeding of the Royal So-

ciety, Series B Biological Sciences 280; 20131128

Murienne J, Daniels SR, Buckley TR, Mayer G, Giribet G (2014)

A living fossil tale of Pangaean biogeography. Proceedings of

©ZFMK

Vasily V. Grebennikov

the Royal Society, Series B Biological Sciences 281:

20132648

Papadopoulou A, Anastasiou I, Vogler AP (2010) Revisiting the

insect mitochondrial molecular clock; the mid-aegean trench

calibration. Molecular Biology and Evolution 27 : 1659-1 672

Pierce RA (1960) Russian Central Asia 1867-1917. A Study in

Colonial Rule. University of California Press, Berkeley

Rambaut A (2014) FigTree, Version 1.4. online at

http://tree.bio.ed.ac.uk/software/figtree last accessed March

15, 2014

Rambaut A, Suchard M, Drummond A. (2014) Tracer, Version

1.6. online at http://tree.bio.ed.ac.uk/software/tracer/ last ac-

cessed March 15, 2014

Reitter E (1913) Analytische Revision der russisch-asiatischen

Alophus = Trichalophus-Arten (Col. Curcul.). Deutsche En-

tomologische Zeitschrift 1913: 446-457

Riedel A (2011) The weevil genus Trigonopterus Fauvel

(Coleoptera, Curculionidae) and its synonyms a taxonomic

study on the species tied to its genus-group names. Zootaxa

2977: 1-49

Riedel A, Sagata K, Suhardjono YR, Tanzler R, Balke M (2013a)

Integrative taxonomy on the fast track-towards more sustain-

ability in biodiversity research. Frontiers in Zoology 10: 15

Riedel A, Sagata K, Surbakti S, Tanzler R, Balke M (2013b) One

hundred and one new species of Trigonopterus weevils from

New Guinea. Zookeys 280: 1-150

Qiu Y-X, Fu C-X, Comes HP (20 11) Plant molecular phylogeog-

raphy in China and adjacent regions: Tracing the genetic im-

prints of Quaternary climate and environmental change in the

world’s most diverse temperate flora. Molecular Phylogenet-

ics and Evolution 59: 225-244

Shorthouse DP (2010) SimpleMappr, an online tool to produce

publication-quality point maps. Online at http://www.sim-

plemappr.net last accessed on March 15, 2014

Sites TW, Marshall JC (2004) Operational criteria for delimit-

ing species. Annual Review of Ecology and Systematics 35;

199-227

Suvorov GE (1915) Novye rody i vidy zhestkokrylikh

(Coleoptera, Curculionidae i Cerambycidae) palaearktich-

eskoy oblasti [New genera and species of beetles (Coleoptera,

Curculionidae and Cerambycidae) from the Palaearctic re-

gion]. Russkoe Entomologicheskoe Obozrenie 15: 327-346

(in Russian)

Tamura K, Stecher G, Peterson D, Filipski A, Kumar S (2013)

MEGA6: Molecular Evolutionary Genetics Analysis Version

6.0. Molecular Biology and Evolution 30: 2725-2729

Tanzler R, Toussaint EFA, Suhardjono YR, Balke M, Riedel A

(2014) Multiple transgressions of Wallace’s Fine explain di-

versity of flightless Trigonopterus weevils on Bali. Proceed-

ings of the Royal Society, Series B Biological Sciences 281;

20132528

Toussaint EFA, Tanzler R, Rahmadi C, Balke M, Riedel A.

(2015) Biogeography of Australasian flightless weevils (Cur-

culionidae, Celeuthetini) suggests permeability of Eydekker’s

and Wallace’s Fines. Zoologica Scripta 44: 632-644

Vences M, Guayasamin JM, Miralles A, De Ea Riva I (2013)

To name or not to name: Criteria to promote economy of

change in Einnaean classification schemes. Zootaxa 3636:

201-244

Vogler AP, Timmermans MJTN (2012) Speciation: don’t fly and

diversify? Current Biology 22: R284-R286

Wanat M (2007) Alignment and homology of male terminalia

in Curculionoidea and other Coleoptera. Invertebrate System-

atics 21: 147-171

Ward PS (2011) Integrating molecular phylogenetic results in-

to ant taxonomy (Hymenoptera: Formicidae). Myrmecologi-

calNews 15: 21-29

Wibmer GJ, O’Brien CW (1986) Annotated checklist of the wee-

vils (Curculionidae sensu lato) of South America (Coleoptera,

Curculionoidea). Memoirs of the American Entomological In-

stitute 39: xvi+563 pp

Yunakov N (2013) Entiminae: Alophini. Pp. 251-254 in: Eobl

I & Smetana A (eds.) Catalogue of Palaearctic Coleoptera, vol.

8. Curculionoidea IT Brill, Eeiden

Yunakov NN, Dedyukhin SV, Filimonov RV (2012) Towards the

survey of Entiminae weevils (Coleoptera: Curculionidae) of

Russia: species occurring in the Volga and Ural Regions. Russ-

ian Entomological Journal 21: 57-72

Bonn zoological Bulletin 64 (2): 59-76

©ZFMK

Bonn zoological Bulletin 64 (2): 77-106

March 2016

An annotated checklist of the inland fishes of Sulawesi

Friedrich Wilhelm Miesen**, Fabian Droppelmann’, Sebastian Hiillen*,

Renny Knrnia Hadiaty^ & Fabian Herder'

'Zoologisches Forschungsmuseum Alexander Koenig, Bonn, Germany

Uchthyology Laboratory, Division of Zoology, Research Center for Biology, Indonesian Institute of Science (LIPI),

Cibinong, Indonesia;

E-mail: fw.miesen@googlemail.com; +49 (0)228 9122 431

Abstract. Sulawesi is the largest island of the Wallacea. Here, we present an annotated checklist of fish species record-

ed in Sulawesi’s inland waters. We recognize a total of 226 species from 112 genera and 56 families. Gobiidae (41 species),

Adrianichthyidae (20 species) and Telmatherinidae (19 species) are most species-rich, making up a total of 43% of the

total species diversity. 65 species are endemic to Sulawesi’s freshwaters, including 1 9 Tematherinidae, 1 7 Adrianichthyi-

dae, and 17 Zenarchopteridae. 44% of the inland fish fauna are obligate freshwater fishes, followed by euryhaline (38%)

and amphi-, ana- or diadromous (29%) taxa. 65 species have been recorded from lacustrine environments. However, we

stress that the data available are not representative for the island’s freshwater habitats. The fish species diversity of the

spectacular lakes is largely explored, but the riverine ichthyofaunas are in clear need of further systematic exploration.

Keywords. Sulawesi, freshwater, fishes, endemism, Wallacea, SE-Asia

INTRODUCTION

Sulawesi is the largest island of the Wallacea, a biodiver-

sity hotspot located between the Sunda and Sahul shelves

(Mokodongan & Yamahira 2015, Myers et al. 2000, Whit-

ten et al. 2002). The onset of the scientific investigation

of Sulawesi's inland waters and its fishes dates back to

the late 19^^ century (Abendanon 1915a, b, Bleeker 1855a,

1858a, b, Boulenger 1897). Exploration of species diver-

sity was, and still is, in the focus of ichthyological research

on the island (e.g. Hadiaty 2007, Hadiaty & Wirjoatmod-

jo 2003, Hadiaty et al. 2004, Kottelat 1989a, b, c, 1990a,

b, c, d, 1991, Larson 2001, Parent! 2008, 2011, Weber

1909, 1913), complemented more recently by studies on

evolutionary biology (e.g. Herder & Schliewen 2010).

Sulawesi’s freshwater environments are home to sev-

eral endemic animal radiations that include gastropods,

crustaceans, and fishes (e.g. de Bruyn et al. 2013, Herder

et al. 2006a, Mokodongan & Yamahira 2015, Parent! 2011,

Parent! et al. 2013, Rintelen et al. 2007a, b, Rintelen et

al. 2012, Tweedley et al. 2013). Much of this diversity is

restricted to species flocks confined to a few ancient lakes,

systems that serve as models for the study of speciation

processes (Herder & Schliewen 2010, Rintelen et al. 2010,

2012, Vaillant et al. 2011). Phylogeographic studies of the

island’s freshwater animals accordingly focused mostly on

lineages of molluscs, shrimps, crabs, and fishes, with fo-

cus on the lake radiations (e.g. Mokodongan & Yamahi-

ra 2015, Rintelen et al. 2007b, 2014, but see also de Bruyn

et al. 2012, 2013; reviewed by Rintelen et al. 2012).

Received: 30.10.2015

Accepted: 09.12.2015

Parent! recognized 57 species of freshwater fishes as en-

demic to Sulawesi, with the majority being restricted to

the ancient lakes (Parent! 2011). Most of the non-endem-

ic fish species are classified as secondary or peripheral

freshwater fishes (Berra 2001), with occasional records of

marine species (Kottelat 1990a, Tweedley et al. 2013).

Sulawesi's lakes include some of the oldest lakes on

earth. Lake Poso and the Malili Lakes in the highlands of

Central Sulawesi are so-called ancient lakes, exception-

ally long-lived lakes that have existed for more than

100,000 years (Brooks 1950, Rintelen et al. 2012). The

Malili Lakes system is known for its endemic species flock

of sailfin silversides (Telmatherinidae) (Herder et al.

2006a), but also includes small radiations of ricefishes

(Oryzias) and gobies {Mugilogobius, Glossogobius;

Hoese et al. 2015, Kottelat 1990d, Larson et al. 2014).

Moreover, the lakes and their surroundings harbour en-

demic species of half beaks (Dermogetys,Nomorhamphus;

Huylebrouck et al. 2012, Meisner 2001). Ricefishes

{Oryzias, Adrianichthys) are the dominant fish radiation

of Lake Poso (Kottelat 1990b). Remote lakes Lindu and

Tiu are substantially smaller than the ancient lakes, and

harbour two (Lindu) or one (Tiu) endemic (rice-) fish

species (Mokodongan et al. 2014, Parent! 2008). Lake

Tondano on the northern tip of North Sulawesi is the on-

ly known habitat of Tondanichthys kottelati (Collette

1995), an endemic genus and species of viviparous half-

beak (Collette 1995). Lake Tempe and Lake Sidereng are

Corresponding editor: P Wagner

Friedrich Wilhelm Miesen et al.

LakeTondano

Lake Limboto

Lake

Lindu

Lake Matano

- Lake Mahalona

Lake

Towuti

.-Tenggara

Lake

Sidenreng

Lake Tern pe

150 km

Fig. 1. Map of Sulawesi showing the island's administrative partition as referred to in this study, major lakes are highlighted.

Bonn zoological Bulletin 64 (2): 77-106

©ZFMK

An annotated checklist of the inland fishes of Sulawesi

shallow lakes in southern Sulawesi, but little remains

known about their fish fauna, and its present status apart

from its use in aquaculture (Hadijah et al. 2014, Tamsil

2000) (for the location of Sulawesi's lakes see Fig. 1).

In contrast to the ancient lakes, Sulawesi's riverine fish

fauna has attracted far less scientific interest, and the num-

ber of studies is limited (de Bruyn et al. 2013, Mokodon-

gan & Yamahira 2015, Schwarzer et al. 2008, Tweedley

et al. 2013). A typical feature of Sulawesi's riverine en-

vironments is the absence of large, slow rivers and

drainages (Kottelat 1990a), in contrast to the hydrology

of most other larger Indonesian islands (Stelbrink et al.

2012). The majority of drainages are rather small and

short, typically with medium to high stream velocity (Kot-

telat 1990a).

Sulawesi's freshwaters have been subject to massive fish

species introduction and alien species invasion (Herder et

al. 2012a, Kottelat & Whitten 1996, Parenti 2011, Whit-

ten et al. 1987). Alien fishes were introduced for food pro-

duction (e.g. Oreochromis spp., Channa spp.) (Whitten et

al. 1987), pest control (Poecilia reticulata), or aquarium

trade (e.g. flowerhorn cichlids) (Herder et al. 2012a). The

spread of populations of alien fish species in freshwater

systems of the island is apparently rapid, and may pose

as a threat to the native communities (e.g. Herder et al.

2012a, Tweedley et al. 2013).

Purpose of this paper

The literature on freshwater fish species of Sulawesi is

complex and dispersed. The most recent comprehensive

source covering the island’s ichthyofauna dates back to

1993 (Kottelat et al. 1993), but is not focussed on the is-

land’s fishes, and meanwhile partially out-dated. Kotte-

lat’s recent catalogue on “The Fishes of the Inland Wa-

ters of Southeast Asia” (Kottelat 2013) includes Sulawe-

si, but focuses on the nomenclature of the whole South-

east Asian ichthyofauna, and the related bibliography. The

present paper aims at summarizing ichthyological records

from Sulawesi’s inland waters. It provides an account of

actual species records, and species that have not actually

been recorded, but are likely to be present according to

their known distribution. It is understood that the authors

do not claim that this list is complete with respect to all

records ever made, but aim at providing a baseline for

analysing species records, required for upcoming inves-

tigations of the island’s fauna.

MATERIAL AND METHODS

Literature records are compiled from the scientific liter-

ature; sources or records considered questionable were not

included. Material examined was mostly collected during

various field campaigns of the senior author's group in Su-

lawesi, since 2002. Field methods applied during field-

work include beach seining, dipnetting, gillnetting,

scubadiving and electrofishing. Samples were either fixed

in formalin (4%) prior to storage in ethanol (80%), or fixed

and stored directly in pure ethanol (-98%). Specimens

were determined to the lowest feasible taxonomical lev-

el using the most recent literature available. The system-

atic division largely follows Kottelat (2013). ZFMK: Fish

collection of Zoologisches Forschungsmuseum Alexander

Koenig Bonn, Germany. Coordinates with reference to

ZFMK specimens are own species records, linked to the

respective voucher. Records that are not linked to vouch-

ers, represent visual records (F.H.); ZMH: Zoologisches

Museum Hamburg in Hamburg, Germany; MZB: Muse-

um Zoologicum Bogoriense in Cibinong, Indonesia.

Species occurrence is classified to: Euryhaline: species

with a broad tolerance towards salinity and thus can be

found in marine, brackish and freshwater environments

(Hiroi & McCormick 2012); anadromous: species with

adults entering marine environments and reproduction in

freshwaters (Daverat et al. 2012); catadromous: species

that migrate into marine environments for reproduction

(Daverat et al. 2012); amphidromous: species that migrate

between marine and freshwater environments for purpos-

es other than reproduction (Daverat et al. 2012); freshwa-

ter: primary or secondary/obligate freshwater species with

no marine stadium or life history phase (Berra 2001); in-

troduced: non-native, introduced species.

Species expected to occur in Sulawesi, but lacking ac-

tual records from the island, are included as “potential”,

and justified. Clearly non-native species are classified as

“introduced”. Synonyms listed are restricted to the most

common ones, with emphasis on publications dealing with

Sulawesi’s ichthyofauna (see Kottelat 2013 for nomencla-

ture).

RESULTS

CHONDRICHTHYES

CARCHARHINIFORMES

Carcharhinidae

Requiem sharks: Marine; all oceans (Nelson 2006); enter

brackish and freshwaters, including lakes (Grace 2001,

Nelson 2006).

Carcharhinus leucas (Muller & Henle, 1839)

Carcharias leucas Muller & Henle, 1839

Potential: Euryhaline; worldwide in subtropical and trop-

ical coastal waters; enter brackish and freshwaters (Com-

pagno 1984, Heupel & Simpfendorfer 2008); no actual

records for Sulawesi.

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PRISTIFORMES

Pristidae

Sawfishes: Euryhaline; enter brackish and freshwaters;

distributed in all tropical and subtropical oceans (Nelson

2006, Wueringer et al. 2009).

Pristis pristis (Linnaeus, 1758)

Squalus pristis hmmQm, 1758

Potential: Euryhaline; worldwide in subtropical and trop-

ical coastal waters; enter brackish and freshwaters; no ac-

tual records for Sulawesi (Einnaeus 1758, McEachran &

Carvalho 2002).

MYLIOBATIFORMES

Dasyatidae

Stingrays: Marine; distributed throughout the Atlantic and

Indo-Pacific; some species enter brackish and freshwaters

(Compagno & Roberts 1982, Nelson 2006).

Himantura leoparda Manjaji-Matsumoto & Last, 2008

Potential: Euryhaline; enter brackish and freshwaters; no

actual records for Sulawesi; distributed throughout the In-

do-West-Pacific (Manjaji-Matsumoto & East 2008).

Himantura uarnak (Gmelin, 1789)

Raja uarnak GmQlm, 1789

Potential: Euryhaline; enter brackish and freshwaters; no

actual records for Sulawesi; distributed throughout the In-

do-Pacific (Compagno et al. 1989, Gmelin 1789).

Himantura undulata (Bleeker, 1852)

Trygon undulata Bleeker, 1 852d

Potential: Euryhaline; enter brackish and freshwaters; no

actual records for Sulawesi; distributed throughout the In-

do-West Pacific (East & Stevens 1994).

Himantura tutul Borsa, Durand, Shen, Alyza, Solihin

& Berrebi, 2013

Potential: Euryhaline; enter brackish and freshwaters; dis-

tributed throughout the Indo Pacific; no actual records

from Sulawesi (Borsa et al. 2013).

ACTINOPTERYGII

ELOPIFORMES

Megalopidae

Tarpons: Euryhaline; enter brackish and freshwaters; dis-

tributed in tropical and subtropical regions (Adams et al.

2013, Nelson 2006).

Megalops cyprinoides (Broussonet, 1782)

Clupea cyprinoides Broussonet, 1782

Euryhaline; enter brackish and freshwaters; record from

Badjoa, Sulawesi Selatan (Adams et al. 2013, Bleeker

1865a, Coates 1987).

ALBULIFORMES

Albulidae

Bonefishes: Euryhaline; enter brackish and freshwaters;

distributed throughout tropical regions (Adams et al. 2013,

Nelson 2006).

Albula glossodonta (Forskal, 1775)

Argentina glossodonta Forskal, 1775

Potential: Euryhaline; enter brackish waters; no actual

records for Sulawesi; distributed throughout the Indo-Pa-

cific (Myers 1991, Randall & Bauchot 1999).

ANGUILLIFORMES

Anguillidae

Freshwater eels: Catadromous; adults inhabit freshwaters

or estuaries; marine reproduction; juveniles enter fresh-

waters after metamorphosis; distributed throughout trop-

ical and subtropical regions except the South Atlantic and

Eastern Pacific (Arai et al. 1999, Nelson 2006).

Anguilla bicolor McClelland, 1844

Catadromous; distributed throughout the Indo-Pacific

(Arai et al. 1999, Kottelat 2013); record from Buton

(Tweedley et al. 2013).

Anguilla celebesensis Kaup, 1857

Anguilla ancestralis Ege, 1939

Catadromous; enter brackish and freshwaters; distributed

throughout the Indo-Pacific (Arai et al. 1999); recorded

from Eake Tondano, Manado, Sulawesi Utara (Ege

1939), Buton and Kabaena (Tweedley et al. 2013), Takes

of Gorontalo (Haryono & Tjakrawidjaja 2004), Sulawe-

si Utara (Arai et al. 2003), Sulawesi Tengah (Arai et al.

2003).

Anguilla interioris Whitley, 1938

Potential: Catadromous; enter brackish and freshwaters;

distributed throughout the Indo-Pacific; no actual records

for Sulawesi (Arai et al. 1999, Kottelat 2013).

Anguilla marmorata Quoy & Gaimard, 1824

Catadromous; enter brackish and freshwaters; distributed

throughout the Indo-Pacific (Arai et al. 1999); records

from Sulawesi Utara (Haryono et al. 2002), Sulawesi

Tenggara (02°56.035’S 121°06.855’E, ZFMK 066057),

Sulawesi Selatan (3°41.589’S 119°38.629’E, ZFMK

69560), Eake Poso, Sulawesi Tengah (visual record F.H.).

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An annotated checklist of the inland fishes of Sulawesi

Anguilla nebulosa McClelland, 1844

Catadromous; enter brackish and freshwaters; distributed

throughout the Indo-Pacific (Arai et al. 1999); record from

Sulawesi Barat (2°37.368S 1 19°08.784’E, ZFMK 69701).

Anguilla obscura Gunther, 1872

Potential: Catadromus; distributed throughout the Pacif-

ic; no actual records for Sulawesi (Arai et al. 1999, Giin-

ther 1872a).

Moringuidae

Worm, Spaghetti eels: Euryhaline; enter brackish and

freshwaters; fossorial lifestyle; distributed throughout the

tropical Western Atlantic and the Indo-Pacific (Nelson

2006, Tsukamoto et al. 2014).

Moringua guthriana (McClelland, 1844)

Ptyobranchus arundinaceus McClelland, 1 844

Potential: Euryhaline; enter brackish waters; distributed

throughout the Indo-Pacific (Kottelat 2013); no actual

records from Sulawesi.

Moringua javanica (Kaup, 1856)

Aphthalmichthys javanicus Kaup, 1856

Euryhaline; enter brackish waters; distributed throughout

the Indo-Pacific (Allen & Steene 1988); record from Ba-

ton (Tweedley et al. 2013).

Moringua microchir Bleeker, 1853

Potential: Euryhaline; enter brackish waters; distributed

throughout the Indo-West Pacific (Keith et al. 2006); no

actual records for Sulawesi.

Moringua raitaborua (Hamilton, 1822)

Moringua latebrosa Schultz, 1953

Euryhaline; enter brackish waters (Kottelat 2013); record

from: Kwandang, Gorontalo (Castle 1968, Kottelat 2013,

Smith 1994).

Muraenidae

Moray eels: Euryhaline; enter brackish and freshwaters;

worldwide distributed in all tropical regions (Nelson 2006,

Tsukamoto et al. 2014).

Gymnothorax polyuranodon (Bleeker, 1853)

Muraena polyuranodon Bleeker, 1853f

Potential: Euryhaline; enter brackish and freshwaters; dis-

tributed throughout the Indo-Pacific; no actual records

from Sulawesi (Ebner et al. 2011, Tsukamoto et al. 2014).

Gymnothorax tile (Hamilton, 1822)

Muraenophis tile Hamilton, 1 822

Euryhaline; enter brackish waters; distributed throughout

the Indo-Pacific (Tsukamoto et al. 2014); record from Ba-

ton (Tweedley et al. 2013).

Strophidon sathete (Hamilton, 1822)

Muraenophis sathete Hamilton, 1 822

Muraena macrurus Bleeker, 1 854b

Euryhaline; enter brackish waters; distributed throughout

the Indo-Pacific; record from Buton (Tweedley et al.

2013).

Ophichthidae

Snake, Worm eels: Marine; some species enter freshwa-

ters; cosmopolitan, distributed throughout tropical regions

(Cosker et al. 2012, Eschmeyer 2015, Froese & Pauly

2014, Nelson 2006).

Cirrhimuraena chinensis Kaup, 1856

Euryhaline; enter brackish waters; distributed throughout

the Indo-Pacific; record from Makassar, Sulawesi Selatan

(Kaup 1857).

Lamnostoma mindora (Jordan & Richardson, 1908)

Coecula mindora Jordan & Richardson, 1908

Euryhaline; enter brackish waters; distributed throughout

the Indo-Pacific; record from Buton (Tweedley et al.

2013).

Muraenichthys gymnopterus Bleeker, 1852

Muraena gymnopterus Bleeker, 1 852b

Muraenichthys microstomus Bleeker, 1 864

Euryhaline; enter brackish waters; distributed throughout

the West-Pacific; record from Makassar, Sulawesi Sela-

tan (Bleeker 1864).

Ophichthus polyophthalmus Bleeker, 1864

Potential: Euryhaline; enter brackish waters; distributed

throughout the Indo-Pacific (Kottelat 2013); no actual

records from Sulawesi.

Pisodonophis cancrivorus (Richardson, 1848)

Ophisurus cancrivorus Richardson, 1848

Ophisurus brachyosoma Bleeker, 1852b

Euryhaline; enter brackish waters; distributed throughout

the Indo-Pacific; record from Makassar, Sulawesi Selatan

(Bleeker 1852b).

Yirrkala kaupii (Bleeker, 1858)

Sphagebranchus kaupii Bleeker, 1858b

Catadromous; enter brackish and freshwaters; distribute

throughout Asia; record from Klabat Diatas, Sulawesi

Utara (Bleeker 1858b).

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GONORHYNCHIFORMES

Chanidae

Milkfishes: Euryhaline; enter brackish waters; distributed

throughout the Indo-Pacific (Berra 2001; Nelson 2006).

Chanos chanos (Forsskal, 1775)

Mugil chanos 111 5

Euryhaline; enter brackish and freshwaters (Allen et al.

2002); record from Sulawesi Selatan (4°07.456’S

119’37.196’E, ZFMK 69759).

CYPRINIFORMES

Cyprinidae

Minnows, Carps: Freshwater; native throughout Africa,

Eurasia and North America, introduced worldwide;

cyprinids are naturally absent from Sulawesi (Kottelat

1990a, Nelson 2006).

Barbonymus gonionotus (Bleeker, 1849)

Barbus gonionotus Bleeker, 1 849a

Introduced: Freshwater; record from Sulawesi Selatan

(3°41.589S 119°38.629’E, ZFMK 69514-69516, 69534,

69552-69555), Eake Poso, Sulawesi Tengah (Kottelat

1990b).

Carassius auratus (Linnaeus, 1758)

Introduced: Freshwater; common ornamental fish; breed-

ing form from East Asia; distributed almost worldwide;

record from the Malili Eakes system, Sulawesi Selatan

(Nasution & Aisyah 2013), Eake Poso, Sulawesi Tengah

(Kottelat 1990b).

Cyprinus carpio (Linnaeus, 1758)

Introduced: Freshwater; native to Central Asia; record

from Malili Lakes system, Sulawesi Selatan; Lake Poso,

Sulawesi Tengah (Kottelat 1990b); aquaculture

escapees or stocked specimens (Herder et al. 2012a).

Cyprinus cf. rubrofuscus (Lacepede, 1803)

Cyprinus rubro-fuscus Lacepede, 1 803

Introduced: Freshwater; ornamental carp varieties; possi-

bly derived from C. fuscus or hybrids (See Kottelat &

Freyhof 2007, and references therein); record from Poso

River, Sulawesi Tengah (visual record F.H.).

Osteochilus vittatus (Valenciennes, in Cuvier und Va-

lenciennes, 1842)

Rohita vittata Valenciennes, in Cuvier und Valenciennes,

1842

Osteochilus hasselti (Valenciennes, in Cuvier und Valen-

ciennes, 1842)

Introduced: Freshwater; native to mainland Southeast

Asia; records from Malili Lakes system and Lake Siden-

reng, Sulawesi Selatan (Omar 2010); Lake Poso, Sulawe-

si Tengah (visual record F.H.).

CHARACIFORMES

Characidae

Characins: Freshwater; native to Central America, South

America, and Africa; without native members in Asia (Es-

chmeyer 2015, Froese & Pauly 2014, Nelson 2006).

Colossoma macropomum (Cuvier, 1816)

Myletes macropomus Cuvier, 1816

Introduced: Freshwater; native South America (Santos et

al. 2007); record from Lake Matano, Sulawesi Selatan

(Herder et al. 2012a) and Lake Poso, Sulawesi Tengah (vi-

sual record F.H.).

SILURIFORMES

Plotosidae

Eeltail catfishes: Euryhaline; enter brackish and freshwa-

ters (Nelson 2006, Usman et al. 2013); distributed

throughout the Indo-West Pacific (Eschmeyer 2015,

Froese & Pauly 2014, Usman et al. 2013).

Plotosus canius Hamilton, 1822

Introduced: Amphidromous; enter brackish and freshwa-

ters (Usman et al. 2013); record from Sulawesi Selatan

(04°07.540S 119°37.295’E, ZFMK 066013).

Clariidae

Airbreathing catfishes: Freshwater; native throughout

Africa, Syria, South and West Asia (Nelson 2006, Ng &

Kottelat 2007); 16 species ocurring in Asia (Ng & Kotte-

lat 2007).

Clarias batrachus (Linnaeus, 1758)

Introduced: Freshwater; native to India, Indochina, Sun-

daland and the Philippines; neotype from Java, Indonesia

(Nelson 2006, Ng & Kottelat 2007); record from the Malili

Lake system, Sulawesi Selatan (Herder et al. 2012a); all

non-Java specimens are considered as related species-

complex (Ng & Kottelat 2007).

CLUPEIFORMES

Engraulididae

Anchovies: Euryhaline; enter brackish and freshwaters;

distributed throughout the Atlantic and Indo-Pacific

(Nelson 2006, Whitehead et al. 1988).

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An annotated checklist of the inland fishes of Sulawesi

Thrissina baelama (Forskal, 1775)

Clupea baelama Forskal, 1775

Thryssa baelama (Forskal, 1775)

Potential: Euryhaline; enter brackish waters (Whitehead

et al. 1988); no actual records for Sulawesi; distributed

throughout the Indo-Pacific (Kottelat 2013).

Thrissina encrasicholoides (Bleeker, 1852)

EngrauUs encrasicholoides Bleeker, 1851a

Thriyssa encrasicholoides Bleeker, 1852a

Potential: Euryhaline; enter brackish waters; no actual

records for Sulawesi; distributed throughout the Indo-Pa-

cific (Whitehead et al. 1988, Wongratana et al. 1999).

Thrissina mystax (Bloch, in Schneider, 1801)

Stolephorus hamiltoni Bleeker, 1 872

Thryssa mystax (Bloch in Schneider, 1801)

Euryhaline; enter brackish waters; distributed throughout

the Indo-West Pacific (Wongratana et al. 1999); record

from Badjoa, Sulawesi Selatan (Bleeker, 1872).

Clupeidae

Herrings, Shads, Sardines: Euryhaline; distributed world-

wide (Nelson 2006, Wongratana et al. 1999).

Herklotsichthys quadrimaculatus (Riippell, 1837)

Clupea quadrimaculata Riippell, 1837

Potential: Euryhaline; enter brackish waters; no actual

records for Sulawesi; distributed throughout the Indo-Pa-

cific (Wongratana et al 1999).

MUGILIFORMES

Mugilidae

Mullets: Euryhaline; enter brackish and freshwaters; dis-

tributed throughout all tropical and temperate regions (Cu-

vier & Valenciennes 1836, Durand 2012, Nelson 2006).

Cestraeus plicatilis Valenciennes, in Cnvier & Valenci-

ennes, 1836

Catadromous; enter brackish and freshwaters; distributed

throughout the Indo-West Pacific (Harrison & Senou

1999); record from Sulawesi freshwaters (Valenciennes,

in Cuvier and Valenciennes 1836).

Mugil cephalus Linnaens, 1758

Catadromous; marine, enter brackish and freshwaters

(Harrison & Senou 1997); record from Sulawesi Selatan

(4°07.456’S 119’37.196’E, ZFMK 69741, 69760-69763).

ATHERINIFORMES

Telmatherinidae

Sailfin silvers ides: With exception of Kalyptatherina

helodes, on islands off the Vogelkop peninsula, restricted

to Sulawesi; adaptive radiations in the Malili Eakes: Eakes

Matano, Mahalona, Towuti, Eontoa, and connecting / sur-

rounding rivers and streams (Hadiaty & Wirjoatmodjo

2002, Hadiaty et al. 2004, Herder et al. 2006a, b, Pfaen-

der et al. 2011; reviewed by Herder & Schliewen (2010)).

Telmatherina abendanoni Weber, 1913

Freshwater; endemic to Eake Matano, Sulawesi Selatan;

predatory, benthic “sharpfin” species of Telmatherina (Ha-

diaty & Wirjoatmodjo 2002, Herder et al. 2006a).

Telmatherina antoniae Kottelat, 1991

Freshwater; endemic to Eake Matano, Sulawesi Selatan;

a “roundfin” Telmatherina., the formal name Telmatheri-

na antoniae is currently applied to two distinct popula-

tions, T antoniae “small” and “large”; males in distinct

colour morphs, females with cryptic polymorphism (Ha-

diaty & Wirjoatmodjo 2002, Herder et al. 2006a, Herder

et al. 2008, Pfaender et al. 2014).

Telmatherina bond Weber & de Beaufort, 1922

Freshwater; endemic to rivers and streams of the Malili

Eakes system, and adjacent systems; enter Eakes Matano,

Mahalona and Towuti, Sulawesi Selatan; the only formal

name currently available for stream-dwelling Telmathe-

rina (Hadiaty & Wirjoatmodjo 2002, Herder et al. 2006a).

Telmatherina celebensis Boulenger, 1897

Freshwater; endemic to Eakes Mahalona and Towuti, Su-

lawesi Selatan; additional record from River Tominanga;

common in both lakes; males in distinct colour morphs

(Hadiaty et al. 2004, Herder et al. 2006a).

Telmatherina cf. celebensis “Lontoa”

Freshwater; endemic to Eake Eontoa [also Eantoa or Wa-

wontoa], and surrounding swamps, Sulawesi Selatan;

smaller and deeper bodied than T celebensis from Eakes

Mahalona and Towuti, Sulawesi Selatan; males in distinct

colour morphs (Herder et al. 2006a).

Telmatherina “elongated”

Freshwater; endemic to Eake Matano, Sulawesi Selatan;

predatory, benthic “sharpfin” Telmatherina with short fins

and slender body (Herder et al. 2006a).

Telmatherina obscura Kottelat, 1991

Freshwater; endemic to Eake Matano, Sulawesi Selatan;

small, blackish inshore “sharpfin” Telmatherina (Hadiaty

& Wirjoatmodjo 2002, Herder et al. 2006a).

Telmatherina opudi Kottelat, 1991

Freshwater; endemic to Eake Matano, Sulawesi Selatan;

small “sharpfin” Telmatherina predominantly inhabiting

well- structured shallows; males in distinct colour morphs

(Hadiaty & Wirjoatmodjo 2002, Herder et al. 2006a).

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Telmatherina prognatha Kottelat, 1991

Freshwater; endemic to Lake Matano Sulawesi Selatan;

large, slender “roundfin” Telmatherina, conspicuous

“beak-like” jaws; epibenthic, predatory ecology; males in

distinct colour morphs (Hadiaty & Wirjoatmodjo 2002,

Herder et al. 2006a).

Telmatherina sarasinorum Kottelat, 1991

Freshwater; endemic to Lake Matano, Sulawesi Selatan;

benthic “sharpfm” Telmatherina specialized on eating eggs

of sailfin silversides including conspecifics; males in dis-

tinct colour morphs (Hadiaty & Wirjoatmodjo 2002, Cer-

wenka et al. 2012, Herder et al. 2006a, Gray et al. 2007,

Gray et al. 2008a, Gray et al. 2008b, Pfaender et al. 2010).

Telmatherina “thicklip”

Freshwater; endemic to Lake Matano, Sulawesi Selatan;

benthic “sharpfin” Telmatherina specialized on eating

shrimps; deep-bodied species, with pronounced “puffy

lips” (Herder et al. 2006a, Pfaender et al. 2010).

Telmatherina wahjui Kottelat, 1991

Freshwater; endemic to Lake Matano, Sulawesi Selatan;

“sharpfm” Telmatherina occurring at the outlet of Lake

Matano to River Petea, and the shallows of the lake (Ha-

diaty & Wirjoatmodjo 2002, Herder et al. 2006a).

Paratherina cyanea Aurich, 1935

Freshwater; endemic to Lake Towuti and Lake Mahalona,

Sulawesi Selatan; slender, conspicuously large-eyed (Ha-

diaty et al. 2004, Herder et al. 2006a).

Paratherina labiosa Aurich, 1935

Freshwater; possibly endemic to Lake Lontoa, Sulawesi

Selatan; holotype destroyed (Kottelat 1990c); Kottelat

(1990c) tentatively assigned four juveniles obtained in

1989 to P. labiosa, further investigations lacking; no

records during recent surveys by F.H. in Lake Lontoa.

Paratherina striata Aurich, 1935

Freshwater; endemic to Lakes Towuti and Mahalona, Su-

lawesi Selatan; largest sailfin silverside species; males in

distinct colour morphs (Hadiaty et al. 2004, Herder et al.

2006a, Kottelat 1990c).

Paratherina wolterecki Aurich, 1935

Freshwater; endemic to Lake Mahalona, Sulawesi Sela-

tan; blackish male colouration (Hadiaty et al. 2004, Herder

et al. 2006a).

Tominanga aurea Kottelat, 1990

Freshwater: endemic to Lake Mahalona, Sulawesi Sela-

tan; enter rivers; Kottelat (1990c) distinguished Tominan-

ga aura from Tominanga sanguicauda by male colour

traits, gill raker counts, and occurrence (Lake Mahalona

vs. Lake Towuti); Herder et al. (2006a) reported less clear

indications for species discrimination based on colour

traits, and highlight the need for more detailed examina-

tions (Kottelat 1990c, Hadiaty et al. 2004, Herder et al.

2006a).

Tominanga sanguicauda Kottelat, 1990

Freshwater; endemic to Lake Towuti, Sulawesi Selatan;

enter rivers; see Tominanga aurea for notes on species dis-

crimination (Hadiaty et al. 2004, Kottelat 1990c).

Marosatherina ladigesi (Ahl, 1936)

Telmatherina ladigesi Ahl, 1936

Freshwater; endemic to the Bantimurung area, Maros

karst, Sulawesi Selatan; dwelling in cool karst streams;

males with conspicuously elongated blackish rays in sec-

ond dorsal and anal fins; popular aquarium species (Ha-

diaty 2007); locally transferred for breeding purposes

(F.H., pers. obs.).

Phallostethidae

Priapium fishes: Euryhaline; enter brackish and freshwa-

ters; distributed in Southeast Asia; distinct reproductive

morphology: males transfer sperm with a conspicuous pri-

aprium, on the underside of the head (Parenti 1996).

Neostethus djajaorum Parenti & Louie, 1998

Euryhaline; endemic to Sulawesi Selatan; brackish waters

of coastal plains (Parenti & Eouie 1998).

BELONIFORMES

Adrianichthyidae

Ricefishes: Brackish and freshwaters; distributed through-

out the West-Pacific; 17 of the 35 species recognized are

endemic to Sulawesi, including species flocks in Eake

Poso and the Malili Eakes system, endemics in small, re-

mote lakes {Oryzias hadiatyae, O. soerotoi), one riverine

pelvic brooder O. eversi, and a riverine lineage species

from Southeast Sulawesi (Herder et al. 2012b, Kottelat

1990d, Mokodongan & Yamahira 2015, Parenti 2008, Par-

enti & Hadiaty 2010, Parenti et al. 2013).

Adrianichthys kruyti Weber, 1913

Adrianichthys kruytii (Soeroto & Tungka, 1991)

Freshwater; endemic to Eake Poso, Sulawesi Tengah;

pelagic; only a few specimens known (Kottelat 1990b,

Parenti 2008).

Adrianichthys oophorus (Kottelat, 1990)

Xenopoecilus oophorus Kottelat, 1990a

Freshwater; endemic to Eake Poso, Sulawesi Tengah;

pelagic pelvic brooder; abundant in open waters of the lake

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An annotated checklist of the inland fishes of Sulawesi

in 1995 (Parent! 2008); confirmed by own fieldwork in

2013; caught in the night at the surface; exploited by sub-

sistence fisheries (F.H., pers. obs.).

Adrianichthys poptae (Weber & de Beaufort, 1922)

Xenopoecilus poptae Weber & de Beaufort, 1 922

Freshwater; endemic to Lake Poso, Sulawesi Tengah;

pelagic, rare; not recorded by the authors (Kottelat 1990a,

Mokodongan & Yamahira 2015, Parent! 2008, Parent! &

Soeroto 2004, Soeroto & Tungka 1991, 1996).

Adrianichthys roseni Parenti & Soeroto, 2004

Adrianichthys kruyti (Weber, 1913)

Freshwater; endemic to Lake Poso, Sulawesi Tengah;

known from a single collection; likely pelagic; abdomi-

nal concavity points towards pelvic brooding (Parenti &

Soeroto 2004).

Oryzias asinua Parenti, Hadiaty, Lumbantobing &

Herder, 2013

Freshwater; endemic to Sulawesi Tenggara; known only

from the type locality: Asinua River, regency of Kendari

(Parenti et al. 2013).

Oryzias bonneorum Parenti, 2008

Xenopoecilus sarasinorum (Rosen, 1964)

Freshwater; endemic to Lake Lindu, Sulawesi Tengah;

probably pelagic (Parenti 2008).

Oryzias celeb ensis (Weber, 1894)

Haplochilus celebensis Weber, 1 894b

Aplocheilus celebensis Weber & de Beaufort, 1912

Freshwater; records from rivers, streams and Lake Tempe

in Sulawesi Selatan (Herder & Chapuis 2010, Parenti

2008), and East Timor (see Parenti 2008).

Oryzias eversi Herder, Hadiaty & Nolte, 2012

Freshwater; endemic; reported only from the type local-

ity in Sulawei Selatan, Tana Toraja; Salo Sadang drainage,

close to village Tilanga, about 8 km south of Rantepao;

the only known riverine Adrianichthyid with pelvic-brood-

ing reproduction (Herder et al. 2012b).

Oryzias hadiatyae Herder & Chapuis, 2010

Freshwater; endemic to Lake Masapi, Malili Lakes

(Larona) system, Sulawesi Selatan; Lake Masapi is a small

and shallow blackwater lake in the hills west of Lake

Towuti (Herder & Chapuis, 2010).

Oryzias javanicus (Bleeker, 1854)

Aplocheilus javanicus Bleeker, 1 854b

Euryhyaline; enter brackish waters; distributed from Thai-

land to Eombok, Borneo and Sulawesi (Parenti 2008);

records from Sulawesi Barat (3°20.143S 119°10.179E,

ZFMK 69890-69947).

Oryzias marmoratus (Aurich, 1935)

Aplocheilus marmoratus Amieb, 1935

Freshwater; endemic to Takes Towuti, Mahalona, Eontoa

[also Eantoa or Wawontoa], and adjacent streams, Sulawe-

si Selatan; possible hybridization among O. marmoratus

and O. profundicola in Take Towuti (Herder & Chapuis

2010, Kottelat 1990d, Mokodongan & Yamahira 2015).

Oryzias matan ensis (Aurich, 1935)

Freshwater; endemic to Take Matano, Malili Takes sys-

tem, Sulawesi Selatan; abundant around the lake; shallows

to deeper habitats along the coast (Kottelat 1990d, F.H.,

pers. obs.).

Oryzias nebulosus Parenti & Soeroto, 2004

Freshwater; endemic to Take Poso, Sulawesi Tengah (Par-

enti & Soeroto, 2004); small Oryzias (up to 33 mm SE);

benthopelagic; male courtship at rocky, open deeper habi-

tats (F.H., pers. obs.).

Oryzias nigrimas Kottelat, 1990

Freshwater; endemic to Take Poso, Sulawesi Tengah; ben-

thopelagic, at open habitats in the shallows (Kottelat

1990d, Parenti & Soeroto, 2004); comparatively abundant

(F.H., pers. obs.).

Oryzias orthognathus Kottelat, 1990

Freshwater; endemic to Take Poso, Sulawesi Tengah (Par-

enti 2008); characterized by conspicuously upwards di-

rected mouth, and rounded body (Kottelat 1990d, F.H.,

pers. obs.); tentatively pelagic, possibly rather ben-

thopelagic (Parenti 2008).

Oryzias profundicula Kottelat, 1990

Freshwater; endemic to Take Towuti, Sulawesi Selatan;

lacustrine, deep-bodied Oryzias with filamentous fm rays;

tends to inhabit deeper inshore habitats (Kottelat 1990d,

F.H., pers. obs.).

Oryzias sarasinorum (Popta, 1905)

Haplochilus sarasinorum Popta, 1905

Xenopoecilus sarasinorum Regan, 1911

Freshwater; endemic to Take Eindu, Sulawesi Tengah;

slender, pelagic pelvic brooder (Parenti 2008); Juveniles

recorded in the shallows of the lake in 2013.

Oryzias soerotoi Mokodongan, Tanaka & Yamahira, 2014

Freshwater; endemic to Take Tiu in Sulawesi Tengah, a

small (approx. 2 km long) blackwater lake draining to the

Eaa River; subadults in structured shallows; habitat of

adults unknown (Mokodongan et al. 2014).

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Oryzias wolasi Parenti, Hadiaty, Lumbantobing &

Herder, 2013

Freshwater; endemic; small, comparatively deep-bodied

Oryzias from streams in Sulawesi Tenggara, south of

Kendari (Parenti et al. 2013).

Oryzias woworae Parenti & Hadiaty, 2010

Freshwater; endemic to Muna island, off Sulawesi Teng-

gara (Parenti & Hadiaty 2010); small comparatively deep-

bodied, with conspicuous, colourful male ornamentation

(Parenti & Hadiaty 2010).

Zenarchopteridae

Halfbeaks: Freshwater; distributed in inland and coastal

habitats of the Indo-West Pacific; four genera in Sulawe-

si; Nomorhamphus and Dermogenys are viviparous, less

is known about the reproductive biology of Tondanichthys

and Zenarchopterus; Nomorhamphus inhabit hillstreams;

12 species endemic to Sulawesi, especially species-rich

(Huylebrouck et al. 2014). Halfbeak taxonomy largely fol-

lows the checklist of Collette 2004, supplemented by re-

cent descriptions (Anderson & Collette 1991, Collette

1995, Grier & Collette 1987; Lovejoy et al. 2004, Meis-

ner 2001)

Dermogenys orientalis (Weber, 1894)

Hemiramphus orientalis Weber, 1 894b

Dermogenys montanus Brembach, 1982

Freshwater: endemic to a hillstream of Maros Karst, Ban-

timurung, Sulawesi Selatan (Collette 2004, Parenti 2011).

Dermogenys vogti Brembach, 1982

Freshwater; most likely endemic to a hillstream on Lime-

stone Mountain at “Topobulu”, Sulawesi Selatan [locali-

ty not confirmed] (Brembach 1982, Collette 2004, Paren-

ti 2011).

Nomorhamphus brembachi Vogt, 1978

Nomorhamphus ravnaki Brembach, 1991

Nomorhamphus ravnaki australe Brembach, 1991

Nomorhamphus sanussii Brembach, 1991

Freshwater; endemic to Maros highland, Sulawesi Sela-

tan (Collette 2004, Parenti 2011).

Nomorhamphus celebensis Weber & de Beaufort, 1922

Freshwater; endemic to Lake Poso, Sulawesi Tengah (Col-

lette 2004, Parenti 2011).

Nomorhamphus ebrardtii (Popta, 1912)

Hemiramphus (Dermatogenus) ebrardtii Popta, 1912

Freshwater; endemic to Sulawesi Tenggara; records from

Wowoni Island (ZMH 7150); Muna Island, off Sulawesi

Tenggara; stream leading to Lake Towuti, Sulawesi Sela-

tan; stream crossing the road Soroako to Malili,

02°38.16LS, 121°12.920’E, ZFMK 49156-49176; Maros

Regency, Sungai Abbalu, Village Camba, Camba District,

MZB 21295 (Huylebrouck et al. 2014).

Nomorhamphus hageni (Popta, 1912)

Hemiramphus hageni Popta, 1912

Freshwater; endemic to Sulawesi Tenggara, Penango and

Rumbia valley (Collette 2004, Parenti 2011).

Nomorhamphus kolonodalensis Meisner & Louie, 2000

Freshwater; Sulawesi Tengah, Nuha drainage, north of

Lake Matano, Sulawesi Selatan, and city of Poso, district

of Kolonodale, Sulawesi Tengah (Collette 2004, Meisner

& Louie 2000, Parenti 2011).

Nomorhamphus lanceolatus Huylebrouck, Hadiaty &

Herder, 2014

Freshwater: endemic to Sungai Wawolambo, Sulawesi

Tenggara (Huylebrouck et al. 2014).

Nomorhamphus liemi Vogt, 1978

Nomorhamphus liemi snijdersi Vogt, 1978

Freshwater; endemic to Maros highland, Sulawesi Sela-

tan (Collette 2004, Parenti 2011, Vogt 1978).

Nomorhamphus megarrhamphus (Brembach, 1982)

Dermogenys megarrhamphus Brembach, 1982

Freshwater; endemic to Lakes Towuti and Mahalona, Su-

lawesi Selatan (Collette 2004; Parenti 2011).

Nomorhamphus rex Huylebrouck, Hadiaty & Herder,

2012

Freshwater; disjunct distribution in Sulawesi; records from

drainage adjacent to Malili Lakes: Stream Wewu,

Cerekang drainage, west of Lake Matano; stream within

Malili Lakes drainage: Toletole River at village Toletole;

and Sulawesi Selatan, Tana Toraja, about 8 km south of

Rantepao (Huylebrouck et al. 2012).

Nomorhamphus Sagittarius Huylebrouck, Hadiaty &

Herder, 2014

Freshwater: Endemic to three streams in Sulawesi Teng-

gara (Huylebrouck et al. 2014).

Nomorhamphus towoetii Ladiges, 1972

Normorhamphus towoetii Eadiges, 1972

Freshwater; records from Eake Towuti, Sulawesi Selatan

and Eake Poso, Sulawesi Tengah (Collette 2004, Eadiges

1972, Parenti 2011).

Nomorhamphus weberi (Boulenger, 1897)

Hemirhamphus weberi Boulenger, 1 897

Freshwater; endemic to Eakes Matano and Mahalona, Su-

lawesi Selatan (Boulenger 1897, Collette 2004, Parenti

2011 ).

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Tondanichthys kottelati Collette, 1995

Freshwater; endemic to Lake Tondano, Sulawesi Utara;

monotypic genus (Collette 1995, 2004, Parent! 2011).

Zenarchopterus gilli Smith, 1945

Freshwater; enter brackish and coastal waters (Donaldson

& Myers 2002); record from Sulawesi Selatan (4°07.456’S

119’37.196’E, ZFMK 69726-69740, 69838).

Zenarchopterus dispar (Valenciennes, in Cnvier & Va-

lenciennes, 1847)

Hemiramphus dispar Valenciennes, in Cuvier & Valenci-

ennes, 1847

Zenarchopterus maculosus Garman, 1903

Zenarchopterus vaisiganus Jordan & Seale, 1906

Potential: Freshwater; enter brackish and coastal waters;

distributed throughout the Indo-Pacific; no actual records

from Sulawesi (Donaldson & Myers 2002, Garman 1903).

CYPRINODONTIFORMES

Aplocheilidae

Rivulines: Freshwater; enter brackish waters; native in the

Neotropics, Africa and Southern Asia (Eschmeyer 2015,

Froese & Pauly 2014, Nelson 2006).

Aplocheilus panchax (Hamilton, 1822)

Esox panchax Hamilton, 1 822

Possibly introduced: Freshwater; native to India and

Southeast Asia; records from Buton (Tweedley et al.

2013), Sulawesi Utara (Haryono et al. 2002), Eakes of

Gorontalo (Haryono & Tjala-awidjaja2004), Sulawesi Se-

latan (3°4 1.589’ S 119°38.629’E, ZFMK 69557), Sulawe-

si Barat (2°39.081’S 119°12.436’E, ZFMK 69651), Eake

Poso, Sulawesi Tengah (visual record F.H.) and the Malili

Eakes system, Sulawesi Selatan (Herder et al. 2012a).

Poeciliidae

Eivebearers: Freshwater; enter brackish waters; native to

North, Central and South America; several worldwide in-

troduced species (Nelson 2006).

Gambusia affinis (Baird & Girard, 1853)

Heterandria affinis Baird & Girard, 1853

Introduced: Freshwater; native to North and Central Amer-

ica; introduced into warm waters almost worldwide (Pyke

2006); record from Eake Poso, Sulawesi Tengah (visual

record F.H.).

Gambusia holbrooki Girard, 1859

Introduced: Freshwater; native to North and Central Amer-

ica; introduced to warm waters almost worldwide; likely

confused with G. affinis (Girard 1859, Pyke 2006).

Poecilia reticulata Peters, 1859

Introduced: Freshwater; native to northern South Ameri-

ca; introduced almost worldwide; record from Malili

Eakes system, Sulawesi Selatan (Herder et al. 2012a),

Eake Poso, Sulawesi Tengah (1°46.29’S 120°42.98’E,

ZFMK 69801-69803).

GASTEROSTEIFORMES

Syngnathidae

Pipefishes, Seahorses: Catadromous; enter marine, brack-

ish and freshwaters; distributed throughout the Atlantic,

Indo-Pacific (Nelson 2006, Wilson & Orr 2011).

Belonichthys mento (Bleeker, 1856)

Syngnathus mento BlQokQY, 1856a

Catadromous; record from Manado, Sulawesi Utara

(Bleeker 1856a), Buton (Tweedley et al. 2013).

Coelonotus biocellatus Gunther, 1870

Potential: Catadromous; no actual records for Sulawesi;

distributed throughout the East Indian Archipelago (Giin-

ter 1870, cited in Kottelat 2013).

Coelonotus leiaspis (Bleeker, 1854)

Syngnathus leiaspis Bleeker, 1854c

Microphis leiaspis Bleeker, 1 854c

Syngnathus budi Bleeker, 1856a

Catadromous; distributed throughout the Indo-Pacific

(Dawson 1985); records from Manado, Sulawesi Utara

(Bleeker 1856a), Buton (Tweedley et al. 2013).

Doryichthys boaja (Bleeker, 1850)

Syngnathus boaja Bleeker, 1 850

Doryichtys spinosus Kaup, 1856

Catadromous; distributed throughout Asia (Dawson

1985); record from Makassar, Sulawesi Selatan (Kaup

1856).

Hippichthys cyanospilos (Bleeker, 1854)

Potential: Catadromous; no actual records for Sulawesi;

distributed throughout the Indo-Pacific (Bleeker 1 854c).

Hippichthys heptagonus Bleeker, 1849

Potential: Catadromous; no actual records for Sulawesi;

distributed throughout the Indo-Pacific (Bleeker 1849b).

Hippichthys spicifer (Riippell, 1838)

Syngnathus spicifer Kuyi^QW, 1838

Catadromous; distributed throughout the Indo-Pacific

(Dawson 1985); record from Sulawesi Selatan

(04°14.475’S 119°36.826’E, ZFMK 066326).

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Hippocampus waleananus Gomon & Kuiter, 2009

Catadromous; endemic to the Togian islands, off Sulawe-

si Tengah (Gomon & Kuiter 2009).

Lophocampus retzii (Bleeker, 1856)

Syngnathus retzii Bleeker, 1856a

Catadromous; distributed throughout the Indo-Pacific;

record from Manado, Sulawesi Utara (Bleeker 1856a).

Microphis ocellatus (Duncker, 1910)

Doryichthys ocellatus Duncker, 1910

Catadromus; distributed throughout the Indo-Pacific

(Dawson 1984); record from Sulawesi Tengah

(00°55.395’S 122°52.962’E, ZFMK 066065).

Oostethus brachyurus (Bleeker, 1854)

Syngnathus brachyurus Bleeker, 1 854c

Syngnathus polyacanthus Bleeker, 1856a

Doryichthys auronitens Kaup, 1 856

Catadromous; distributed throughout the Indo-Pacific;

record from Manado, Sulawesi Utara (Bleeker 1856a),

Makassar, Sulawesi Selatan (Dawson 1985, Kaup 1856).

Oostethus manadensis (Bleeker, 1856)

Syngnathus manadensis Bleeker, 1856a

Catadromous; distributed throughout the Indo-Pacific

(Dawson 1985); record from Manado, Sulawesi Utara

(Bleeker 1856a).

SYNBRANCHIFORMES

Synbranchidae

Swamp eels: Freshwater; entering brackish waters; distrib-

uted throughout Central and South America, Mexico, the

Indo Australian Archipelago, Asia and West Africa (Nel-

son 2006, Rosen & Greenwood 1976).

Monopterus albus (Zuiew, 1793)

Muraena alba Zuiew, 1793

Introduced: Freshwater; enter brackish waters; records

from Sulawesi Utara (Haryono et al. 2002), Malili Lake

drainage, Sulawesi Selatan (Herder et al. 2012a).

SCORPAENIFORMES

Tetrarogidae

Wasp fishes: Euryhaline; enter brackish and freshwaters;

distributed throughout the Indo-West Pacific (Eschmey-

er 2015, Froese & Pauly 2014 Nelson 2006).

Neovespicula depressifrons (Richardson, 1848)

Apistus plagiometopon Bleeker, 1853a

Euryhaline; entering brackish waters; record from Bu-

lukumba, Sulawesi Selatan (Bleeker 1853a).

Tetraroge barbata (Cuvier, in Cuvier & Valenciennes,

1829)

Apistus barbatus Cuvier, in Cuvier & Valenciennes, 1 829

Euryhaline; enter brackish and freshwaters (Fricke et al.

2011); record from Sulawesi Tengah (00°55.395’S

122°52.962’E, ZFMK 066003).

Tetraroge nigra (Cuvier, in Cuvier & Valenciennes,

1829)

Apistus nigra Cuvier, in Cuvier & Valenciennes, 1 829

Euryhaline; enter brackish waters; record from Buton

(Tweedley et al. 2013).

Platycephalidae

Flatheads: Euryhaline; enter brackish waters; distributed

throughout the Indo-Pacific (Nelson 2006).

Grammoplites scaber (Linnaeus, 1758)

Cottus scaber Linnaeus, 1758

Potential: Euryhaline; no actual records for Sulawesi; dis-

tributed throughout the Indo-Pacific (Knapp 1999).

PERCIFORMES

PERCOIDEI

Ambassidae

Asiatic glassfishes: Euryhaline; enter brackish and fresh-

waters; distributed throughout the Indo-West Pacific (An-

derson & Heemstra 2003, Nelson 2006).

Ambassis gymnocephala (La Cepede, 1802)

Lutjan gymnocephale La Cepede, 1 802

Ambassis dussumieri Cuvier, in Cuvier & Valenciennes,

1828

Euryhaline; enter brackish and freshwaters; distributed

throughout the Indo-Pacific (Anderson & Heemstra

2003); record from Sulawesi Tengah (00°55.395’S

122°52.962’E, ZFMK 066031-066039).

Ambassis interrupta Bleeker, 1853

Euryhaline; enter brackish and freshwaters; distributed

throughout the Indo-Pacific (Anderson & Heemstra

2003); type locality in Sulawesi (Bleeker 1853c).

Ambassis miops Gunther, 1872

Euryhaline; enter brackish and freshwaters; distributed

throughout the Indo-Pacific (Kottelat 2013); record from

Buton (Tweedley et al. 2013).

Ambassis urotaenia Bleeker, 1852

Potential: Euryhaline; enter brackish and freshwaters (An-

derson & Heemstra 2003); no actual records for Sulawe-

si; distributed throughout the Indo-West Pacific (Bleeker

1852c).

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Ambassis vachellii Richardson, 1846

Ambassis telkaraWhitlQy, 1935a

Potential: Euryhaline; enter braekish and freshwaters; no

actual records for Sulawesi; distributed throughout the In-

do-Pacific (Kottelat 2013).

Opistognathidae

Jawfishes: Euryhaline; enter brackish waters; distributed

throughout the Western and Central Atlantic and Indo-Pa-

cific (Kottelat 2013; Nelson 2006, Smith- Vaniz 1999).

Stalix moenensis (Popta, 1922)

Gnathypops moenensisVo^tdL, 1922

Euryhaline; enter brackish waters (Smith- Vaniz 1999);

record from Muna Island, off Sulawesi Tenggara (Pop-

tal922).

Family Carangidae

Jacks, Pompanos: Marine; juveniles enter brackish waters;

distributed throughout the Indo-Pacific and Atlantic

(Holland et al. 1996, Nelson 2006).

Caranx melampygus Cuvier, in Cuvier & Valenciennes,

1833

Potential: Euryhaline; enter brackish waters; no actual

records for Sulawesi; distributed throughout the Indo-Pa-

cific (Holland et al. 1996).

Caranx papuensis Alleyne & Macleay, 1877

Euryhaline; enter brackish waters; distributed throughout

the Indo-Pacific (Holland et al. 1996); record from Ba-

ton (Tweedley et al. 2013).

Caranx sexfasciatus Quoy & Gaimard, 1825

Potential: Euryhaline; enter brackish waters; no actual

records for Sulawesi; distributed throughout the Indo-Pa-

cific (Holland et al. 1996).

Scontberoides lysan (Forskal, 1775)

Scomber lysan Forskal, 1775

Potential: Euryhaline; enter brackish waters; no actual

records for Sulawesi; distributed throughout the Indo-Pa-

cific (Smith- Vaniz 1999).

Selaroides leptolepis (Cuvier, in Cuvier & Valenciennes,

1833)

Caranx leptolepis Cuvier, in Cuvier & Valenciennes, 1 833

Potential: Euryhaline; enter brackish waters; no actual

records for Sulawesi; distributed throughout the Indo-Pa-

cific (Smith- Vaniz 1999).

Leiognathidae

Slimys, Slipmouths, Ponyfishes: Euryhaline; enter brack-

ish and freshwaters; distributed throughout the Indo-West

Pacific (Eschmeyer 2015, Froese & Pauly 2014; Nelson

2006).

Eubleekeria splendens (Cuvier, 1829)

Equula splendens Cuvier, 1 829

Potential: Euryhaline; no actual records for Sulawesi; dis-

tributed throughout the Indo-Pacific; enter brackish wa-

ters (Kottelat 2013).

Lutjanidae

Snappers: Euryhaline; enter brackish waters; distributed

throughout the Atlantic and Indo-Pacific (Eschmeyer

2015, Froese & Pauly 2014, Nelson 2006).

Lutjanus argentimaculatus (Forskal, 1775)

Sciaena argentimaculata Forskal, 1775

Mesoprion taeniops Valenciennes, in Cuvier & Valenci-

ennes, 1830

Euryhaline; enter brackish waters; distributed throughout

the Indo-West Pacific (Anderson & Allen 1999); record

from Sulawesi freshwaters (Valenciennes, in Cuvier & Va-

lenciennes 1830).

Lutjanus bohar (Forskal, 1775)

Sciaena bohar ForskM, 1775

Potential: Euryhaline; enter brackish waters; no actual

records for Sulawesi; distributed throughout the Indo-West

Pacific (Anderson & Allen 1999).

Lutjanus ehrenbergii (Peters, 1869)

Mesoprion ehrenbergii Peters, 1 869

Lutjanus oligolepis Bleeker, 1873a

Euryhaline; enter brackish waters; distributed throughout

the Indo-West Pacific (Anderson & Allen 1999); record

from Makassar, Sulawesi Selatan (Bleeker 1873a).

Lutjanus fulviflamma (Forskal, 1775)

Sciaena fulviflamma Forskal, 1775

Potential: Euryhaline; enter brackish waters; no actual

records for Sulawesi; distributed throughout the Indo-Pa-

cific (Anderson & Allen 1999).

Lutjanus fulvus (Forster, in Schneider, 1801)

Holocentrus fulvus Forster, in Schneider, 1801

Potential: Euryhaline; enter brackish waters; no actual

records for Sulawesi; distributed throughout the Indo-Pa-

cific (Anderson & Allen 1999).

Lutjanus fuscescens (Valenciennes, in Cnvier & Valen-

ciennes, 1830)

Mesoprion fuscescens Valenciennes, in Cuvier & Valen-

ciennes, 1830

Euryhaline; enter brackish waters; distributed throughout

the Indo-West Pacific (Anderson & Allen 1999); record

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Friedrich Wilhelm Miesen et al.

from Sulawesi freshwaters (Valenciennes, in Cuvier & Va-

lenciennes 1830).

Lutjanus maxweberi Popta, 1921

Lutianus max weberi Popta, 1921

Euryhaline; enter brackish and freshwaters; distributed

throughout the Indo-Pacific (Anderson & Allen 1999);

record from Kali La River, Muna Island, off Sulawesi

Tenggara (Popta 1921).

Haemulidae

Grunts: Euryhaline; enter brackish and freshwaters distrib-

uted throughout the Atlantic and Indo-Pacific (Eschmey-

er 2015, Froese & Pauly 2014).

Pomadasys argenteus (Forskal, 1775)

Sciaena argentea Forskal, 1775

Pristipoma manadense Gunther, 1 872b

Euryhaline; enter brackish waters; distributed throughout

the Indo-West Pacific (McKay 2001); record from Man-

ado, Sulawesi Utara (Gunther 1872b).

Nemipteridae

Threadfin breams: Euryhaline; enter brackish waters; dis-

tributed throughout the Indo-West-Pacific (Eschmeyer

2015, Froese & Pauly 2014, Nelson 2006).

Nemipterus peronii (Valenciennes, in Cnvier & Valen-

ciennes, 1830)

Dentex peronii Valenciennes, in Cuvier & Valenciennes,

1830

Dentex Smithii Steindachner, 1 868

Euryhaline; enter brackish waters; distributed throughout

the Indo-West Pacific (Russell 2001); record from Tiworo,

Muna Island, off Sulawesi Tenggara (Steindachner 1 868).

Lethrinidae

Emperors, Scavengers: Euryhaline; enter brackish waters;

distributed from West Africa to the Indo-West Pacific (Es-

chmeyer 2015, Froese & Pauly 2014, Nelson 2006).

Lethrinus nebulosus (Forskal, 1775)

Sciaena nebulosaVoxs\M, Ml 5

Potential: Euryhaline; enter brackish waters; no actual

records for Sulawesi; distributed throughout the Indo-Pa-

cific (Carpenter 2001a).

Polynemidae

Threadfins: Euryhaline; enter brackish and freshwaters;

distributed in all tropical and subtropical regions (Moto-

mura 2004, Nelson 2006).

Polydactylus microstoma (Bleeker, 1851)

Polynemus microstoma Bleeker, 1851a

Euryhaline; enter brackish waters; distributed throughout

the Indo-Pacific (Feltes 2001); record from Bulukumba,

Sulawesi Selatan (Bleeker 1851a).

Sciaenidae

Drums: Euryhaline; enter brackish and freshwaters; Dis-

tributed throughout the Atlantic and Indo-Pacific (Es-

chmeyer 2015, Froese & Pauly 2014, Nelson 2006).

Nibea soldado (La Cepede, 1802)

Holocentrus soldado La Cepede, 1 802

Corvina celebica Bleeker, 1 854d

Euryhaline; enter brackish waters; distributed throughout

the Indo-West Pacific (Sasaki 2001); record from Makas-

sar, Sulawesi Selatan (Bleeker 1854d).

Mullidae

Goatfishes: Euryhaline; enter brackish waters; distributed

throughout the Atlantic and Indo-Pacific (Eschmeyer

2015, Froese & Pauly 2014; Nelson 2006).

Upeneus tragula Richardson, 1846a

Upeneus sundaicus yar. caudalis Popta, 1921

Euryhaline; enter brackish waters; distributed throughout

the Indo-Pacific (Randall 2001); record from Tiworo, Mu-

na Island, off Sulawesi Tenggara (Popta 1921).

Toxotidae

Archerfishes: Euryhaline; enter brackish and freshwaters;

distributed throughout the Indo-Pacific (Berra 2001 , Nel-

son 2006).

Toxotes chatareus (Hamilton, 1822)

Coins chatareus Hamilton, 1 822

Potential: Euryhaline; enter brackish waters; no actual

records for Sulawesi; distributed throughout the Indo-Pa-

cific (Allen 1991, 2001).

Toxotes jaculatrix (Pallas, in Schlosser, 1767)

Sciaena jaculatrix Pallas, in Schlosser, 1767

Potential: Euryhaline; enter brackish waters; no actual

records for Sulawesi; distributed throughout the Indo-Pa-

cific (Allen 1991, 2001).

Terapontidae

Grunters, Tigerperches: Euryhaline; enter brackish and

freshwaters; distributed throughout the Indo-West Pacif-

ic (Berra 2001, Nelson 2006, Vari 2001).

Lagusia micracanthus (Bleeker, 1860)

Datnia micracanthus Bleeker, 1860

Therapon {Datnia) micracanthus Bleeker, 1873b

Terapon micracanthus Fowler, 1931

Papuservus micracanthus Munro, 1958

Euryhaline; endemic to Sulawesi; enter brackish and fresh-

waters; records from Lagusi, Amparang, Bantimurung,

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An annotated checklist of the inland fishes of Sulawesi

Cendrana, Leang-leang, Maros, Menralang, Samanggi and

Saripa rivers, Manjali Spring, Sulawesi Selatan (Bleeker

1860, Fowler 1931, Vari & Hadiaty 2012).

Ter upon jarbua (Forskal, 1775)

SciaenajarbuaYoxskkX, 111 5

Euryhaline; enter brackish and freshwaters; distributed

throughout the Indo-Pacific (Vari 2001); record from Su-

lawesi Selatan (04°14.475’S 119°36.826’E, ZFMK

066043).

Kuhliidae

Flagtails: Euryhaline; enter brackish and freshwaters; dis-

tributed throughout the Indo-Pacific (Berra 2001, Nelson

2006).

Kuhlia marginata (Cuvier, in Cuvier and Valenciennes,

1829)

Dules marginatus Cuvier, in Cuvier & Valenciennes, 1 829

Euryhaline; enter brackish and freshwaters; distributed

throughout the Indo-West Pacific (Carpenter 2001b);

records from Buton (Tweedley et al. 2013), Sulawesi Utara

(Haryono et al. 2002), Sulawesi Selatan (3°30.822’S

119°32.267’E, ZFMK 69614-69615).

Kulia rupestris (La Cepede, 1802)

Centropomus rupestris Ea Cepede, 1 802

Euryhaline; enter brackish and freshwaters; distributed

throughout the Indo-West Pacific (Carpenter 2001b);

record from Buton (Tweedley et al. 2013).

LABROIDEI

Cichlidae

Cichlids: Diverse group in marine, brackish and freshwa-

ter environments; distributed throughout the Neo- and

Palaeotropics; not native to Sulawesi; several species in-

troduced worldwide (Berra 2001, Eschmeyer 2015,

Froese & Pauly 2014; Nelson 2006).

“Flowerhorn” cichlid

Introduced: Freshwater; hybrid of neotropical species;

records from the Malili Eakes system, Sulawesi Selatan

(Herder et al. 2012a), Poso River, Sulawesi Tengah (vi-

sual record F.H.).

Melanochromis cyaneorhabdos (Bowers & Stauffer,

1997)

Introduced: Freshwater; native to Eake Malawi, East

Africa; record from the Malili Eakes system, Sulawesi Se-

latan (Herder et al. 2012) and Eake Poso, Sulawesi Ten-

gah (visual record F.H.).

Oreochromis mossambicus (Peters, 1852)

Chromis mossambicus Peters, 1 852

Introduced: Freshwater; native to Africa; record from the

Malili Eakes system, Sulawesi Selatan (Herder et al.

2012a, Nasution & Aisyah 2013), Eake Poso, Sulawesi

Tengah, and various streams (visual record F.H.).

Oreochromis niloticus (Linnaeus, 1758)

Introduced: Freshwater; native to Africa; record from the

Malili Lakes system (Nasution & Aisyah 2013), Sulawe-

si Selatan and Sulawesi Barat (2°39.08r S 1 19°12.436’E,

ZFMK 69650).

Scaridae

Parrotfishes: Euryhaline; enter brackish waters; distributed

throughout the Atlantic and Indo-Pacific (Eschmeyer

2015, Froese & Pauly 2014, Nelson 2006).

Chlorurus sordidus (Forskal, 1775)

Scarus sordidusVoxskkX, 1775

Scarus celebicus Bleeker, 1 854d

Euryhaline; enter brackish waters (Bellwood 2001),

record from Makassar, Sulawesi Selatan (Bleeker 1854d).

BLENNIOIDEI

Blenniidae

Combooth blennies: Euryhaline; enter brackish and

freshwaters; distributed throughout the Atlantic and Indo-

Pacific (Eschmeyer 2015, Froese & Pauly 2014, Nelson

2006).

Meiacanthus anema (Bleeker, 1852)

Petroskirtes anema Bleeker, 1 852c

Euryhaline; enter brackish waters; distributed throughout

the Indo-Pacific (Allen 1991); record from Kabaena

(Tweedley et al. 2013).

CALLIONYMOIDEI

Callionymidae

Dragonets: Euryhaline; two species in freshwaters; distrib-

uted throughout the Indo-West Pacific (Eschmeyer 2015,

Froese & Pauly 2014, Nelson 2006).

Eleutherochir opercularis (Valenciennes, in Cuvier &

Valenciennes, 1837)

Callionymus opercularis Valenciennes, in Cuvier & Va-

lenciennes, 1837

Brachycallionymus mirus Herre, 1936

Euryhaline; enter brackish waters; distributed throughout

the Indo-West Pacific (Talwar & Jhingran 1991); record

from Lembeh Strait, north coast of Sulawesi (Herre, 1936).

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Friedrich Wilhelm Miesen et al.

GOBIOIDEI

Rhyacichthyidae

Loach gobies: Freshwater; distributed throughout the In-

do-West Pacific (Berra 2001, Nelson 2006).

Rhyacichthys aspro (Valenciennes, in Cnvier and Va-

lenciennes, 1837)

Platyptera aspro Valenciennes, in Cuvier & Valenciennes,

1837

Anadromous; enter brackish and freshwaters; distributed

throughout Indo-Pacific (Allen 1991); records from Ba-

ton (Tweedley et al. 2013), Sulawesi Utara (Haryono et

al. 2002), Sulawesi Barat (3°16.65rS 118°51.929’E,

ZFMK 6848-6850).

Eleotrididae

Sleepers: Euryhaline; enter brackish and freshwaters; dis-

tributed worldwide in tropical and subtropical regions

(Berra 2001, Eschmeyer 2015, Froese & Pauly 2014, Nel-

son 2006).

Belobranchus belobranchus (Valenciennes, in Cnvier &

Valenciennes, 1837)

Eleotris belobrancha Valenciennes, in Cuvier & Valenci-

ennes, 1837

Anadromous; distributed throughout the Indo-Pacific

(Allen 1991); records from Manado, Sulawesi Utara (Va-

lenciennes, in Cuvier & Valenciennes, 1 837), Buton and

Kabaena (Tweedley et al. 2013), Sulawesi Utara (Hary-

ono et al. 2002) and Sulawesi Barat (2°39.308’S

119°12.095’E, ZFMK 69631; 2°39.08rS 119 12.436’E,

ZFMK 69642-69647; 2°38.428’S 119°09.294’E, ZFMK

69670; 2°37.368’S 119°08.784’E, ZFMK 69699).

Belobranchus segura Keith, Hadiaty & Lord, 2012

Freshwater; enter brackish and freshwaters; described

from Halmahera and Irian Jaya (Keith et al. 2012); record

from Sulawesi Barat (3°16.65US 118°51.929’E, ZFMK

69814-69815).

Bostrychus microphthalmus Hoese & Kottelat, 2005

Freshwater; endemic to Gua Tanette cave, Sulawesi Se-

latan; genus poorly defined and likewise poorly known

(Hoese & Kottelat 2005).

Bunaka gyrinoides (Bleeker, 1853)

Eleotris gyrinoides Bleeker, 1 853c

Anadromous; distributed throughout the Indo-West Pacif-

ic (Allen 1991); records from Buton (Tweedley et al.

2013), Sulawesi Selatan (3°41.589’S 119°38.629’E,

ZFMK 69556, 69558-69559), Sulawesi Barat (3° 16. 65 US

118°51.929’E, ZFMK 69823).

Butis amboinensis (Bleeker, 1854)

Eleotris amboinensis Bleeker, 1 854a

Anadromous; enter brackish and freshwaters; distributed

throughout the Indo-West Pacific (Yokoo et al. 2006);

record from Buton (Tweedley et al. 2013).

Butis butis (Hamilton, 1822)

Cheilodipterus butis Hamilton, 1 822

Potential: Anadromous; enter braekish and freshwaters; no

actual records for Sulawesi; distributed throughout the In-

do-Pacific (Allen et al. 2002).

Eleotris fusca (Schneider, 1801)

Poecilia fusca Schneider, 1801

Potential: Anadromous; enter brackish and freshwaters as

adults; no actual records for Sulawesi; distributed through-

out the Indo-Pacific (Maeda et al. 2007).

Eleotris melanosoma Bleeker, 1853

Anadromous; enter brackish and freshwaters; distributed

throughout the Indo-Pacific (Bleeker 1853c, Maeda et al.

2007); records from Sulawesi Selatan (3°30.822'S

119°32.267’E, ZFMK 69595, 69616-69617), Sulawesi

Barat (3°16.65rS 118°51.929’E, ZFMK 69806).

Giuris margaritaceus (Valenciennes, in Cuvier & Va-

lenciennes, 1837)

Eleotris margaritacea Valenciennes, in Cuvier & Valen-

ciennes, 1837

Anadromous; enter brackish and freshwaters; distributed

throughout the Indo-Pacific (Allen et al. 2002); record

from Sulawesi Barat (3°16.65rS 118°51.929’E, ZFMK

69804-69805, 69822).

Oxyleotris marmorata (Bleeker, 1852)

Eleotris marmorataBlQokQr, 1852e

Anadromous; enter brackish and freshwaters; distributed

throughout the Indo-Pacific (Earson & Murdy 2001);

record from Sulawesi Selatan (3°41.589'S 119°38.629’E,

ZFMK 69512-69513, 69550-69551).

Gobiidae

Gobies: Euryhaline, catadromous, anadromous and fresh-

water; includes a total of 1725 species in 25 1 genera (Berra

2001, Eschmeyer 2015, Froese & Pauly 2014, Nelson

2006).

Acentrogobius janthinopterus (Bleeker, 1853)

Gobius janthinopterus Bleeker, 1 853b

Gobius hemigymnopomus Bleeker, 1856a

Amphidromous; enter brackish and freshwaters; distrib-

uted throughout the Indo-Pacific (Donaldson & Myers

2002); record from Makassar, Sulawesi Selatan (Bleeker

1853b).

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An annotated checklist of the inland fishes of Sulawesi

Acentrogobius moloanus (Herre, 1927)

Anadromous; enter brackish and freshwaters; distributed

throughout the Western Pacific (Blaber & Milton 1990);

record from Sulawesi Selatan (119°36.826’E 04°14.475’S,

ZFMK 066064).

Amblygobius decussatus (Bleeker, 1855)

Gobius decussatus Bleeker, 1855a

Anadromous; enter brackish and freshwaters; distributed

throughout the Western Central Pacific (Myers 1991);

record from Manado, Sulawesi Utara (Bleeker 1855a).

Awaous grammepomus (Bleeker, 1849)

Gobius grammepomus Bleeker, 1 849c

Anadromous; enter brackish and freshwaters; distributed

throughout Asia (Watson 1992); records from Sulawesi

Selatan (3°41.589'S 119°38.629’E, ZFMK 69502-69511,

69535-69549; 3°30.822’S 119°32.267’E, ZFMK 69602-

69603, 69628), Sulawesi Barat(2°38.428'S 119°09.294’E,

ZFMK 69653-69659; 2°37.915'S 119°09.488’E, ZFMK

69673-69676).

Bathy gobius petrophilus (Bleeker, 1853)

Gobius petrophilus Bleeker, 1853d

Gobius villosus Weber, 1909

Anadromous; entering brackish and freshwaters; distrib-

uted throughout the Indo-West Pacific; record from Man-

ado, Sulawesi Utara (Weber 1 909, Weber & de Beaufort

1953).

Cryptocentroides insignis (Seale, 1910)

Amblyogobius insignis Seale, 1910

Cryptocentroides dentatus Popta, 1922

Anadromous; entering brackish and freshwaters; distrib-

uted throughout the Western Pacific; record from Raha,

Muna Island, off Sulawesi Tenggara (Popta 1922).

Drombus bontii (Bleeker, 1849)

Gobius bontii Bleeker, 1 849c

Acentrogobius elberti Popta, 1921

Anadromous; entering brackish and freshwaters; distrib-

uted throughout the Indo-West Pacific (Kottelat 2013);

record from Raha, Muna Island, off Sulawesi Tenggara

(Popta 1921).

Glossogobius celebius (Valenciennes, in Cnvier & Va-

lenciennes, 1837)

Freshwater; endemic to Sulawesi; records from Eake

Towuti, Sulawesi Selatan (119°37.295’E 04°07.540’S,

ZFMK 066014); Sulawesi Selatan (3°41.589'S

119°38.629’E, ZFMK 69517; 3°30.822’S 119°32.267’E,

ZFMK 69596-69601, 69612-69613, 69626-69627), Su-

lawesi Barat (2°37.368’S 119°08.784’E, ZFMK 69697-

69698).

Bonn zoological Bulletin 64 (2): 77-106

Glossogobius flavipinnis (Anrich, 1938)

Freshwater; endemic to Eake Towuti, Sulawesi Selatan;

lacustrine dwarf species (Hoese et al. 2015).

Glossogobius intermedius (Anrich, 1938)

Freshwater; endemic to Eakes Mahalona and Towuti, Su-

lawesi Selatan (Kottelat et al. 1993).

Glossogobius matanensis (Weber, 1913)

Freshwater; endemic to Eakes Matano, Mahalona, Towu-

ti, and Eontoa, Sulawesi Selatan (Kottelat et al. 1993).

Glossogobius mahalonensis Hoese, Hadiaty & Herder,

2015

Freshwater; endemic to Eake Mahalona, Sulawesi Sela-

tan; so far known from one single site within the lake

(Hoese et al. 2015).

Gnatholepis anjerensis (Bleeker, 1851)

Gobius anjerensis Bleeker, 1851b

Anadromous; enter brackish waters; distributed through-

out the Indo-Pacific; record from Bunaken Island, off

Manado, Sulawesi Utara (Bleeker, 1851b).

Lentipes mekonggaensis Keith, Hadiaty, Hubert, Bns-

son & Lord, 2014

Presumably amphidromous; terra typica is a fast flowing

stream in Sulawesi Tenggara (Keith et al. 2014).

Lentipes watsoni Allen, 1997

Presumably anadromous; record from Sulawesi Tenggara

(02°65.035'S 121°06.855'E, ZFMK 45041); type locali-

ty in Papua New Guinea; determination based on one male

specimen matching the diagnosis of the species descrip-

tion (Allen 1997).

Lophogobius bleekeri Popta, 1921

Anadromous; enter brackish and freshwaters; distributed

throughout the Western Pacific; recorded from Raha, Mu-

na Island, off Sulawesi Tenggara (Popta 1921).

Mugilogobius adeia Larson & Kottelat, 1992

Freshwater; endemic to Lake Matano, Malili Lakes sys-

tem, Sulawesi Selatan (Larson 2001, Larson & Kottelat

1992).

Mugilogobius amadi (Weber, 1913)

Gobius amadi Weber, 1913

Weberogobius amadi Koumans, 1953

Freshwater; endemic to Lake Poso, Sulawesi Tengah; orig-

inally abundant, not reported since 1985 (Larson 2001).

Mugilogobius chulae (Smith, 1932)

Vaimosa chulae Smith, 1932

Freshwater; distributed throughout the South-East Asian

©ZFMK

Friedrich Wilhelm Miesen et al.

archipelago, southern Japan, Sri Lanka, Thailand,

Malaysia, Taiwan and Singapore (Huang et al. 2013, Tan

& Lim, 2004); record from Boloang, Sulawesi Utara (Lar-

son 2001, Larson et al. 2008).

Mugilogobius hitam Larson, Geiger, Hadiaty & Her-

der, 2014

Freshwater; most likely endemic to Lake Towuti, Sulawe-

si Selatan (Larson et al. 2014).

Mugilogobius latifrons (Boulenger, 1897)

Gobius latifrons Boulenger, 1 897

Gobius latifrons WQhQX, 1913

Vaimosa latifrons Aurich, 1938

Freshwater; endemic to streams and lakes of the Malili

Lakes system, Sulawesi Selatan (Larson et al. 2014).

Mugilogobius lepidotus Larson, 2001

Freshwater; endemic to Lake Towuti, Malili Lakes sys-

tem, Sulawesi Selatan; recorded by F.H. in 2002 and 2004.

Mugilogobius mertoni (Weber, 1911)

Gobius mertoni Wober, 1911

Tamanka mindora Herre, 1945

Vaimosa layia Herre, 1953

Tamanka mertoni Koumans, 1953

Potential: Anadromous; distributed throughout the Indo-

Pacific; no actual records for Sulawesi (Heemstra et al.

2004, Huang et al. 2013, Larson 2001, Larson et al. 2013,

Manilo & Bogorodsky 2003).

Mugilogobius rexi Larson, 2001

Freshwater; endemic to Lake Mahalona and Lake Towu-

ti, Sulawesi Selatan (Larson 2001).

Mugilogobius sarasinorum (Bonlenger, 1897)

Gobius sarasinorum Boulenger, 1 897

Tamanka sarasinorum Koumans, 1953

Freshwater; endemic to Lake Poso, Sulawesi Tengah;

highly abundant in 2013 (F.H. pers. obs.).

Oxyurichthys tentacularis (Valenciennes, in Cnvier &

Valenciennes, 1837)

Gobius tentacularis Valenciennes, in Cuvier & Valenci-

ennes, 1837

Oxyurichthys rumbia Popta, 1922

Anadromous; enters brackish and freshwaters; distributed

throughout the Indo-West Pacific (Mauge 1986); record

from Rumbia, Sulawesi Tenggara (Popta 1922).

Periophthalmus kalolo Lesson, 1831

Periophthalmus argentilineatus Valenciennes, in Cuvier

& Valenciennes, 1 837

Anadromous; enter brackish and freshwaters; records from

Buton (Tweedley et al. 2013), Sulawesi Selatan

(04°14.475'S 119°36.826'E, ZFMK 066001-066002).

Redigobius bikolanus (Herre, 1927)

Vaimosa bikolana Herre, 1927

Anadromous; enter brackish and freshwaters; distributed

throughout Borneo, Sulawesi and the Philippines (Kotte-

lat et al. 1993); record from Buton (Tweedley et al. 2013).

Redigobius penango (Popta, 1922)

Pseudogobius penango Popta, 1922

Anadromous; enter brackish and freshwaters; distributed

in Indonesia; record from Penango, Sulawesi Tenggara

(Larson 2010, Popta 1922).

Schismatogobius bruynisi de Beaufort, 1912

Anadromous; enter brackish and freshwaters; distributed

throughout Indonesia and the Philippines (de Beaufort

1912, Keith & Lord 2011, Kottelat & Whitten 1996, Kot-

telat et al. 1993); record from Sulawesi Tengah

(121°06.855’E 02°56.035’S, ZFMK 45049).

Schismatogobius marmoratus (Peters, 1868)

Gobiosoma marmorataVQtQVS, 1868

Anadromous; enter brackish and freshwaters; distributed

throughout Sulawesi, Philippines and Japan (Keith & Eord

2011, Kottelat et al. 1993); record from Sulawesi Tengah

(02°56.035’S 121°06.855’E, ZFMK 066059; 00°55.395’S

122°52.962’E, ZFMK 066060).

Sicyopterus cynocephalus (Valenciennes, in Cuvier &

Valenciennes, 1837)

Sicydium cynocephalum Valenciennes, in Cuvier & Valen-

ciennes, 1837

Anadromous; enter brackish and freshwaters; distributed

throughout Indonesia and the Philippines (Keith & Eord

2011, Koumans 1953); records from Manado harbour, Su-

lawesi Utara (Valenciennes, in Cuvier & Valenciennes

1837), Sulawesi Utara (Haryono et al. 2002) and Buton

(Tweedley et al. 2013).

Sicyopterus longifilis de Beaufort, 1912

Anadromous; enter brackish and freshwaters (Allen

1991, Keith & Eord 2011); distributed throughout Seram,

Sulawesi, Sumatra and the Philippines (Koumans 1953);

records from Sulawesi Utara (Haryono et al. 2002), Su-

lawesi Selatan (3°27.242’S 119°32.357’E, ZFMK 69563-

69573, 69575-69577; 3°30.822’S 119°32.267’E, ZFMK

69604-69609, 69618-69619), Sulawesi Barat(2°39.08 US

119°12.436’E, ZFMK 69632-69641; 2°38.428’S 119

09.294’E, ZFMK 69667-69673; 2°37.915’S

119°09.488’E, ZFMK 69677-69678; 2°37.368’S

119°08.784’E, ZFMK 69689-69696; 3°16.65LS

118°5L929’E, ZFMK 69816-69821, 69837).

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An annotated checklist of the inland fishes of Sulawesi

Sicyopterus macrostetholepis (Bleeker, 1853)

Sicydium macrostetholepis Bleeker, 1853e

Sicydium gymnauchen Bleeker, 1858a

Anadromous; enter brackish and freshwaters; distributed

throughout Indonesia and the Philippines ( Allen 1991,

2011, Keith & Lord 2011); record from Manado, Sulawe-

si Utara (Bleeker 1858a), Buton and Kabaena (Tweedley

et al. 2013), Sulawesi Utara (Haryono et al. 2002).

Sicyopterus microcephalus (Bleeker, 1855)

Sicydium microcephalus Bleeker, 1855b

Anadromous; enter brackish and freshwaters; distributed

throughout Asia (Allen 1991, Keith & Lord 2011); record

from Buton (Tweedley et al. 2013).

Sicyopterus micrurus (Bleeker, 1854)

Sicydium micrurus Bleeker, 1854a

Anadromous; enter brackish and freshwaters; distributed

throughout Asia (Allen 1991, Keith & Lord 2011); records

from Buton and Kabaena (Tweedley et al. 2013).

Sicyopus zosterophorus (Bleeker, 1856)

Sicydium zosterophorum Bleeker, 1856b

Anadromous; enter brackish and freshwaters; distributed

throughout Asia (Allen 1991); record from Sulawesi Barat

(3°16.65LS 118°5L929’E, ZFMK 69810-69813, 69835-

69836).

Stenogobius ophthalmoporus (Bleeker, 1854)

Gobius ophthalmoporus Bleeker, 1 854a

Chonophorus lachrymosus Weber, 1 894a

Anadromous; enter brackish and freshwaters; distributed

throughout Asia; records from Sulawesi Selatan (Watson

1991, Weber 1894a) and Buton (Tweedley et al. 2013).

Stiphodon elegans (Steindachner, 1879)

Anadromous; enter brackish and freshwaters; distributed

throughout the Indo-Pacific (Watson 1995); records from

Buton and Kabaena (Tweedley et al. 2013), Sulawesi Ten-

gah(02°56.035’S 121°06.855’E, ZFMK 066027-066028),

Sulawesi Selatan (3°27.242’S 119°32.357’E, ZFMK

69578-69584; 3°30.822’S 119°32.267’E, ZFMK 69620-

69622), Sulawesi Barat (2°39.08LS 119°12.436’E,

ZFMK 69648-69649; 2°38.428'S 119°09.294’E, ZFMK

69667-69669; 2°37.915’S 119°09.488’E, ZFMK 69702-

69704; 3° 1 6.65 1 ’ S 1 1 8°5 1 .929’E, ZFMK 69832-69834);

specimens recorded outside the Society, Tubuai and Samoa

Islands are considered as closely related to S. elegans (Kot-

telat 2013).

Stiphodon semoni Weber, 1895

Anadromous; enter brackish and freshwaters; distributed

throughout the Indo-Pacific (Watson 1996); records from

Buton (Tweedley et al. 2013), Sulawesi Utara; (Haryono

et al. 2002).

Yongeichthys nebulosus (Forskal, 1775)

Gobius nebulosusYoxskkX, 111 5

Acentrogobius nebulosus (Forsskal, 1775)

Potential: Anadromous; enter brackish and freshwaters; no

actual records for Sulawesi; distributed throughout the In-

do-Pacific (Randall et al. 1990).

ACANTHUROIDEI

Scatophagidae

Scats: Euryhaline; enter brackish waters; distributed

throughout the Indo-West Pacific (Berra 2001, Eschmey-

er 2015, Froese & Pauly 2014, Nelson 2006).

Scatophagus argus (Linnaeus, 1766)

Chaetodon argus Einnaeus, 1766

Euryhaline; enter brackish waters; distributed throughout

the Indo-Pacific (Allen 1984); record from Buton (Tweed-

ley et al. 2013).

SCOMBROIDEI

Sphyraenidae

Barracudas: Marine; enter brackish waters; distributed in

all tropical and subtropical parts of the Atlantic, Indian and

Pacific Ocean (Eschmeyer 2015, Froese & Pauly 2014,

Nelson 2006).

Sphyraena barracuda (Edwards, in Catesby, 1771)

Esox barracuda Edwards, in Catesby, 1771

Euryhaline; juveniles enter brackish waters (Kottelat 2013;

Senou 2001); record from Sulawesi Selatan (4°07.456’S

1 19’37. 196’E, ZFMK 69758); original description is a re-

jected work and not available as a source, author of the

species description follows Kottelat (2013).

Sphyraena obtusata Cuvier, in Cuvier & Valenciennes,

1829

Potential: Euryhaline; enter brackish waters; no actual

records for Sulawesi; distributed throughout the Indo-Pa-

cific (Senou 2001).

ANABANTOIDEI

Anabantidae

Climbing gouramies: Freshwater; enter brackish waters;

distributed throughout the Indo-West Pacific (Berra 2001 ,

Eschmeyer 2015, Froese & Pauly 2014, Nelson 2006).

Anabas testudineus (Bloch, 1792)

Anthias testudineus Bloch, 1792

Anabas variegatus Bleeker, 1851a

Introduced: Freshwater; enter brackish waters; native to

India, South and Southeast Asia; record from Kema, Su-

lawesi Utara (Bleeker 1851a), Malili Eakes system (Na-

Bonn zoological Bulletin 64 (2): 77-106

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Friedrich Wilhelm Miesen et al.

Table 1. Summary of species and records according to i) region and ii) salt tolerance / ecology. Total numbers of records and

species are highlighted, numbers of introduced and endemic species refer to these total numbers. Species with no actual record for

Sulawesi are listed as "potential". Note that obligate and primary freshwater fishes are combined as freshwater fishes. The islands

of Buton and Kabaena, off Sulawesi Tenggara, are treated as individual regions (Tweedley et al. 2013).

sution & Aisyah 2013), Sulawesi Selatan, Lake Poso, Su-

lawesi Tengah (Kottelat 1990b, confirmed in 2012 by

F.H.).

Osphronemidae

Gouramies: Freshwater; distributed throughout Sumatra,

Java and Borneo; able to breath atmospheric air using a

specialised respiratory organ (Berra 2001, Kottelat et al.

1993, Nelson 2006).

Trichopodus (Trichogaster) pectoralis Regan, 1910

Introduced: Freshwater; native to Thailand; record from

the Malili Lakes system, Sulawesi Selatan (Herder et al.

2012a, Kottelat et al. 1993, ).

Trichopodus (Trichogaster) trichopterus Pallas, 1770

Introduced: Freshwater; native to Sundaland and Indochi-

na; record from the Malili Lakes system, Sulawesi Sela-

tan (Herder et al. 2012a, Kottelat et al. 1993), Lake Poso,

Sulawesi Tengah (Kottelat 1990b, visual record F.H.).

Channidae

Snakeheads: Freshwater; native to tropical Africa and

Southern Asia (Berra 2001; Nelson 2006).

Channa lucius (Cuvier, in Cuvier and Valenciennes,

1831)

Ophicephalus lucius Cuvier, in Cuvier and Valenciennes,

1831

Introduced: Freshwater; native to Sundaland and Indochi-

na; record from the Malili Lakes system, Sulawesi Sela-

tan (Kottelat et al. 1993).

Channa striata (Bloch, 1793)

Ophicephalus striatus Bloch, 1793

Introduced: Freshwater; native to India, China and South-

east Asia; records from the Malili Lake system, Sulawe-

si Selatan (Hadiaty & Wirjoatmodjo 2002, Hadiaty et al.

2004, Herder et al. 2012a), Lake Poso, Sulawesi Tengah

(3°4L589’S 119°38.629’E, ZFMK 69518; 3°30.822’S

119°32.267’E, ZFMK 69625) and Sulawesi Barat

(2°38.428’S 119°09.294’E, ZFMK 69671-69672).

PLEURONECTIFORMES

Paralichthyidae

Sand flounders: Euryhaline, enter brackish and freshwa-

ters; distributed throughout the Atlantic and Indo-Pacific

(Froese & Pauly 2014, Eschmeyer 2015, Nelson 2006).

Pseudorhombus malayanus Bleeker, 1865

Euryhaline; enter brackish waters (Amaoka & Hensley

2001), record from Makassar, Sulawesi Selatan, (Bleek-

er 1865a).

Pseudorhombus neglectus Bleeker, 1865

Euryhaline; enter brackish waters (Amaoka & Hensley

2001); record from Makassar, Sulawesi Selatan (Bleek-

er, 1865a).

Soleidae

Soles: Euryhaline; enter brackish and freshwaters; distrib-

uted throughout tropical and temperate regions; usually

flat, bottom dwelling fishes (Froese & Pauly 2014, Nel-

son 2006).

Bonn zoological Bulletin 64 (2); 77-106

©ZFMK

An annotated checklist of the inland fishes of Sulawesi

Achirus poropterus (Bleeker, 1851)

Euryhaline; enter brackish and freshwaters (Bleeker

1851c); record from Sulawesi Selatan (4°07.456’S

119’37.196’E, ZFMK 69766-69767).

TETRAODONTIFORMES

Triacanthidae

Triplespines: Euryhaline; enter brackish waters; distrib-

uted throughout the Indo-Pacific (Nelson 2006, Santini &

Tyler 2002).

Triacanthus biaculeatus (Bloch, 1786)

Balistes biaculeatus Bloch, 1786

Triacanthus russellii Bleeker, 185 Id

Euryhaline; enter brackish waters (Matsuura 2001);

record from Makassar, Sulawesi Selatan (Bleeker 1851d).

Tetraodontidae

Puffers: Euryhaline; enter brackish and freshwaters; dis-

tributed throughout all tropical and subtropical parts of the

Atlantic and Indo-Pacific (Nelson 2006, Yamanoue et al.

2011 ).

Arothron manilensis (Marion de Proce, 1822)

Tetrodon Manilensis Marion de Proce, 1 822

Euryhaline; enter brackish waters; distributed throughout

the Indo-Pacific (Randall 1985); record from Sulawesi Se-

latan (04°07.540’S 119°37.295’E, ZFMK 066046-

066052).

Chelonodontops patoca (Hamilton, 1822)

Tetrodon patoca Hamilton, 1 822

Chelonodon patoca (Hamilton, 1 822)

Euryhaline; enter brackish waters; distributed throughout

the Indo-Pacific (Talwar & Jhingran 1991); record from

Sulawesi Selatan (04°07.540’S 119°37.295’E, ZFMK

060060).

Dichotomy ctere erythrotaenia (Bleeker, 1853)

Tetraodon erythrotaenia Bleeker, 1853e

Euryhaline; enter brackish and freshwaters; distributed

throughout the Indo-West Pacific (Allen, 1991); record

from Maros, Sulawesi Selatan (Bleeker 1853e).

DISCUSSION

Actual records and likely occurrences of fishes in inland

waters of Sulawesi sum up to a total of 226 species (see

Table 1 for details). This ichthyofauna is composed of 1 12

genera and 56 families, dominated by Gobiidae (41

species, 18%), Adrianichthyidae (20 species, 9%), Tel-

matherinidae (19 species, 8%), and Zenarchopteridae (17

species, 7%). Taken together, these four families account

for 43% of the island’s total inland fish species diversity.

Sulawesi's native inland ichthyofauna is heterogeneous

in terms of salt tolerance: Only 89 species (44% of all na-

tive species) are obligate freshwater fishes, whereas 77

species (38% of all native species) are euryhaline. 60

species (29% of all native species) are amphi-, ana- or

catadromous, migrating between marine and freshwater

environments. 65 species (32% of all native species) of

the species inventory are endemic. 46 (71% of all endem-

ic species) of these endemic species are from radiations

in the ancient lakes of Sulawesi while only 1 8 riverine in-

cluding three euryhaline species are considered endemic.

In addition, endemism also appears to be unevenly dis-

tributed among the families. Telmatherinidae (19 species),

Adrianichthyidae (17 species) and Zenarchopteridae (17

species) contain in sum 86% of all endemic Sulawesi fish-

es.

In 2011, Parenti reported a total of 76 native freshwa-

ter fish species from Sulawesi, of which 56 were consid-

ered endemic. This significantly higher number reported

here arises to a smaller proportion from additional, recent

species descriptions (e.g. Hoese et al. 2015, Huylebrouck

et al. 2012, 2014, Earson et al. 2014, Mokodongan et al.

2014, Parenti et al. 2013). However, it is largely due to

the wider focus of the present list, which includes all fish

species recorded from Sulawesi's inland waters, also wide-

spread fish species that are very likely to be expected in

the island’s inland waters, but without actual records.

Sulawesi’s freshwater and coastal habitats are facing

substantial and manifold threats from habitat degradation

(e.g. urbanization, damming, surface mining), and stock-

ing with alien fish species (African cichlids, Asian carps,

snakeheads, or gouramis, and others - see Kottelat et al.

1993, Herder, et al. 2012, Tweedley et al. 2013). The list

presented here includes 22 introduced species, some of

which have been recognized as potential threats to the in-

digenous fauna of Sulawesi's ancient lakes (see Herder

et al. 2012a for alien fish species recorded in the Malili

Fakes area).

The total number of fish species of Sulawesi’s freshwa-

ter and brackish habitats is significantly lower than that

of the Sundaic islands, like closeby Borneo with its at least

430 fish reported species (Kottelat et al. 1993, McGinley

& Hogan 2003, Tan 2006; but note that no actual check-

list of the inland fishes is available). However, the actu-

al size of Sulawesi is four times smaller than that of Bor-

Bonn zoological Bulletin 64 (2): 77-106

©ZFMK

Friedrich Wilhelm Miesen et al.

neo (Rachman et al. 201 5). With values between 37% (160

species, McGinley & Hogan 2003) to 62% (267 species,

Tan 2006) Bomeos rate of endemism is exceptionally high,

however the four times smaller island of Sulawesi with

its endemism rate of 32% (65 species), appears surpris-

ingly close to its larger neighbour.

The actual state of exploration of the inland ichthyofau-

na of Sulawesi shows clear regional sampling biases.

However, it appears clear that the total species account

of fishes occurring in Sulawesi’s inland waters is strong-

ly dominated by the lake species flocks (see also Tweed-

ley et al. 2013, Parent! 2011, Herder & Schliewen 2010),

but the exploration of the riverine fish species diversity,

and its distribution across the island, remains in a gener-

ally fragmentary stage.

Acknowledgements. We thank the Indonesian Institute of Sci-

ences (LIPI), and the State Ministry of Research and Technol-

ogy (RISTEK) for the permit to conduct research in Indonesia.

Fieldwork benefited from help by B. Stelbrink, J. Pfaender, S.

Chapuis and M. Milanovic. F.H. thanks A.W. Nolte for fruitful

ricefish discussions and joint fieldwork. We acknowledge T. von

Rintelen for providing access to digitized maps. Fieldwork was

funded by a research grant of the DFG (to F. Herder; DFG HE

5707/2-1), and by the ARCBC 2001-2003 project to S. Wirjoat-

modjo.

REFERENCES

Abendanon EC (1915a) Midden-Celebes-Expeditie. Geologis-

che en geographische Doorkruising van Midden-Celebes

(1909-1910), Atlas. Brill, Eeiden

Abendanon EC (1915b) Midden-Celebes-Expeditie. Geologis-

che en geographische Doorkruising van Midden-Celebes

(1909-1910) 2. Brill, Eeiden

Adams AJ, Horodysky AZ, McBride RS, Guindon K, Shenker

J, MacDonald TC, Harwell HD, Ward R, Carpenter K (2013)

Global conservation status and research needs for tarpons

(Megalopidae), ladyfishes (Elopidae) and bonefishes (Albu-

lidae). Fish and Fisheries 15: 280-311. DOE

10.1111/faf 12017

Ahl E (1936) Beschreibung eines neuen Fisches der Familie

Atherinidae aus Celebes. Zoologischer Anzeiger 114: 175-177

Allen GR (1984) Scatophagidae. In; Fischer W & Bianchi G

(eds) FAO species identification sheets for fishery purposes.

Western Indian Ocean (Fishing Area 51). Volume 4. FAO,

Rome

Allen GR (1991) Field guide to the freshwater fishes of New

Guinea. Publication 9. Christensen Research Institute,

Madang, Papua New Guinea

Allen GR (1997) Lentipes watsoni, a new species of freshwater

goby (Gobiidae) from Papua New Guinea. Ichthyological Ex-

plorations of Freshwaters 8: 33-40

Allen GR (2001) TOXOTIDAE, Archerfishes. Pp. 3212-3215

in: Carpenter KE & Niem VH (eds.) FAO species identifica-

tion guide for fishery purposes. The living marine resources

of the Western Central Pacific. Volume 5. Bony fishes part 3

(Menidae to Pomacentridae). FAO, Rome

Allen GR, Midgley SH, Allen M (2002) Field guide to the fresh-

water fishes of Australia. Western Australian Museum, Perth,

Western Australia

Bonn zoological Bulletin 64 (2); 77-106

Allen GR, Steene RC (1988) Fishes of Christmas Island Indian

Ocean. Christmas Island Natural History Association, Christ-

mas Island, Indian Ocean, 6798, Australia

Alleyne HG, Macleay W (1877) The ichthyology of the Chev-

ert expedition. Proceedings of the Einnean Society of New

South Wales 1: 261-281, 321-359

Amaoka K, Hensley DA (2001) PARAEYCHTHYIDAE Sand

flounders. Pp.3842-3862 in; Carpenter KE & Niem VH (eds)

FAO species identification guide for fishery purposes. The liv-

ing marine resources of the Western Central Pacific. Volume

6. Bony fishes part 4 (Eabridae to Eatimeriidae), estuarine

crocodiles, sea turtles, sea snakes and marine mammals. FAO,

Rome

Anderson WD, Allen GR (1999) Eutjanidae, Snappers (jobfish-

es). Pp. 2840-2918 in; Carpenter KE & Niem VH (eds) FAO

species identification guide for fishery purposes. The living

marine resources of the Western Central Pacific. Volume 5.

Bony fishes part 3 (Menidae to Pomacentridae). FAO, Rome

Anderson WD, Collette BB (1991) Revision of the freshwater

halfbeaks of the genus Hemirhamphodon (Teleostei, Hemi-

ramphidae). Ichthyological Exploration of Freshwaters 2;

151-176

Anderson ME, Heemstra PC (2003) Review of the glassfishes

(Perciformes: Ambassidae) of the Western Indian ocean. Cy-

bium27: 199-209

Aral T, Eimbong D, Otake T, Tsukamoto K (1999) Metamorpho-

sis and inshore migration of tropical eels Anguilla spp. in the

Indo-Pacific. Marine Ecology Progress Series 182; 283-293

Aral T, Miller MJ, Tsukamoto K (2003) Earval duration of the

tropical eel Anguilla celebesensis from Indonesia and the

Philippine coasts. Marine Ecology Progress Series 251 : 255-

261

Aurich H (1935) Mitteilung der Wallacea-Expedition Woltereck.

Mitteilung XIII. Fische I. Zoologischer Anzeiger 112: 97-107

Aurich H (1938) Die Gobiiden (Ordnung; Gobioidea). Mittei-

lung XXVIII der Wallacea-Expedition Woltereck. Interna-

tionale Revue der gesamten Hydrobiologie und Hydrographic,

Eeipzig 38: 125-183

Baird SF, Girard CF (1853) Descriptions of new species of fish-

es collected by Mr. John H. Clark, on the U. S. and Mexican

Boundary Survey, under Et. Col. Jas. D. Graham. Proceedings

of the Academy of Natural Sciences of Philadelphia 6:

387-390

Beaufort EF de (1912) On some new Gobiidae from Ceram and

Waigen. Zoologischer Anzeiger 39: 136-143

Bellwood DR (2001) SCARIDAE, Parrotfishes. Pp. 3468-3492

in: Carpenter KE & Niem VH (eds) FAO species identifica-

tion guide for fishery purposes. The living marine resources

of the Western Central Pacific. Volume 6. Bony fishes part 4

(Eabridae to Eatimeriidae), estuarine crocodiles, sea turtles,

sea snakes and marine mammals. FAO, Rome

Berra TM (2001) Freshwater fish distribution. Academic Press,

San Diego

Blaber SJM, Milton DA (1990) Species composition, commu-

nity structure and zoogeography of fishes of mangrove estu-

aries in the Solomon Islands. Marine Biology 105: 259-267

Bleeker P (1 849a) Bijdrage tot de kermis der ichthyologische fau-

na van Midden- en Oost-Java, met beschrijving van eenige

nieuwe species. Verhandelingen van het Bataviaasch Genoot-

schap van Kunsten en Wetenschappen 23 (12); 1-23

Bleeker P (1849b) Bijdrage tot de kermis der ichthyologische fau-

na van het eiland Bali, met beschrijving van eenige nieuwe

species. Verhandelingen van het Bataviaasch Genootschap van

Kunsten en Wetenschappen 22 (7); 1-11

©ZFMK

An annotated checklist of the inland fishes of Sulawesi

Bleeker P (1849c) Bijdrage tot de kermis der Blennioiden en Go-

bioiden van den Soenda-Molukschen archipel, met be-

schrijving van 42 nieuwe species. Verhandelingen van het Ba-

taviaasch Genootschap van Kunsten en Wetenschappen 22 (6):

1-40

Bleeker P (1850) Bijdrage tot de kermis der ichthyologische fau-

na van Borneo, met beschrijving van 16 nieuwe soorten van

zoetwatervisschen. Natuurkundig Tijdschrift voor Neder-

landsch Indie 1; 1-16

Bleeker P (1851a) Nieuwe bijdrage tot de kennis der ichthyo-

logische fauna van Celebes. Natuurkundig Tijdschrift voor Ne-

derlandsch Indie 2: 209-224

Bleeker P (1 85 lb) Over eenige nieuwe soorten van Blennioiden

en Gobioiden van den indischen archipel. Natuurkundig Tijd-

schrift voor Nederlandsch Indie 1: 236-258

Bleeker P (1851c) Bijdrage tot de kennis der Pleuronecteoiden

van den Soenda-Molukschen archipel. Verhandelingen van het

Bataviaasch Genootschap van Kunsten en Wetenschappen 24

(9): 1-28

Bleeker P (185 Id) Bijdrage tot de kennis der Balistini en Ostra-

ciones van den indischen archipel. Verhandelingen van het Ba-

taviaasch Genootschap van Kunsten en Wetenschappen 24

(11): 1-36

Bleeker P (1852a) Bijdrage tot de kennis der clupeoiden van den

Soenda-Molukschen archipel [title on cover; Bijdrage tot de

kennis der haringachtige visschen van den Soenda-Moluk-

schen archipel. Verhandelingen van het Bataviaasch Genoot-

schap van Kunsten en Wetenschappen 24; 1-48

Bleeker P (1852b) Bijdrage tot de kennis der Muraenoiden en

Symbranchoiden van den indischen archipel. Verhandelingen

van het Bataviaasch Genootschap van Kunsten en Weten-

schappen 25: 1-62

Bleeker P (1852c) Bijdrage tot de kennis der ichthijologische fau-

na van de Moluksche eilanden. Visschen van Amboina en Ce-

ram. Natuurkundig Tijdschrift voor Nederlandsch Indie 3:

229-309

Bleeker P (1852d) Bijdrage tot de kermis der Plagiostomen van

den Indischen Archipel. Verhandelingen van het Bataviaasch

Genootschap van Kunsten en Wetenschappen 24 (12): 1-92

Bleeker P (1852e) Zesde bijdrage tot de kennis der ichthyolo-

gische fauna van Borneo. Visschen van Pamangkat, Bandjer-

massing, Praboekarta en Sampit. Natuurkundig Tijdschrift

voor Nederlandsch Indie 3: 407-442

Bleeker P (1853a) Derde bijdrage tot de kennis der ichthijolo-

gische fauna van Celebes. Natuurkundig Tijdschrift voor Ne-

derlandsch Indie 3: 739-782

Bleeker P (1853b) Nieuwe bijdrage tot de kennis der ichthijo-

logische fauna van Ceram. Natuurkundig Tijdschrift voor Ne-

derlandsch Indie 3: 689-714

Bleeker P (1853c) Diagnostische beschrijvingen van nieuwe of

weinig bekende vischsoorten van Sumatra. Tiental V-X. Na-

tuurkundig Tijdschrift voor Nederlandsch Indie 4; 243-302

Bleeker P (1853d) Diagnostische beschrijvingen van nieuwe of

weinig bekende vischsoorten van Batavia. Natuurkundig Tijd-

schrift voor Nederlandsch Indie 4: 451-516

Bleeker P (1853e) Vierde bijdrage tot de kennis der ichthyolo-

gische fauna van Celebes. Natuurkundig Tijdschrift voor Ne-

derlandsch Indie 5: 153-174

Bleeker P (18531) Aanhangsel op de bijdrage tot de kennis der

Muraenoiden en Symbranchoiden van den indischen archipel.

Verhandelingen van het Bataviaasch Genootschap van Kuns-

ten en Wetenschappen 25 (5): 63-76

Bleeker P (1854a) Vierde bijdrage tot de kennis der ichthyolo-

gische fauna van Amboina. Natuurkundig Tijdschrift voor Ne-

derlandsch Indie 5: 317-352

Bonn zoological Bulletin 64 (2): 77-106

Bleeker P (1854b) Ichthyologische waarnemingen, gedaan op

verschillende reizen in de Residentie Banten. Natuurkundig

Tijdschrift voor Nederlandsch Indie 7; 309-326

Bleeker P (1854c) Bijdrage tot de kennis der troskieuwige viss-

chen van den indischen archipel. Verhandelingen van het Ba-

taviaasch Genootschap van Kunsten en Wetenschappen 25 (6):

1-30

Bleeker P (1854d) Vijfde bijdrage tot de kennis der ichthyolo-

gische fauna van Celebes. Natuurkundig Tijdschrift voor Ne-

derlandsch Indie 7: 225-260

Bleeker P (1855a) Zevende bijdrage tot de kennis der ichthyo-

logische fauna van Celebes. Natuurkundig Tijdschrift voor Ne-

derlandsch Indie 8: 435-444

Bleeker P (1855b) Specierum piscium Javanensium novarum vel

minus cognitarum diagnoses adumbratae. Natuurkundig Tijd-

schrift voor Nederlandsch Indie 7; 415-448

Bleeker P (1856a) Beschrijvingen van nieuwe en weinig beken-

de vischsoorten van Manado en Makassar, grootendeels ver-

zameld op eene reis door den Molukschen archipel, gedaan

in het gevolg van den Gouverneur Generaal Duymaer van

Twist, in September en Oktober 1855. Acta Societatis Scien-

tiarum Indo-Neerlandicae 1 (6): 1-80

Bleeker P (1856b) Nieuwe bijdrage tot de kennis der ichthyo-

logische fauna van Bali. Natuurkundig Tijdschrift voor Ne-

derlandsch Indie 12; 291-302

Bleeker P (1858a) Tiende bijdrage tot de kennis der vischfauna

van Celebes. Acta Societatis Scientiarum Indo-Neerlandicae

3: 1-16

Bleeker P (1858b) Twaalfde bijdrage tot de kennis der vischfau-

na van Celebes. Visschen van Manado. Acta Societatis Sci-

entiarum Indo-Neerlandicae 5; 1-4

Bleeker P (1860) Dertiente bijdrage tot de kennis der vischfau-

na van Celebes (visschen van Bonthain, Badjoa, Sindjai, La-

goesi en Pompenoea). Acta Societatis Regiae Scientiarum In-

do-Neerlandicae 8 (7); 1-60

Bleeker P (1864) Poissons inedits indo-archipelagiques de Tor-

dre des murenes. Nederlandsch Tijdschrift voor de Dierkun-

de 2: 38-54

Bleeker P (1865a) Sur la pluralite des especes indoarchipelagi-

ques du genre Megalops Lac. Nederlandsch Tijdschrift voor

de Dierkunde; 278-292

Bleeker P (1865b-1875) Atlas ichthyologique des Indes Orien-

tales Neerlandaises. Tome VI. Pleuronectes, scombresoces,

clupees, clupesoces, chauliodontes, saurides. Muller, Amster-

dam

Bleeker P (1872) See Bleeker (1865b)

Bleeker P (1873a) Revision des especes indo-archipelagiques des

genres Lutjanus et Aprion. Verhandelingen der Koninklijke

Akademie van Wetenschappen te Amsterdam 13; 1-102

Bleeker P (1873b) Revision des especes insulindiennes du gen-

re Therapon. Nederlandsch Tijdschrift voor de Dierkunde 4:

372-393

Bloch ME (1786) Naturgeschichte der auslandischen Fische.

Berlin, v. 2

Bloch ME (1792) Naturgeschichte der auslandischen Fische.

Berlin, v. 6

Bloch ME (1793) Naturgeschichte der auslandischen Fische.

Berlin, v. 7

Borsa P, Durand JD, Shen KN, Arlyza IS, Solihen DD, Berrebi

P (2013) Himantura tutuJ sp. nov. (Myliobatoidei; Dasyati-

dae), a new ocellated whipray from the tropical Indo-West Pa-

cific, described from its cytochrome-oxidase I gene sequence.

Comptes Rendus Biologies 336: 82-92

Boulenger GA (1897) An account of the freshwater fishes col-

lected in Celebes by Drs. P. & F. Sarasin. Proceedings of the

©ZFMK

100

Friedrich Wilhelm Miesen et al.

General Meetings for Scientific Business of the Zoological So-

ciety of London 1897; 426-429

Bowers NJ, Stauffer JR Jr (1997) Eight new species of rock-

dwelling cichlids of the genus Melanochromis (Teleostei; Ci-

chlidae) from Lake Malawi, Africa. Ichthyological Explora-

tion of Freshwaters 8: 49-70

Brembach M (1982) Drei neue Dermogenys-AxtQn aus Sulawe-

si; D. montamis, D. vogti, D. megarrhamphus. Die Aquarien-

und Terrarien-Zeitschrift 35; 51-55

Brembach M (1991) Lebendgebarende Halbschnabler. Untersu-

chungen zur Verbreitung, Morphologic, Systematik und Fort-

pflanzungsbiologie der lebendgebarenden Halbschnabler der

Gattungen Dermogenys und Nomorhamphus (Hemirhamphi-

dae; Pisces). Verlag Natur und Wissenschaft, Solingen

Brooks JL (1950) Speciation in ancient lakes. Quarterly Review

of Biology 25, 30-60; 131-76

Broussonet PMA (1782) Ichthyologia sistens piscium descrip-

tiones et icones. Elmsly, London

Carpenter KE (2001a) LETHRINIDAE, Emperors (emperor

snappers). Pp. 3004-3050 in; Carpenter KE & Niem VH (eds)

FAO species identification guide for fishery purposes. The liv-

ing marine resources of the Western Central Pacific. Volume

5. Bony fishes part 3 (Menidae to Pomacentridae). FAO, Rome

Carpenter KE (2001b) KUHLIIDAE, Flagtails. Pp. 3317-3320

in; Carpenter KE & Niem VH (eds) FAO species identifica-

tion guide for fishery purposes. The living marine resources

of the Western Central Pacific. Volume 5. Bony fishes part 3

(Menidae to Pomacentridae). FAO, Rome

Castle PHJ (1968) A contribution to a revision of the moringuid

eels. Special Publication, Department of Ichthyology, Rhodes

University, Grahamstown 3; 1-29

Cepede E La (1802) Histoire naturelle des poissons. Tome qua-

trieme. Plassan, Paris

Cerwenka AF, Wedekind JD, Hadiaty RK, Schliewen UK,

Herder F (2012) Alternative egg-feeding tactics in Telmath-

erina sarasinorum, atrophic specialist of Lake Matano’s evolv-

ing sailfin silversides fish radiation. Hydrobiologia 693;

131-139

Coates D (1987) Observations on the biology of tarpon. Mega-

lops cyprinoides (Broussonet) (Pisces; Megalopidae), in the

Sepik River, northern Papua New Guinea. Australian Journal

of Marine & Freshwater Research 38; 529-535

Collette B (1995) Tondanichthys kotteJati, a new genus and

species of freshwater halfbeak (Teleostei; Hemiramphidae)

from Sulawesi. Ichthyological Exploration of Freshwaters 6

(2); 171-174

Collette BB (2004) Annotated checklist of fishes, family Hemi-

ramphidae Gill 1859. California Academy of Sciences 22;

1-35

Compagno LJV (1984) FAO Species Catalogue. Vol. 4. Sharks

of the world. An annotated and illustrated catalogue of shark

species known to date. Part 2 - Carcharhiniformes. FAO Fish-

eries Synopsis 125; 251-655

Compagno LJV, Ebert DA, Smale MJ (1989) Guide to the sharks

and rays of southern Africa. New Holland Ltd., London

Compagno LJV, Roberts TR (1982) Freshwater stingrays

(Dasyatidae) of Southeast Asia and New Guinea, with descrip-

tion of a new species of Himantura and reports of unidenti-

fied species. Environmental Biology of Fishes 7; 321-339

Cosker JE, Ide S, Endo H (2012) Three new species of

Ophichthid Eels (Anguilliformes; Ophichthidae) from Japan.

Bulletin of the National Museum of Nature and Science. Se-

ries A, Supplement 6; 1-16

Cuvier G (1816) Le regne animal distribue d’apres son organi-

sation, pour servir de base a Fhistoire naturelle des animaux

Bonn zoological Bulletin 64 (2); 77-106

et d’ introduction a Fanatomie comparee. Tome IV, contenant

les zoophytes, les tables, et les planches. Deterville, Paris

Cuvier G (1829) Le Regne Animal, distribue d’apres son organ-

isation, pour servir de base a Fhistoire naturelle des animaux

et d’ introduction a Fanatomie comparee. Edition 2. Fortin,

Masson et Cie, Paris

Cuvier G., Valenciennes A (1828) Histoire naturelle des pois-

sons. Tome second. Livre Troisieme. Des poissons de la

famille des perches, ou des percoides 2. Imprimeur-Libraire,

Paris

Cuvier G, Valenciennes A (1829) Histoire naturelle des poissons.

Tome troisieme. Levrault, Paris

Cuvier G, Valenciennes A (1830) Histoire naturelle des poissons.

Tome Sixieme. Livre sixieme. Partie I. Des Sparoides; Partie

IT Des Menides 6. Levrault, Paris

Cuvier G, Valenciennes A (1831) Histoire naturelle des poissons.

Tome septieme. Levrault, Paris

Cuvier G, Valenciennes A (1833) A Histoire naturelle des pois-

sons. Tome neuvieme. Suite du livre neuvieme. Des Scom-

beroides 9 Levrault, Paris

Cuvier G, Valenciennes A (1836) Histoire naturelle des poissons.

Tome onzieme. Levrault, Paris

Cuvier G, Valenciennes A (1837) Histoire naturelle des poissons.

Tome douzieme. Levrault, Paris

Cuvier G, Valenciennes A (1842) Histoire naturelle des poissons.

Tome seizieme. Livre dix-huitieme. Les Cyprinoides 16. P.

Bertrand, Paris

Cuvier G, Valenciennes A (1847) Histoire naturelle des poissons.

Tome dix-neuvieme. Suite du livre dix-neuvieme. Brochets ou

Lucioides. Livre vingtieme. De quelques families de Malacop-

terygiens, intermediaires entre les Brochets et les Clupes. P.

Bertrand, Paris

Daverat F, Morais P, Dias E, Babaluk J, Martin J, Eon M, Fa-

blet R, Peycheran C, Antunes C (2012) Plasticity of European

flounder life history patterns discloses alternatives to

catadromy. Marine Ecology Progress Series 465; 267-280

Dawson CE (1984) Revision of the Genus Microphis Kaup

(Pisces; Syngnathidae). Bulletin of Marine Sciences 35;

117-181

Dawson CE (1985) Indo-Pacific pipefishes (Red Sea to the

Americas). The Gulf Coast Research Laboratory Ocean

Springs, Mississippi, USA

de Bruyn M, Rintelen T von, Rintelen K von, Mather PB, Car-

valho GR (2012) Molecular biogeography and phylogeogra-

phy of the freshwater fauna of the Indo- Australian Archipel-

ago. Pp. 316-348 in; Gower DJ (ed.) Biotic Evolution and En-

vironmental Change in Southeast Asia. Cambridge Universi-

ty Press, Cambridge

de Bruyn M, Riiber L, Nylinder S, Stelbrink B, Lovejoy NR,

Lavoue S, Tan HH, Nugroho E, Wowor D, Ng PKL, Siti Az-

izah MN, Rintelen T von. Hall R, Carvalho GR (2013) Pa-

leo-drainage basin connectivity predicts evolutionary relation-

ships across three Southeast Asian biodiversity hotspots. Sys-

tematic Biology 62; 398-410

Donaldson TJ, Myers RF (2002) Insular freshwater fish faunas

of Micronesia; patterns of species richness and similarity. En-

vironmental Biology of Fishes 65; 139-149

Duncker G. (1910) On some syngnathids (“pipe fish”) from Cey-

lon. Spolia Zeylanica 7 (25); 25-34

Durand JD, Shen KN, Chen WJ, Jamandre BW, Biel H, Diop

K, Borsa P (2012) Systematics of the grey mullets (Teleostei;

Mugiliformes; Mugilidae); molecular phylogenetic evidence

challenges two centuries of morphology-based taxonomy. Mo-

lecular Phylogenetics and Evolution 64; 73-92

©ZFMK

An annotated checklist of the inland fishes of Sulawesi

101

Ebner BC, Kroll B, Godfrey P, Thuesen PA, Vallance T, Pusey

B, Allen GR, Rayner IS, Perna N (2011) Is the elusive Gym-

nothorax polyuranodon really a freshwater moray? Journal of

Fish Biology 79: 70-79

Ege V (1939) A revision of the genus Anguilla Shaw. A system-

atic, phylogenetic and geographical study. Dana Reports:

1-256

Eschmeyer WN (ed.) (2015) Catalog of fishes: Genera, species,

references. World Wide Web electronic publication. Online at:

http://researcharchive.calacademy.org/research/ichthyology/ca

talog/fishcatmain.asp last accessed on May 26, 2015

Feltes RM (2001) POEYNEMIDAE, Threadfins. Pp. 3090-3116

in: Carpenter KE & Niem VH (eds.) FAO species identifica-

tion guide for fishery purposes. The living marine resources

of the Western Central Pacific. Volume 5. Bony fishes part 3

(Menidae to Pomacentridae). FAO, Rome

Forskal P (1775) Descriptiones animalium avium, amphibiorum,

piscium, insectorum, vermium; quae in itinere orientali obser-

vavit Petrus Forskal [...]. Post mortem auctoris edidit Carsten

Niebuhr. Adjuncta est materia medica kahirina atque tabula

Maris Rubra geographica. Moller, Hauniae

Fowler HW (1931) Contribution to the biology of the Philippine

Archipelago and adjacent regions. The fishes of the families

Pseudochromitidae, lobotidae, Pempheridae, Priacanthidae,

Eutjanidae, Pomadasyidae, and Terapontidae, collected by the

United States bureau of Fisheries steamer “Albatross”,

chiefly in Philippine seas and adjacent waters. Bulletin of the

United States National Museum 100 (11): 1-388

Fricke R, Kulbicki M, Wantiez F (2011) Checklist of the fish-

es of New Caledonia, and their distribution in the Southwest

Pacific Ocean (Pisces). Stuttgarter Beitrage zur Naturkunde

A, Neue Serie 4: 341-463

Froese R, Pauly D (2014) FishBase. World Wide Web electron-

ic publication. Online at www.fishbase.org last accessed on

May 26, 2015

Garman S (1903) Some fishes from Australasia. Bulletin of the

Museum of Comparative Zoology 39: 229-241

Girard CF (1859) Ichthyological notices. Proceedings of the

Academy of Natural Sciences of Philadelphia 11: 56-68

Gmelin, JF (1789) Carol! a Finne systema naturae per regna tria

naturae, secundum classes, ordines, genera, species cum char-

acteribus, differentiis, synonymis, locis. Beer, Fipsiae vol. 1

pars 3: 1033-2224

Gomon MF, Kuiter RH (2009) Two new pygmy seahorses

(Teleostei: Syngnathidae: Hippocampus) from the Indo-West

Pacific, aqua. International Journal of Ichthyology 15: 37-44

Grace M (2001) Field Guide to Requiem Sharks (Elasmobra-

chiomorphi: Carcharhinidae) of the Western North Atlantic.

NOAA Technical Report NMFS 153: 1-32

Gray SM, Dill FM, McKinnon JS, 2007. Cuckoldry incites can-

nibalism: male fish turn to cannibalism when perceived cer-

tainty of paternity decreases. The American Naturalist 169:

258-263

Gray SM, Dill FM, Tantu FY, Foew ER, Herder F, McKinnon

JS (2008b) Environment-contingent sexual selection in a

colour polymorphic fish. Proceedings of the Royal Society of

FondonB 275: 1785-1791

Gray SM, McKinnon JS, Tantu FY, Dill FM (2008a) Sneaky egg-

eating in Telmatherina sarasinorum, an endemic fish from Su-

lawesi. Journal of Fish Biolo^ 73: 728-731

Grier HJ, Collette BB (1987) Unique spermatozeugmata in testes

of halfbeaks of the genus Zenarchopterus (Teleostei: Hemi-

ramphidae). Copeia 1987: 300-311

Gunther A (1870) Catalogue of the fishes in the British Muse-

um. Catalogue of the Physostomi, containing the families

Bonn zoological Bulletin 64 (2): 77-106

Gymnotidae, Symbranchidae, Muraenidae, Pegasidae, and of

the Fophobranchii, Plectognathi, Dipnoi, ...[thru] ... Fepto-

cardii, in the British Museum. Catologue of Fishes 8: 1-512

Gunther A (1 872a) Notice of two new fishes from Celebes. An-

nals and Magazine of Natural History 4: 438-440

Gunther A (1872b) Report on several collections of fishes re-

cently obtained for the British Museum. Proceedings of the

Scientific Meetings of the Zoological Society of Fondon 1871:

652-675

Hadiaty RK (2007) Kajian ilmiah ikan pelangi, Marosatherina

ladigesi (Ahl 1936), Fauna endemik Sulawesi. Berita Biolo-

gi 8: 473-479

Hadiaty RK, Wirjoatmodjo S (2002) Studi Pendahuluan: Bio-

diversitas dan Distribusi Ikan Endemik di Danau Matano, Su-

lawesi Selatan. Jurnal Iktiologi Indonesia 2 (2): 23-29

Hadiaty RK, Wirjoatmodjo S, Sulistiono (2004) Koleksi ikan

di Danau Mahalona, Fantoa dan Masapi, Sulawesi Selatan. Ju-

mal Iktiologi Indonesia 4 (I): 31-42

Hadijaj S, Tuwo A, Jusoff K, MallawaA, Malina AC, Tamsil A

(2014) Genetic Diversity Population of Beloso (Glossogob-

ius aureus) in Fake Tempe, Sulawesi, Indonesia. World Jour-

nal of Fish and Marine Sciences 6 (1): 87-97

Hamilton F (1822) An account of the fishes found in the river

Ganges and its branches. Constable, Edinburgh

Harrison IJ, Senou H (1999) Order Mugiliformes. Mugilidae.

Mullets. Pp. 2069-2108 in: Carpenter KE & Niem VH (eds.)

FAO species identification guide for fishery purposes. The liv-

ing marine resources of the Western Central Pacific. Volume

4. Bony fishes part 2 (Mugilidae to Carangidae). FAO, Rome

Haryono AHT, Tjakrawidjaja D (2004) Komunitas ikan di

perairan danau wilayah Sulawesi utara dan Gorontalo. Biota:

54-62

Haryono AHT, Tjakrawidjaja D, Riyanto A (2002) Iktiofauna di

perairan sekitar gunung kabela taman nasional Bogani Nani

wartabone Sulawesi utara. Jurnal Iktiologi Indonesia: 31-40

Heemstra E, Heemstra PC, Smale MJ, Hooper T, Pelicier D

(2004) Preliminary checklist of coastal fishes from the Mau-

ritian island of Rodrigues. Journal of Natural History 38:

3315-3344

Herder F, Chapuis S (2010) Oryzias hadiatyae, a new species

of ricefish (Atherinomorpha: Beloniformes: Adrianichthyidae)

endemic to Fake Masapi, Central Sulawesi, Indonesia. Raf-

fles Bulletin of Zoology 58: 269-280

Herder F, Hadiaty R, Nolte A (2012b) Pelvic-fm brooding in a

new species of riverine ricefish (Atherinomorpha: Beloni-

formes: Adrianichthyidae) from Tana Toraja, Central Sulawe-

si, Indonesia. Raffles Bulletin of Zoology 60: 467-476

Herder F, Pfaender J, Hadiaty R, Schliewen U (2006a) Prelim-

inary checklist of sailfm silversides (Teleostei: Telmatherini-

dae) in the Malili Fakes of Sulawesi (Indonesia), with a syn-

opsis of systematics and threats. Verhandlungen der Gesell-

schaft fur Ichthyologic 5: 139-163

Herder F, Nolte AW, Pfaender J, Schwarzer J, Hadiaty RK, &

Schliewen UK (2006b) Adaptive radiation and hybridization

in Wallace’s Dreamponds: evidence from sailfin silversides

in the Malili Fakes of Sulawesi. Proceedings of the Royal So-

ciety of Fondon B: Biological Sciences 273: 2209-2217

Herder F, Pfaender J, Schliewen UK (2008) Adaptive sympatric

speciation of polychromatic “roundfin” sailfm silverside fish

in Fake Matano (Sulawesi). Evolution 62: 2178-95. doi:

10. 1 1 1 1/j. 1 558-5646.2008.00447.X

Herder F, Schliewen U (2010) Beyond sympatric speciation: Ra-

diation of sailfin silverside fishes in the Malili Fakes (Sulawe-

si). Pp. 465-483 in: Glaubrecht M (ed.). Evolution in Action -

Adaptive Radiations and the Origins of Biodiversity. Springer

©ZFMK

102

Friedrich Wilhelm Miesen et al.

Verlag, Berlin Heidelberg, doi: 10.1007/978-3-642-12425-

9_22

Herder F, Schliewen UK, Geiger MF, Hadiaty RK, Gray SM,

McKinnon JS, Walter RP, Pfaender J (2012a) Alien invasion

in Wallace’ s Dreamponds; records of the hybridogenic “flow-

erhom” cichlid in Lake Matano, with an annotated checklist

of fish species introduced to the Malili Lakes system in Su-

lawesi. Aquatic Invasions 7: 521-535

Herre AWCT (1927) Gobies of the Philippines and the China

Sea. Monographs of the Bureau of Sciences, Manila

Herre AWCT (1936) Notes on fishes in the Zoological Muse-

um of Stanford University. IV. A new catostomid from Mex-

ico and a new callionymid from Celebes and the Philippines.

Proceedings of the Biological Society of Washington 49:

11-13

Herre AWCT (1945) Notes on fishes in the Zoological Muse-

um of Stanford University. XVIII. Two new species of Taman-

ka, with a key to the species from the Philippines and China.

Proceedings of the Biological Society of Washington 58:

73-76

Herre AWCT (1953) Eight additions to the Philippine fish fau-

na, including three new species. Philippine Journal of Science

82: 9-14

Heupel MR, Simpfendorfer CA (2008) Movement and distribu-

tion of young bull sharks Carcharhinus leucas in a variable

estuarine environment. Aquatic Biology 1: 277-289

Hiroi J, McCormick SD (2012) New insights into gill ionocyte

and ion transporter function in euryhaline and diadromous fish.

Respiratory Physiology & Neurobiology 184: 257-268

Hoese DF, Hadiaty RK, Herder F (2015) Review of the dwarf

Giossogohius lacking head pores from the Malili lakes, Su-

lawesi, with a discussion of the definition of the genus. Raf-

fles Bulletin of Zoology 63: 14-26

Hoese DF, Kottelat M (2005) Bostrychus microphthalmus, a new

microphthalmic cavefish from Sulawesi (Teleostei: Gobiidae).

Ichthyological Explorations of Freshwaters 16: 183-191

Holland KN, Eowe CG, Wetherbee BM (1996) Movements and

dispersal patterns of blue trevally (Carawc melampygus) in a

fisheries conservation zone. Fisheries Research 25. 279-292

Huang SP, van Oijen MJP, Huang KY, Tsai CC, Chen IS (2013)

Redescription of Gobiiis gastrospilos Bleeker, 1853 with com-

ments on four newly recorded species of brackish gobies from

Taiwan. Journal of Marine Science and Technology Supple-

ment 21: 94-105

Huylebrouck J, Hadiaty RK, Herder F (2012) Nomorhamphus

rex, a new species of viviparous halfbeak (Atherinomorpha:

Beloniformes: Zenarchopteridae) endemic to Sulawesi Sela-

tan, Indonesia. Raffles Bulletin of Zoology 60: 477-485

Huylebrouck J, Hadiaty RK, Herder F (2014) Two new species

of viviparous halfbeaks (Atherinomorpha : Beloniformes :

Zenarchopteridae) endemic to Sulawesi Tenggara, Indonesia.

Raffles Bulletin of Zoology 62: 200-209

Jordan DS, Richardson RE (1908) Fishes from islands of the

Philippine Archipelago. Bulletin of the Bureau of Fisheries 27:

233-287

Jordan DS, Seale A (1906) The fishes of Samoa. Description of

the species found in the archipelago, with a provisional check-

list of the fishes of Oceania. Bulletin of the Bureau of Fish-

eries 25: 173-455 + 457-488

Kaup J (1856) Catalogue of lophobranchiate fish in the collec-

tion of the British Museum. British Museum, Eondon

Kaup J (1857) Catalogue of apodal fish, in the collection of the

British Museum. British Museum, Eondon

Keith P, Hadiaty R, Hubert N, Busson F, Eord C (2014) Three

new species of Lentipes from Indonesia (Gobiidae). Cybium

38: 133-146

Keith P, Hadiaty RK, Eord C (2012) A new species of Belo-

branchus (Teleostei: Gobioidei: Eleotridae) from Indonesia.

Cybium 36:479-484

Keith P, Eord C (2011) Tropical Freshwater Gobies: Am-

phidromy as a Fife Cycle. Pp. 243-278 in: Patzner R, Van Tas-

sell JE, Kovacic M & Kapoor BG (eds) (2011) The biology

of gobies. CRC Press, Boca Raton

Keith P, Marquet G, Valade P, Bose P, Vigneux E (2006) Atlas

des poissons et des crustaces d’eau douce des Comores, Mas-

careignes et Seychelles. Museum national d’Histoire naturelle,

Paris. Patrimoines naturels 65: 250 pp.

Knapp EW (1999) Platycephalidae. Flatheads. Pp. 2385-2421

in Carpenter KE & Niem VH (eds) FAO species identifica-

tion guide for fishery purposes. The living marine resources

of the Western Central Pacific. Volume 4. Bony fishes part 2

(Mugilidae to Carangidae). FAO, Rome

Kottelat M (1989a) Der Matano-See. DATZ 42: 616-618

Kottelat M (1989b) Der Towuti-See. DATZ 42: 681-684

Kottelat M (1989c) Die Susswasser-Fauna von Sulawesi.

DATZ 42: 555-558

Kottelat M (1990a) Sulawesi: Flussfische. Aquarien und Ter-

rarien Zeitschrift 43: 725-738

Kottelat M (1990b) Synopsis of the endangered buntingi (Os-

teichthyes: Adrianichthyidae and Oryziidae) of Take Poso,

Central Sulawesi, Indonesia, with a new reproductive guild

and descriptions of three new species. Ichthyological Explo-

ration of Freshwaters 1: 49-67

Kottelat M (1990c) Sailfm silversides (Pisces: Telmatherinidae)

of Eakes Towuti, Mahalona and Wawontoa (Sulawesi, Indone-

sia) with descriptions of two new genera and two new species.

Ichthyological Exploration of Freshwaters 1: 35-54

Kottelat M (I990d) The ricefishes (Oryziidae) of the Malili

Eakes, Sulawesi, Indonesia, with description of a new

species. Ichthyological Exploration of Freshwaters I: 151-166

Kottelat M (1991) Sailfin silversides (Pisces: Telmatherinidae)

of Take Matano, Sulawesi, Indonesia, with descriptions of six

new species. Ichthyological Exploration of Freshwaters 1:

321-344

Kottelat M, Freyhof J (2007) Handbook of European Freshwa-

ter Fishes. Publications Kottelat, Cornol and Freyhof, Berlin

Kottelat M, Whitten T (1996) Freshwater biodiversity in Asia:

with special reference to fish. World Bank Publications 343:

1-72

Kottelat M, Whitten T, Kartikasari SN, Wirjoatmodjo S (1993)

Freshwater fishes of western Indonesia and Sulawesi. Hong

Kong: Periplus Editions

Kottelat M (201 1) Pieter Bleeker in the Netherlands East Indies

(10 March 1842 -ca. 21 September 1860): new biographical

data and a chronology of his zoological publications. Ichthy-

ological Explorations of Freshwaters 22 (1): 11-94

Kottelat M (2013) The Fishes of the Inland Waters of Southeast

Asia: A Catalogue and Core Bibliography of the Fishes Known

to Occur in Freshwaters, mangroves and Estuaries. Raffles

Bulletin of Zoology Supplement 27: 1-663

Koumans FP (1953) Gobioidea. In: Weber M & Beaufort EF de

(eds). The Fishes of the Indo-Australian Archipelago X. Brill,

Eeiden

Eadiges W (1972) Zwei neue Hemirhamphiden von Celebes und

Cebu (Philippinen). Mitteilungen aus dem Hamburgischen Zo-

ologischen Museum und Institut 68: 207-212

Bonn zoological Bulletin 64 (2): 77-106

©ZFMK

An annotated checklist of the inland fishes of Sulawesi

103

Larson HK (2001) A revision of the gobiid fish genus Mugilo-

gobius (Teleostei; Gobioidei), and its systematic placement.

Records of the Western Australian Museum Supplement 62:

1-235

Larson HK (2010) A review of the gobiid fish genus Redigob-

ius (Teleostei: Gobionellina), with descriptions of two new

species. Ichthyological Explorations of Freshwaters 21:

123-191

Larson HK, Geiger ME, Hadiaty RK, Herder F (2014) Mugilo-

gobius hitam, a new species of freshwater goby (Teleostei: Go-

bioidei: Gobiidae) from Fake Towuti, Central Sulawesi, In-

donesia. Raffles Bulletin of Zoology 62: 718-725

Farson HK, Kottelat M (1992) A new species

of Mugilogobius (Pisces: Gobiidae) from Fake Matano, cen-

tral Sulawesi, Indonesia. Ichthyological Exploration of Fresh-

waters 3: 225-234

Farson HK, Murdy EO (2001) Eleotrididae. Sleepers (gudgeons).

Pp. 3574-3577 in: Carpenter KE & Niem VH (eds.) FAO

species identification guide for fishery purposes. The living

marine resources of the Western Central Pacific. Vol. 6. Bony

fishes part 4 (Fabridae to Fatimeriidae), estuarine crocodiles.

FAO, Rome

Farson HK, Williams RD, Hammer MP (2013) An annotated

checklist of the fishes of the Northern Territory, Australia.

Zootaxa 3696: 1-293

Fast PR, Stevens JD (1994) Sharks and rays of Australia. CSIRO

Publishing, Melbourne

Fesson RP (1829-1831) Poissons. Pp. 66-238 in: Fesson RP

(ed.) Voyage autour du monde, execute par Ordre du Roi, sur

la corvette de Sa Majeste, Fa Coquille, pendant les annees

1822, 1823, 1824 et 1825, par M. F. 1. Duperrey. Zoologie 2

(1). Atlas. Arthus Bertrand, Paris

Finnaeus C ( 1758) Systema naturae per regna tria naturae, secun-

dum classes, ordines, genera, species, cum characteribus, dif-

ferentiis, synonymis, locis. Tomus 1. Editio decima, reforma-

ta. Holmiae. Faurentius Salvius, Stockholm

Finnaeus C (1766) Systema naturae per regna tria naturae, secun-

dum classes, ordines, genera, species, cum characteribus, dif-

ferentiis, synonymis, locis. Tomus 1. Editio duodecima, refor-

mata. Holmiae. Faurentius Salvius, Stockholm

Fovejoy NR, Iranpour M, Collette BB (2004) Phylogeny and jaw

ontogeny of beloniform fishes. Integrative and Comparative

Biology 44: 366-377

Maeda K, Yamasaki N, Tachihara K (2007) Size and age at Re-

cruitment and Spawning Season of Sleeper, Genus Eleotris

(Teleostei: Eleotridae) on Okinawa Island, Southern Japan.

Raffles Bulletin of Zoology Supplement 14: 199-207

Manilo EG, Bogorodslcy SV (2003) Taxonomic composition, di-

versity and distribution of coastal fishes of the Arabian Sea.

Journal of Ichthyology 43: 75-149

Manjaji-Matsumoto BM, Fast PR (2008) Himantura

leoparda sp. nov., a new whipray (Myliobatoidei: Dasyatidae)

from the Indo-Pacific. CSIRO Marine and Atmospheric Re-

search Paper 22: 292-301

Marion de Proce PM (1822) Sur plusieurs especes nouvelles de

poissons et de crustaces observees. Bulletin de la Societe

Philomathique de Paris 1822: 129-134

Matsuura K (2001) Order Tetraodontiformes, TriaCanthodida

Spikefishes. Pp. 3902-3910 in: Carpenter KE & Niem VH

(eds.) FAO species identification guide for fishery purposes.

The living marine resources of the Western Central Pacific.

Volume 6. Bony fishes part 4 (Fabridae to Fatimeriidae), es-

tuarine crocodiles, sea turtles, sea snakes and marine mam-

mals. FAO, Rome

Mauge FA (1986) Gobiidae, Pp. 358-388 in: Daget J, Gosse JP

& Thys van den Audenaerde DFE (eds) Check-list of the fresh-

water fishes of Africa (CFOFFA) Vol. 2. ISNB, Brussels;

MRAC, Tervuren; and ORSTOM, Paris

McClelland J (1844) Apodal fishes of Bengal. Calcutta Journal

of Natural History 5: 151-226

McEachran JD, Carvalho MD (2002) Batoid fishes. The living

marine resources of the Western Central Atlantic 1: 507-589

McGinley M, Hogan CM (2003) Biological diversity in Sunda-

land. Encyclopedia of Earth. Online at

http://www.eoearth.org/view/article/150630/ last accessed on

October 20, 2015

McKay RJ (2001) Haemulidae (=Pomadasyidae), Grunts (also

sweetlips, rubberlips, hotlips, and velvetchins). Pp. 2961-2989

in: Carpenter KE & Niem VH (eds) FAO species identifica-

tion guide for fishery purposes. The living marine resources

of the Western Central Pacific. Volume 5. Bony fishes part 3

(Menidae to Pomacentridae). FAO, Rome

Meisner AD (2001) Phylogenetic systematics of the viviparous

halfbeak genera Dermogenys and Nomorhamphus (Teleostei:

Hemiramphidae: Zenarchopterinae). Zoological Journal of the

Finnean Society 133: 199-283

Meisner AD, Fouie KD (2000) Nomorhamphus koJonodaJensis,

a new species of viviparous halfbeak from Sulawesi (Teleostei:

Hemiramphidae). Ichthyological Exploration of Freshwaters

11: 361-368

Mokodongan DF, Tanaka R, Yamahira K (2014) A new ricefish

of the genus Oryzias (Beloniformes, Adrianichthyidae) from

Fake Tiu, Central Sulawesi, Indonesia. Copeia 3: 561-567

Mokodongan DF & Yamahira K (2015) Origin and intra-island

diversification of Sulawesi endemic Adrianichthyidae Molec-

ular Phylogenetics and Evolution 93: 150-160

Motomura H (2004) Threadfins of the world (Family Polyne-

midae): An annotated and illustrated catalogue of polynemid

species known to date (No. 3). FAO, Rome

Muller J, Henle F (1838-1841) Systematische Beschreibung der

Plagiostomen. Veit, Berlin

Munro ISR (1958) The fishes of the New Guinea region. A

check-list of the fishes of New Guinea incorporating records

of species collected by the Fisheries Survey Vessel “Fairwind”

during the years 1949 to 1950. Territory of Papua and New

Guinea. Fisheries Bulletin 1: 97-369

Myers RF (1991) Micronesian reef fishes. Second Ed. Coral

Graphics, Barrigada, Guam

Myers N, Mittermeier RA, Mittermeier CG, da Fonseca GA,

Kent J (2000) Biodiversity hotspots for conservation priori-

ties. Nature 403: 853-858. doi: 10.1038/35002501

Nasution SH, Aisyah S (2013) Zonation of endemic ichthyofau-

na conservation area in ancient Fake Towuti, South Sulawe-

si-Indonesia. FIMNOTEK 20: 207-216

Nelson JS (2006) Fishes of the World. Second Ed. John Wiley

& Sons, New York Chichester, Brisbane, Toronto, Singapore

Ng HH, Kottelat M (2007) The identity of Clarias batrachus

(Finnaeus 1758) with the designation of a neotype (Teleostei:

Clariidae). The Finnean Society of Fondon, Zoological Jour-

nal of the Finnean Society 153: 725-732

Omar SBA (2010) Reproductive biology of bonylip barb, Os-

teochilus vittatus Valenciennes, 1842) in Sidenreng Fake,

South Sulawesi. Jurnal Iktiologi Indonesia 10 (2): 111-122

Pallas PS (1767) in: Schlosser JA (1767) Some further intelli-

gence relating to the jaculator fish, mentioned in the Philo-

sophical Transactions for 1764, Art. XIV, from Mr. Hommel,

at Batavia, together with the description of another species,

by Dr. Pallas. Philosophical Transactions 56 (1766 [1767]):

187-188

Bonn zoological Bulletin 64 (2): 77-106

©ZFMK

104

Friedrich Wilhelm Miesen et al.

Pallas PS (1770) Spicilegia Zoologica quibus novae imprimis

et obscurae animalium species iconibus, descriptionibus atque

commentariis illustrantur. Gotti. August. Lange

Parent! LR (1996) Phylogenetic systematics and biogeography

of phallostethid fishes (Atherinomorpha, Phallostethidae) of

northwestern Borneo, with description of new species.

Copeia 3; 703-712

Parent! LR (2008) A phylogenetic analysis and taxonomic revi-

sion of ricefishes, Oryzias and relatives (Beloniformes, Adri-

anichthyidae). ZoologicaJJournal of the Linnean Society 154:

494-610

Parenti LR (2011) Endemism and conservation of the native

freshwater fish fauna of Sulawesi, Indonesia. Prosiding Sem-

inar Nasional Ikan 1; 1-10

Parenti LR, Hadiaty RK (2010) A new, remarkably colorful,

small ricefish of the genus Oryzias (Beloniformes, Adri-

anichthyidae) from Sulawesi, Indonesia. Copeia 2010 (2);

268-273

Parenti L, Hadiaty R, Lumbantobing D, Herder F (2013) Two

New Ricefishes of the Genus Oryzias (Atherinomorpha; Be-

loniformes; Adrianichthyidae) Augment the Endemic Fresh-

water Fish Fauna of Southeastern Sulawesi. Copeia 2013 (3):

403-414

Parenti ER, Eouie KD (1998) Neostethus djajaorum, new

species, from Sulawesi, Indonesia, the first phallostethid fish

(Teleostei: Atherinomorpha) known from east of Wallace’s

line. Raffles Bulletin of Zoology 43: 139-150

Parenti ER, Soeroto B (2004) Adrianichthys roseni and Oryzias

nebulosus, two new ricefishes (Atherinomorpha: Beloni-

formes: Adrianichthyidae) from Eake Poso, Sulawesi, Indone-

sia. Ichthyological Research 51: 10-19

Peters W (1852) Diagnosen von neuen Flussfischen aus Mos-

sambique. Monatsberichte der Koniglichen Preussischen

Akademie der Wissenschaften zu Berlin 1852: 275-276 +

681-685

Peters W (1859) Eine neue vom Herrn Jagor im atlantischen

Meere gefangene Art der Gattung Leptocephalus, und ilber ei-

nige andere neue Fische des Zoologischen Museums. Monats-

berichte der Koniglichen Preussischen Akademie der Wissen-

schaften zu Berlin 1859: 411-413

Peters W (1868) Uber die von Hrn. Dr. F. Jagor in dem ostindi-

schen Archipel gesammelten und dem Konigl. zoologischen

Museum iibergebenen Fische. Monatsberichte der Koniglichen

Preussischen Akademie der Wissenschaften zu Berlin 1868:

254-281

Peters W (1869) Uber neue oder weniger bekannte Fische des

Berliner Zoologischen Museums. Monatsberichte der Konig-

lichen Preussischen Akademie der Wissenschaften zu Berlin

1869: 703-711

Pfaender J, Gray SM, Rick IP, Chapuis S, Hadiaty RK, Herder

F (2014) Spectral data reveal unexpected cryptic colour po-

lymorphism in female sailfm silverside fish. Hydrobiologia

739: 155-161

Pfaender J, Miesen FW, Hadiaty RK, Herder F (201 1) Adaptive

speciation and sexual dimorphism contribute to diversity in

form and functional in the adaptive radiation of Eake

Matano’s sympatric roundfm sailfln silversides. Journal of

Evolutionary Biology 24: 2329-2345

Pfaender J, Schliewen UK, Herder F (2010) Phenotypic traits

meet patterns of resource use in the radiation of “sharpfm” sail-

fin silverside fish in Eake Matano. Evolutionary Ecology 24:

957-974

Popta CME (1905) Note XXII. Haplochilus sarasinorum, n.sp.

Notes from the Eeyden Museum 25: 239-247

Bonn zoological Bulletin 64 (2); 77-106

Popta CME (1912) Fortsetzung der Beschreibung von neuen

Fischarten der Sunda-Expedition. Notes from the Eeyden Mu-

seum 34: 185-193

Popta CME (1921) Dritte Fortsetzung der Beschreibung von neu-

en Fischarten der Sunda-Expedition. Zoologische Mededelin-

gen 6: 203-214

Popta CME (1922) Vierte und letzte Fortsetzung der Beschrei-

bung von neuen Fischarten der Sunda-Expedition. Zoologis-

che Mededelingen 7: 27-39

Pyke GH (2006) A review of the biology of Gambusia affinis

and G. holbrooki. Review in Fish Biology and Fisheries 15;

339-365

Quoy JRC, Gaimard JP (1824-25) Description des Poissons.

Chapter IX. In; Freycinet, E. de. Voyage autour du

Monde... execute sur les corvettes de E. M. “E’Uranie” et “Ea

Physicienne,” pendant les annees 1817, 1818, 1819 et 1820.

Paris

Rachman A, Rianse U, Musaruddin M, Pasolon Y (2015) The

potential of delivering clean locally available limitless rice

husk energy in the Celebes Island Indonesia. Energy Proce-

dia 00; 1-6

Randall JE (1985) On the validity of the Tetraodontid Fish

Arothron manilensis (proce). Japanese Journal of Ichthyolo-

gy 32: 347-354

Randall JE, Bauchot ME (1999) Clarification of the two Indo-

Pacific species of bonefishes, Albula glossodonta and A.

forsteri. Cybium 23; 79-83

Randall JE (2001) MUEEIDAE, Goatfishes (surmullets). Pp.

3175-3200 in: Carpenter KE & Niem VH (eds.) FAO species

identification guide for fishery purposes. The living marine

resources of the Western Central Pacific. Volume 5. Bony fish-

es part 3 (Menidae to Pomacentridae). FAO, Rome

Randall JE, Allen GR and Steene RC (1990) Fishes of the Great

Barrier Reef and Coral Sea. University of Hawaii Press, Hon-

olulu, Hawaii

Regan CT (1910) The Asiatic fishes of the family Anabantidae.

Proceedings of the Zoological Society of Eondon 1909 (4);

767-787

Regan CT (1911) On some fishes of the family Poeciliidae. 2.

The poeciliid fishes of Celebes and Eombok. Annals and Mag-

azine of Natural History 8; 374

Richardson J (1844-48) Ichthyology of the voyage of H. M. S.

Erebus & Terror,... Pp. 1-139 in; Richardson J & Gray JE (eds)

The zoology of the voyage of H. H. S. “Erebus & Terror,” un-

der the command of Captain Sir J. C. Ross ... during ...

1839-43 (see Bauchot et al. 1982; 66)

Richardson J (1846a) Report on the ichthyology of the seas of

China and Japan Reports of the meetings of the British Asso-

ciation for the Advancement of Science 1845; 187-320

Rintelen K von, Rintelen T von, Glaubrecht M (2007a) Molec-

ular phylogeny and diversification of freshwater shrimps (De-

capoda, Atyidae, Caridina) from ancient Eake Poso (Sulawe-

si, Indonesia) - The importance of being colourful. Molecu-

lar Phylogenetics and Evolution 45: 1033-1041

Rintelen T von, Bouchet P, Glaubrecht M (2007b) Ancient lakes

as hotspots of diversity: a morphological review of an endem-

ic species flock of Tylomelania (Gastropoda; Cerithioidea;

Pachychilidae) in the Malili lake system on Sulawesi, Indone-

sia. Hydrobiologia 592: 11-94. doi: 10. 1007/s 10750-007-

0765-8

Rintelen T von, Rintelen K von, Glaubrecht M (2010) The

Species Flocks of the Viviparous Freshwater Gastropod Ty-

lomelania (Mollusca: Cerithoidea; Pachychilidae) in the An-

cient Takes of Sulawesi, Indonesia: The role of Geography,

Trophic Morphology and Color as Driving Forces in Adap-

©ZFMK

An annotated checklist of the inland fishes of Sulawesi

105

tive Radiation. Pp. 485-512 in; GlaubrechtM (ed.) Evolution

in Action -Adaptive Radiations and the Origins of Biodiver-

sity. Springer, Heidelberg

Rintelen T von, Rintelen K von, Glaubrecht M, Schubart CD,

Herder F (2012) Aquatic biodiversity hotspots in Wallacea:

the species flocks in the ancient lakes of Sulawesi, Indone-

sia. Pp. 290-315 in: Biotic Evolution and Environmental

Change in Southeast Asia. Cambridge University Press, Cam-

bridge

Rosen DE (1964) The relationships and taxonomic position of

the halfbeaks, killifishes, silversides, and their relatives. Bul-

letin of the American Museum of Natural History 127: arti-

cle 5

Rosen DE, Greenwood PH (1976) A fourth Neotropical species

of synbranchid eel and the phytogeny and systematics of syn-

branchiform fishes. Bulletin of the American Museum of Nat-

ural History 157: article 1

Riippell WPES (1835-38) Neue Wirbelthiere zu der Fauna von

Abyssinien gehorig. Fische des Rothen Meeres. Siegmund

Schmerber, Frankfurt am Main (For dates see Sawyer 1952)

Russel BC (2001) NEMIPTERIDAE, Threadfm breams (also

whiptail breams, monocle breams, dwarf monocle breams and

coral breams). Pp. 3051-3089 in; Carpenter KE & Niem VH

(eds) FAO species identification guide for fishery purposes.

The living marine resources of the Western Central Pacific.

Volume 5. Bony fishes part 3 (Menidae to Pomacentridae).

FAO, Rome

Santini F, Tyler JC (2002) Phytogeny and biogeography of the

extant species of triplespine fishes (Triacanthidae, Tetraodon-

tiformes) Zoologica Scripta 31: 321-330

Santos MCF, Ruffino ME, Farias IP (2007) High levels of ge-

netic variability and panmixia of the tambaqui Colossoma

macropomum (Cuvier, 1 8 1 6) in the main channel of the Ama-

zon River. Journal of Fish Biology 71 (Supplement A): 33-44

Sasaki K (2001) SCIAENIDAE; Croakers (drums). Pp.

3117-3174 in: Carpenter KE & Niem VH (eds) FAO species

identification guide for fishery purposes. The living marine

resources of the Western Central Pacific. Volume 5. Bony fish-

es part 3 (Menidae to Pomacentridae). FAO, Rome

Sawyer FC (1952) The dates of publication of Wilhelm Peter Ed-

uard Simon Riippelf s [1794-1884] “Neue Wirbelthiere zu der

Fauna von Abyssinien gehorig” (fob, Frankfurt a. M.,

1835-1840). Journal of the Society for the Bibliography of

Natural History 2: 407

Schlosser JA (1767) Some further intelligence relating to the jac-

ulator fish, mentioned in the Philosophical Transactions for

1764, Art. XIV, from Mr. Hommel, at Batavia, together with

the description of another species, by Dr. Pallas. Philosophi-

cal Transactions 56 (1766 [1767]); 187-188

Schneider JG (1801) M. E. Blochii, Systema Ichthyologiae Ico-

nibus ex Ilustratum. Post obitum auctoris opus inchoatum ab-

solvit, correxit, interpolavit Jo. Gottlob Schneider, Saxo.

Berolini. Sumtibus Auctoris Impressum et Bibliopolio Sande-

riano Commissum

Schultz FP, Herald ES, Fachner FA, Welander AD, Woods

FP (1953) Fishes of the Marshall and Marianas islands. Vol.

I. Families from Asymmetrontidae through Siganidae. Bulletin

of the United States National Museum 202: 1-685

Schwarzer J, Herder F, Misof B, Hadiaty RK, Schliewen UK

(2008) Gene flow at the margin of Fake Matano’s adaptive

sailfin silverside radiation; Telmatherinidae of River Petea in

Sulawesi. Hydrobiologia 615: 201-213

Seale A (1910) Descriptions of four new species of fishes from

Bantayan Island, Philippine Archipelago. The Philippine Jour-

nal of Science Section D 5: 115-119

Bonn zoological Bulletin 64 (2): 77-106

Senou H (2001) Suborder Scombroidei, Sphyraenidae Barracu-

das. Pp. 3685-3697 in: Carpenter KE & Niem VH (eds) FAO

species identification guide for fishery purposes. The living

marine resources of the Western Central Pacific. Volume 6.

Bony fishes part 4 (Fabridae to Fatimeriidae), estuarine croc-

odiles, sea turtles, sea snakes and marine mammals. FAO,

Rome

Sherborn CD, Woodward BB (1901). Notes on the dates of pub-

lication of the natural history portions of some French voy-

ages. - Part I. ‘Amerique meridionale’; Tndes orientates’;

‘Pole Sud’ (‘Astrolabe’ and ‘Zelee’); ‘Fa Bonite’; ‘Fa Co-

quille’; and ‘F’Uranie et Physiciennes’ . Annals and Magazine

of Natural History 7: 388-392

Smith HM (1932) Contributions to the ichthyology of Siam. I.

Descriptions of a new genus and three new species of Siamese

gobies. Journal of the Siam Society, Natural History Supple-

ment 8: 255-262

Smith HM (1945) The fresh- water fishes of Siam, or Thailand.

Bulletin of the United States National Museum 188: 1-622

Smith DG (1994) Catalog of type specimens of recent fishes in

the National Museum of Natural History, Smithsonian Insti-

tution, 6: Anguilliformes, Saccopharyngiformes, and Notacan-

thiformes (Teleostei: Elopomorpha). Smithsonian Contribu-

tions to Zoology 566; 1-50

Smith- Vaniz WF (1999) Carangidae. In: Fischer W & Bianchi

G (eds) FAO species identification sheets for fishery purpos-

es. Western Indian Ocean (fishing area 5 1 ). Vol. 1 . FAO, Rome

Soeroto B, Tungka F (1991) Fish fauna, fisheries and Adri-

anichthyoidei in Fake Poso. Pp. 12-14 in; Phylogeny and

species differentiation of Adrianichthyoidei in Indonesia.

Study Report. Monbusho International Scientific Research

Program

Soeroto B, Tungka F (1996) The inland fishes and the distribu-

tion of Adrianichthyoidea of Sulawesi Island, with special

comments on the endangered species in Fake Poso. Pp. 1-5

in: Kichener DJ & Suyanto A (eds) Proceedings of the first

international conference on eastern Indonesian- Australian ver-

tebrate fauna, Manado, Indonesia, November 22-26, 1994.

Perth: Western Australian Museum for Fembaga Ilmu Penge-

tahuan Indonesia

Steindachner F (1868) Ichthyologische Notizen (VII). Sitzungs-

berichte der KaiserlichenAkademie der Wissenschaften. Ma-

thematisch-Naturwissenschaftliche Classe 57; 965-1008

Steindachner F (1879) Ichthyologische Beitrage (VIII). Sitzungs-

berichte der Kaiserlichen Akademie der Wissenschaften.

Mathematisch-Naturwissenschaftliche Classe 80: 119-191

Stelbrink B, Albrecht C, Hall R, Rintelen T von (2012) The bio-

geography of Sulawesi revisited: is there evidence for a vic-

ariant origin of taxa on Wallace’s “anomalous island”? Evo-

lution 66: 2252-2271

Talwar PK, Jhingran AG (1991) Inland fishes of India and ad-

jacent countries. Volume 2. A.A. Balkema, Rotterdam

Tamsil A (2000) Studi Beberapa Karakteristik Reproduksi

Prapemijahan dan Kemungkinan Pemijahan Buatan Ikan Bun-

go {Glossogohius cf aureus) di Danau Tempe dan Danau

Sidenreng Sulawesi Selatan. Program Pascasarjana Institut

Pertanian Bogor

Tan HH (2006) The Borneo Suckers. Natural history Publica-

tions (Borneo), Kinabalu

Tan HH, Fim KKP (2004) Inland fishes from the Anambas and

Natuna islands. South China Sea. With description of a new

species of Betta (Teleostei: Osphronemidae). Raffles Bulletin

of Zoology Supplement 11: 107-115

Tsukamoto K, Watanabe S, Kuroki M, Aoyama J, Miller MJ

(2014) Freshwater habitat use by a moray eel species, Gym-

©ZFMK

106

Friedrich Wilhelm Miesen et al.

nothorax polyuranodon, in Fiji shown by otolith microchem-

istry. Environmental Biology of Fishes 97: 1377-1385. DOI

10. 1007/s 1064 1-0 14-0228-9

Tweedley JR, Bird DJ, Potter IC, Gill HS, Miller PJ, O’Dono-

van G, TJakrawidJaJa AH (2013) Species compositions and

ecology of the riverine ichthyofaunas in two Sulawesian is-

lands in the biodiversity hotspot of Wallacea. Journal of fish

biology 82: 1916-50. doi: 10.1111/jfb.l2121

Usman BI, Amin SMN, Arshad AA, Aminur Rahman M (2013)

Review of Some Biological Aspects and Fisheries of Grey-

Eel Catfish Plotosius canius (Hamilton, 1822). Asian Journal

of Animal and Veterinary Advances 8: 154-167

Vaillant JJ, Haffner GD, Cristescu ME (2011) The ancient lakes

of Indonesia: Towards integrated research on speciation. In-

tegrative and comparative biology 51: 634-43. doi:

10.1093/icb/icrl01

Vari RP (2001) TERAPONTIDAE (= Therapontidae, Thera-

ponidae, Teraponidae), Terapon perches (terapon grunters). Pp.

3305-3316 in: Carpenter KE & Niem VH (eds) FAO species

identification guide for fishery purposes. The living marine

resources of the Western Central Pacific. Volume 5. Bony fish-

es part 3 (Menidae to Pomacentridae). FAO, Rome

Vari RP, Hadiaty RK (2012). The endemic Sulawesi fish genus

Eagusia (Teleostei, Terapontidae). Raffles Bulletin of Zoolo-

gy 60: 157-162

Vogt D (1978) Kennen sie Nomorhamphusl Eine Ubersicht und

vorlaufige Beschreibungen einer neuen Unterart, namlich No-

morhamphus Jiemi snijdersi und einer Art, namlich Nomor-

hamphus brembachi. Aquarien und Terrarien-Zeitschrift 3 1 :

222-227

Watson RE (1991) A provisional review of the genus Stenogo-

bius with descriptions of a new subgenus and thirteen new

species (Pisces: Teleostei: Gobiidae). Records of the Western

Australian Museum 15 (3): 627-710

Watson RE (1992) A review of the gobiid fish genus Awaous

from insular streams of the Pacific Plate. Ichthyological Ex-

plorations of Freshwaters 3: 161-176

Watson RE (1995) Gobies of the genus Stiphodon from French

Polynesia, with descriptions of two new species (Teleostei:

Gobiidae: Sicydiinae). Ichthyological Explorations of Fresh-

waters 6: 33-48

Watson RE ( 1 996) A review of Stiphodon from New Guinea and

adjacent regions, with descriptions of five new species

(Teleostei: Gobiidae: Sicydiinae). Revue franqaise d’aquario-

logie 23: 113-132

Weber M (1894a) Die Siisswasser-Fische des Indischen Archi-

pels, nebst Bemerkungen iiber den Ursprung der Fauna von

Celebes. Zoologische Ergebnisse einer Reise inNiederlandisch

Ost-Indien 3: 405-476

Weber M (1894b) Die Siisswasser-Fische des Indischen Archi-

pels, nebst Bemerkungen iiber den Ursprung der Fauna von

Celebes. Zoologische Ergebnisse einer Reise in Niederlan-

disch Ost-Indien: 405-476

Weber M (1895) Fische von Ambon, Java, Thursday Island, dem

Burnett-Fluss und von der Sud-Kuste von Neu-Guinea. In:

Zoologische Forschungsreisen in Australien und dem ma-

layischen Archipel; mit Unterstutzung des Herrn Dr. Paul von

Ritter ausgeftihrt ...Jahren 1891-1893 von Dr. Richard Semon

5: 259-276

Weber M (1909) Diagnosen neuer Fische der Siboga-Expediti-

on. Notes from the Eeyden Museum 31: 143-169

Weber M (1911) Die Fische der Aru- und Kei-Inseln. Ein Bei-

trag zur Zoolographie dieser Inseln. Abhandlungen der Sen-

ckenbergischen Naturforschenden Gesellschaft Frankfurt am

Mam 34: 1-49

Weber M (1913) Die Fische der Siboga-Expedition. Siboga- Ex-

peditie. Brill EJ, Eeiden

Weber M, de Beaufort EF (1912) Over de zoetwatervisshen van

Timor en Babber. Verslagen van de gewone vergaderingen der

Wissen Natuurkundige Afdeeling der Koninklijke Akademie

van Wetenschappen te Amsterdam 1912: 133-138

Weber M, de Beaufort, EF (1922) The fishes of the Indo-Aus-

tralian Archipelago. IV. Heteromi, Solenichthyes, Synentog-

nathi, Percesoces, Eabyrinthici, Microcyprini. Brill, Eeiden

Weber M, de Beaufort EF (1953) The fishes of the Indo-Aus-

tralian archipelago Vol. X Gobioidea. Brill, Eeiden

Whitehead PJP, Nelson GJ, Wongratana T (1988) FAO Species

Catalogue. Vol. 7. Clupeoid fishes of the world (Suborder Clu-

peoidei). An annotated and illustrated catalogue of the her-

rings, sardines, pilchards, sprats, shads, anchovies and wolf-

herrings. FAO, Rome

Whitley GP (1935a) Fishes from Princess Charlotte Bay, North

Queensland. Records of the South Australian Museum 5:

345-365

Whitley GP (1938) Descriptions of some New Guinea fishes.

Records of the Australian Museum 20: 223-233

Whitten AJ, Bishop KD, Nash SV, Clayton E (1987). One or

More Extinctions born Sulawesi, Indonesia. Conservation Bi-

ology 1: 42-48

Whitten AJM, Mustafa M, Henderson GS (2002) The Ecology

of Sulawesi. Periplus, Singapore

Wilson AB, Orr JW (2011) The evolutionary origins of Syng-

nathidae: pipefishes and seahorses. Journal of Fish Biology

78: 1603-1623

Wongratana T, Munroe TA, Nizinski MS (1999) Order Clu-

peiformes, Engraulididae, Anchovies. In: Carpenter KE &

Niem VH (eds) FAO species identification guide for fishery

purposes. The living marine resources of the Western Central

Pacific. Volume 3. Batoid fishes, chimaeras and bony fishes

part 1 (Elopidae to Einophrynidae). FAO, Rome

Wueringer BE, Squire E, Collin SP (2009) The biology of ex-

tinct and extant sawfish (Batoidea: Sclerorhynchidae and Pris-

tidae). Reviews in Fish Biology and Fisheries 19: 445-464

Yamanoue Y, Miya M, Doi H, Mabuchi K, Sakai H, & Nishida

M (2011) Multiple invasions into freshwater by pufferfishes

(Teleostei: Tetraodontidae): A mitogenomic perspective.

PEoS One 6 (2): el 74 10

Yokoo T, Kanou K, Moteki M, Hiroshi K, Tongnunui P, Kuroku-

ra H (2006) Juvenile morphology and occurrence patterns of

three Butis species (Gobioidei: Eleotridae) in a mangrove es-

tuary, southern Thailand. Ichthyological Research 53: 330-336

Zuiew B (1793) Biga Mvraenarvm, novae species descriptae.

Nova Acta Academiae Scientiarum Imperialis Petropolitanae

7 (for 1789): 296-301

Bonn zoological Bulletin 64 (2): 77-106

©ZFMK

Bonn zoological Bulletin 64 (2): 107-108

March 2016

Correction & update

On the Linck collection and specimens of snakes

figured by Johann Jakob Scheuchzer (1735) -

the oldest fluid-preserved herpetological collection in the world?

Bauer, A.M. & R. Wahlgren (2013). Bonn Zoological Bulletin 62: 220-252

Aaron M. Bauer

Department of Biology, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085, U.S.A.

aaron. bauer@villanova. edu; + 1-610-519-485 7

The contributions of Dr. Konstantin Wopke to the reor-

ganization of the Linck collection in the Natural ienkabi-

nett Waldenburg have been outlined by Bauer and

Wahlgren (2013). Unfortunately, Dr. Wopke was inadver-

tently misidentified in Figure 7 (p. 227) of this paper. Orig-

inally identified as the figure on the left, he is, in fact, the

figure on the right of the photo, wearing a white lab coat.

The figure on the left of the photo is the mineralogist Al-

fred Seifert. This photograph had earlier appeared, correct-

ly labeled, in Budig (1999).

Through the kindness of Mr. Dietrich Wopke, who has

shared some biographical data with me, I am able to pro-

vide some further information regarding his father. Kon-

stantin Wopke was born 29 July 1905 in Crimmitschau,

17 km west of Waldenburg, as the son of Richard and He-

lene (nee Brendel) Wopke. He attended school in Leipzig

until the outbreak of World War I and then continued his

schooling in Gotha. He studied zoology, botany, chemistry

in Jena, Freiburg im Breisgau and in Leipzig. In 1930 he

was promoted to the degree of Dr. phil. on the basis of

his dissertation “Die Kloake und die Begattungsorgane der

mannlichen Zauneidechse {Lacerta agilis L.)” complet-

ed in Leipzig under the direction of Prof Dr. Johannes

Meisenheimer (1873-1933), a specialist on the develop-

ment of invertebrates, and published in Jena (Wopke

1930). He subsequently worked as a teaching assistant at

the Zoological Institute of the University of Leipzig and

as a research assistant at the Anatomical Institute in

Wurzburg. From 29 May 1933 to 6 July 1935 he re-or-

ganized the approximately 5000 zoological objects in the

Furstlich-Schonburg-Waldenburgische Naturalienkabi-

nett in Waldenburg, struggling with misidentifications,

mislabeling, and a century or more of the intermixing of

more recently acquired specimens with the 17* and 18*

century material of the original Linck collection.

Received: 23.03.2015

Accepted: 06.11.2015

In January 1934 Wopke passed the state examination

for secondary school teachers in the subjects of zoology,

botany and chemistry. After graduating from his intern-

ship year at the Furstlich-Schonburgischen Deutschen

Oberschule he was employed as a private tutor at the Kam-

mergut in Hardisleben in Thuringia. He was subsequent-

ly excluded from school employment by the Saxon Min-

istry of Education on political grounds and instead, from

1936 to 1939 he served as a research assistant in the Im-

perial Biological Institute in Naumburg/Saale, where he

worked on “Phylloxera in the wine growing areas of Saale

and Unstrut.” The documentation and results of this work,

although apparently unpublished, are maintained in the

German Federal Archives. Although he had moved on

from Waldenburg, in 1937 Wopke’s opinion was solicit-

ed about the renovation of the collections of the Franck-

esche Stiftungen in Haale (Mojsejenko 2013) and in the

same year his guide to the Naturalienkabinett in Walden-

burg was published (Wopke 1937). In August 1939 he was

sent to military service and in April 1944 he was killed

in Russia. Although Wopke’s scientific career was cut

short, his contribution to the reorganization of the Linck

collection in Waldenburg is a lasting legacy.

REFERENCES

Budig U (1999) Ein Kabinett voher Raritaten - Die Einck

Sammlung. Pp. 16-31 in: Budig U, Zenker R & Stadt-

lander G. (eds) Heimatmuseum und Naturalienkabinett

Waldenburg, SachsischeMuseen7. Sachsische Eandes-

stehe fur Museumswesen, Chemnitz

Mojsejenko, A (2013) Die Kunst- und Naturalienkammer

der Franckeschen Stiftung in Halle an der Saale. Sem-

inararbeit im Erganzungsmodul des Masterstudien-

ganges, Universitat Duisburg-Essen, 24 pp.

Corresponding editor: Ph. Wagner

108

Aaron M. Bauer

Wopke K (1930) Die Kloake und die Begattungsorgane

der mannlichen Zauneidechse (Lacerta agilis L.). Je-

naische Zeitschrift fiir Naturwissenschaft 65 : 275-3 1 8,

pis iv-vi

Wopke K (1937) Fiihrer durch das Fiirstlich Schonburgi-

sche Natural ienkabinett in Waldenburg/Sa. und die in

ihm enthaltenen Sammlungen. Fiirstlich Schonburgis-

che Naturalienkab inert in Waldenburg, Waldenburg, 38

pp.

Bonn zoological Bulletin 64 (2); 107-108

©ZFMK

Bonn zoological Bulletin 64 (2): 109-116

March 2016

A new genus and new species of Neotropical Thoracophorini

(Coleoptera: Staphylinidae: Osoriinae)

Ulrich Irmler

Institut for Ecosystem Research, Dept. Applied Ecology, University of Kiel, Olshausenstrasse 40, D-24098 Kiel, Germany;

E-mail: uirmler@ecology. uni-kiel.de

Abstract. Three new species of the new genus Geotrochopsis are described: G. pubescens, G. collaris, and G.flaveolus.

The genus is placed in the subtribe Clavilispinina of the tribe Thoracophorini. G. pubescens seems to be distributed all

over the Neotropical region. Both G. collaris and G.flaveolus are from Peru and Brazil, respectively. Additionally, one

new species is described from the Central Amazon: Geomitopsis amazonensis. Furthermore, Ashnaosorius Makhan, 2008

is recognised as a new synonym of Geomitopsis.

Key words. New species, Osoriinae, Thoracophorini, Neotropics

INTRODUCTION

During the studies on the Neotropical Osoriinae few spec-

imens of blind species were found that belong to the tribe

Thoracophorini. Hitherto among this subtribe, only the

blind genera Geomitopsis Scheerpeltz, 1931 and Ash-

naosorius Makhan, 2008 were known from the Neotrop-

ical region. A more detailed study came to the result that

some specimens belong to a new genus. The present pa-

per describes this new genus with its new species. Fur-

thermore, Ashnaosorius is a new synonym of Geomitop-

sis. According to Herman (2001) the genus Geomitopsis

is recorded also from the Mediterranean region from

Libanon to Canary Islands with nine species and from

Africa with six species. Including the new Geomitopsis

species, a total of six species is also known from the

Neotropics. Therefore, a key to the species of the

Neotropical region is provided.

MATERIAL AND METHODS

The material studied in this investigation is presently de-

posited in the following museums and private collections.

AMNH American Museum of Natural History,

New York

BMNH British Museum, Natural History, London

INPA Collections of Instiuto National de

Pesquisas da Amazonia, Manaus, Brazil

KNHM Kansas Natural History Museum, Lawrence

NHMP National Museum of Natural History,

Czec Republic, Prague

Received: 15.05.2014

Accepted: 18.03.2016

ZFMK Museum Alexander Koenig, Bonn, Germany

JJC Private collection of Jifi Janak, Prague,

Czech Republic

UIC Private collection of U. Irmler, Plon, Germany

The photographs were taken using a Makroskop M 420

(Wild, Herbrugg) in combination with a digital camera

(LeicaECS). CombineZS (Hadley 2006) was used to op-

timise depth of focus. Length was measured in the mid-

dle of tagmata: head from clypeus to posterior edge,

pronotum from anterior to posterior edge along midline,

elytra from anterior edge of shoulders to posterior edge;

width at the widest part of tagmata (head width includes

eyes). In the measurement of total length, the abdominal

inter-segmental space is subtracted. The aedeagus was dis-

sected and drawings were made using a microscope un-

der 250 X magnification.

DESCRIPTION OF THE NEW SPECIES

Geotrochopsis n. gen.

Type species. Geotrochopsis pubescens n.sp. is here des-

ignated as the type species

Diagnosis. Geotrochopsis is similar to the other blind

genus in the tribe Thoracophorini, i.e. Geomitopsis

Scheerpeltz, 193 1. In contrast to Geomitopsis Scheerpeltz,

1931, Geotrochopsis is densely pubescent on the whole

body and the antennomere six is not narrower than anten-

nomeres five and seven. Moreover, the tarsi of Geotro-

chopsis are composed of five tarsomeres, whereas they are

Corresponding editor: D. Ahrens

110

Ulrich Irmler

composed of four tarsomeres in Geomitopsis . Addition-

ally, the aedeagus of Geotrochopsis is symmetric and with-

out a ventral prominence such as in Geomitopsis.

Description. Length and habitus: elongate; blind with re-

duced elytra; small species of about 1.3-1. 6 mm total

length.

Head approximately square; clypeus semicircular;

labrum divided into two lobes separated by a deep emar-

gination; eyes absent; setiferous punctation; without dis-

crete neck; gular sutures combined.

Antennae not geniculate; width of antennomeres in-

creasing from second antennomere to apex of antenna;

penultimate antennomere wider than long.

Pronotum wider than long; sides smoothly rounded; lat-

eral margin fine; with dense setiferous punctation; in pos-

terior half with longitudinal medial impression.

Elytra much wider than long; not longer than pronotum;

shoulders widely rounded; divergent from shoulders to

posterior angles; hind wings reduced; sutural striae weak

or absent; with setiferous punctation.

Abdomen elongate; conically narrowed posteriad;

densely pubescent; with microsculpture.

Protibia slightly wider than meso- and metatibia; pro-

coxae slightly elongate; tarsi composed of five tarsomeres.

Aedeagus with broad and stout central lobe; central lobe

nearly straight; not or weakly curved; paramera as long

as or longer than central lobe: spermatheca oval with short

and straight ductus.

Etymology. The specific name is a combination of the

Greek words geo meaning earth or soil, trochus meaning

circlet, and opsis standing for appearance. The name refers

to the similarity to the gmus Allotrochus Fagel, 1955, and

to the soil dwelling life.

Geotrochopsis pubescens n. sp.

Figs 2A, F, 5A, D

Type material. Holotype, male: Peru . Huanuco, Pangua-

na (74°56’W, 9°37’S), rain forest, collected by pitfall trap,

April 1984, leg. M. Verhaagh (UIC).

Paratypes: Mexico. 1 female, Veracruz, Cordoba, Para-

je Nueve Nacimento, tropical evergreen forest, collected

by Berlese method, 7.8.1969, leg. S. & J. Peck (AMNH);

Costa Rica. 1 male, 2 females, Vulcan Arenal, rd. to Are-

nal Observ. Fodge (84°43.58’W, 10°26.5rN), forest bor-

der, litter & dead wood, sifted, 3.12.2012, leg. M. Schiilke

(UIC, MSC); 3 females, Puntarenas, OS A Peninsula, 5km

W Rincon de OSA (83°3rW, 8°42’N), 50 m elevation,

forest floor, collected by Berlese method, 24.-30.3. 1973,

leg. J. Wagner & J. Kethley (AMNH); 1 male, Osa Penn.,

Fundacion Neotrop., 10 km W. Rincon (83°31.30’W,

8°42.30’N), 20 m elevation, collected from forest litter by

berlese, 23.6. 1997, leg. R. Anderson (KNHM); Peru. 1 fe-

male, Rio Tambopata Reseve, 30 km SW Puerto Maldon-

ado (69°16’W, 12°12’S), trop. Moist forest, on fungi Rigi-

doporus microporus, 19.9.-10.10.1984, leg. D.A. Grimal-

di (AMNH); Brazil. 1 male, Sao Paulo, leg. Mraz (NMP).

Diagnosis. The species resembles Geotrochopsis flaveo-

lus n.sp. (see below) in the structure of the aedeagus and

the widely rounded posterior angles of the pronotum. It

can be differentiated from G. flaveolus by the shorter in-

ner lateral lobes and the longer paramera. The pronotum

of G. flaveolus is more strongly arched than that of G pu-

bescens, in particular, in the posterior half

Description. Fength: 1.4 mm. Colouration: yellow, legs

and antennae light yellow.

Head: 0. 19 mm long, 0.29 mm wide; without eyes; tem-

ples behind base of antennae approximately as long as

clypeus; sides of temples parallel; clypeus semicircular;

extremely finely punctate; pubescent; weak netlike mi-

crosculpture; surface moderately shiny; small area at base

of antennae without punctation and microsculpture; sur-

face shiny.

Antennae as long as head and pronotum combined; first

antennomere thick and as long as second and third com-

bined; second antennomere nearly as thick as first, but

shorter; third antennomere narrower than preceding and

following antennomeres; nearly half as long as second an-

tennomere; following antennomeres increasing in width;

penultimate antennomere nearly twice as wide as long.

Pronotum: 0.23 mm long, 0.36 mm wide; widest in mid-

dle; evenly narrowed in smooth curve to anterior and pos-

terior margin; anterior angles obtuse; anterior edge not

margined; posterior angles widely rounded; posterior edge

widely emarginate and distinctly margined; in dorsal as-

pect, lateral margin fine in anterior half, widened in pos-

terior half and continuous in equal width to wide poste-

rior margin; in central posterior half with finely impressed

furrow; punctation deeper and coarser than on head; on

average, interstices between punctures as wide as diam-

eter of punctures; pubescent; netlike microsculpture

deeper than on head; meshes on average wider than di-

ameter of punctures; surface moderately shiny.

Elytra: 0.22 mm long, 0.38 mm wide; widest at poste-

rior angles; shoulders evenly rounded; with slight obtuse

angle, only; sides distinctly margined; posterior edge of

elytra straight; punctation slightly finer than on pronotum;

irregular microsculpture deeper than on pronotum; pubes-

cent; surface matt; less shiny than on pronotum.

Abdomen as densely and finely punctate as elytra; punc-

tures partly granulate; pubescent; deep netlike mi-

crosculpture; surface as matt as surface of elytra.

Aedeagus short and broad; apical lobe shortly curved;

paramera distinctly projecting apical lobe; lateral inner

lobes transverse and only slightly projecting apical lobe.

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A new genus and new species of Neotropical Thoracophorini

111

Fig l-4.Geotrochopsis jlaveolus (1), G. pubescens (2), G. collaris (3), and Geomitopsis amazonensis (4); Aedeagus in ventral and

lateral aspect (A); protibia and tarsi (B); mesotibia and tarsi (C), metatibia and tarsi (D), spermatheca (E), antenna (F); scale bar:

0 . 1 .

Etymology. The specific name pubescens derived from Geotrochopsis collaris n. sp.

the same Latin name and means pubescent. It refers to the Figs 3A-F, 5C, E

finely hairy punctation.

Type material. Holotype, male: Peru. Madre de Dios,

Cuzco Amazonica (69°02.06’W, 12°36.48’S), 300 m el-

evation, secondary forest Wl, F95420, 17.5.1995, leg. D.

Agosti (AMNH).

Bonn zoological Bulletin 64 (2): 109-116

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Ulrich Irmler

Fig. 5. Dorsal aspect of head, pronotum, and elytra of Geotrochopsis pubescens (A), G. flaveolus (B), G collaris (C), lateral

aspect of G pubescens (D), and ventral aspect of G collaris (E); dorsal aspect of head, pronotum, and elytra of Geomitopsis ama-

zonensis (F), and G. campanae (G), G remilleti (H), lateral aspect of Geomitopsis amazonensis (I); scale bar: 0.2 mm.

Bonn zoological Bulletin 64 (2): 109-116

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A new genus and new species of Neotropical Thoracophorini

113

Paratypes: 4 males, 16 females with same data as for

holotype; 2 females, same location as holotype, but

17.5.1994 (AMNH,UIC).

Diagnosis. The species can be easily differentiated from

the other two species by the specific structure of the prono-

tum. The pronotal sides are slightly emarginate in front

of the posterior angles, whereas they are smoothly round-

ed in the two other species. Furthermore, the posterior

edge of the elytra is slightly retreated at suture as trian-

gular emargination and the aedeagus has not the pair of

inner lobes as found in G. pubescens and G flaveolus.

Description. Length: 1.35 mm. Colouration: dark yellow;

antennae and legs lighter yellow.

Head: 0. 19 mm long, 0.26 mm wide; without eyes; tem-

ples as long as semi-circular clypeus; punctation fine; pu-

bescent; on average, interstices between punctures twice

as wide as diameter of punctures; netlike microsculpture

moderately deep; meshes distinctly wider than diameter

of punctures; surface moderately shiny.

Antennae longer than head and pronotum combined;

first antennomere thick and nearly as long as second and

third antennomere combined; second antennomere slight-

ly narrower than first, but thicker than third antennomere;

twice as long as third antennomere; following anten-

nomeres increasing in width; third antennomere approx-

imately square; penultimate antennomere nearly twice as

wide as long.

Pronotum: 0.23 mm long, 0.33 mm wide; widest in mid-

dle; evenly narrowed to anterior angles in smooth curve;

in front of posterior angles with slight emargination; pos-

terior angles obtuse, but nearly rectangular; anterior edge

not margined; lateral margin fine and continuous to pos-

terior edge; margined posterior edge slightly emarginate;

punctation slightly denser and deeper than on head; pu-

bescent; on average, interstices between punctures slight-

ly wider than diameter of punctures; netlike microsculp-

ture as deep and wide as on head; surface moderately

shiny.

Elytra: 0.21 mm long, 0.35 mm wide; widest at poste-

rior angles; narrowed to shoulders in smooth curve; in dor-

sal aspect, lateral margin visible in its total length; shoul-

ders obtuse without forming angles; posterior edge of ely-

tra retreated to suture as wide triangular emargination;

punctation as deep and dense as on pronotum; microsculp-

ture slightly deeper; surface as shiny as on pronotum.

Abdomen as densely and finely punctate as elytra; punc-

tures partly granulate; pubescent; deep netlike mi-

crosculpture; surface as matt as surface of elytra.

Aedeagus slender with long central lobe; apical lobe as

long as basal lobe; nearly straight; in slight obtuse angle

to basal lobe; slender paramera slightly projecting central

lobe; with numerous sensillae; inner lobe projecting.

Etymology. The specific name derived from the Latin

word collum for pronotum and refers to the specific struc-

ture of the posterior angles of the pronotum.

Geotrochopsis flaveolus n. sp.

Figs lA, F, 5B

Type material. Holotype, male: Brazil. Rio de Janeiro,

Moro de Babilonia (43°10’W, 22°57’S), 100-200 m ele-

vation, 25.10.2002, leg. J. Janak (INPA). Paratypes: 172

specimens with same data and from same location as holo-

type (NHMP, JJC, UIC, ZFMK).

Diagnosis. The species is characterised by the arched

pronotum, in particular, in the posterior half In dorsal as-

pect, the lateral margin is covered at posterior angles. Fur-

thermore, the pair of inner lobe are longer than in G pu-

bescens and the paramera are shorter.

Description. Length: 1.6 mm. Colouration: Dark yellow;

posterior margin of pronotum darker, light brownish; legs

and antennae light yellow.

Head: 0.25 mm long, 0.30 mm wide; without eyes; tem-

ples as long as semi-circular clypeus; punctation moder-

ately deep and dense; pubescent; on average, interstices

between punctures twice as wide as diameter of punctures;

netlike microsculpture moderately deep; meshes distinct-

ly wider than diameter of punctures; surface moderately

shiny.

Antennae as long as head and pronotum combined; first

antennomere long and thick; slightly shorter than second

and third antennomeres combined; second antennomere

slightly narrower and shorter than first antennomere; con-

ical third antennomere much narrower and shorter than

second antennomere; following antennomeres increasing

in width; fourth antennomere approximately square;

penultimate antennomere nearly twice as wide as long.

Pronotum: 0.25 mm long, 0.39 mm wide; widest in mid-

dle; narrowed in even and smooth curve to anterior and

posterior angles; posterior angles obtuse; rounded in wide

smooth curve without forming distinct angles; anterior

edge without margin; lateral margin fine; continuous to

posterior edge; in dorsal aspect, margin covered in ante-

rior half and at posterior angles; punctation as dense and

deep as on head; pubescent; netlike microsculpture mod-

erately deep; meshes wider than diameter of punctures;

with short impressed furrow in posterior half of midline;

netlike microsculpture as deep and wide as on head; sur-

face moderately shiny.

Elytra: 0.24 mm long, 0.40 mm wide; widest at poste-

rior angles; slightly narrowed in posterior half; more

strongly narrowed in anterior half; shoulders widely

rounded; posterior edge of elytra straight; in dorsal aspect,

lateral margin visible throughout its total length; contin-

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Ulrich Irmler

ued to shoulders and ending shortly in front of scutellum;

punctation deeper and denser than on pronotum, pubes-

cent; netlike microsculpture slightly deeper than on prono-

tum, but meshes as wide as on pronotum and wider than

diameter of punctures; surface moderately shiny.

Abdomen conically narrowed posteriad; densely punc-

tate and pubescent; punctation still denser than on elytra;

punctures partly granulate; netlike microsculpture dense

and deep.

Aedeagus broad; apical lobe placed in wide obtuse an-

gle to basal lobe; inner lateral lobes distinctly projecting

central lobe; paramera longer than central lobe, but short-

er than central lobe and projecting inner lobes combined;

group of three sensillae in middle of paramera.

Etymology. The specific name derived from the Latin

word 'Jlaveolus' meaning yellowish and refers to the over-

all yellowish colouration of the species.

Key to the species of Geotrochopsis n. gen.

1. Posterior angles of pronotum obtusely rounded;

aedeagus with pair of inner lobes 2

- Posterior angles of pronotum nearly rectangular,

pronotal sides in front of posterior angles slightly sin-

uate (Fig. 5C), aedeagus with one inner lobe (Fig. 3 A)

G. collaris n. sp.

2. Pronotum strongly arched; in dorsal aspect, margin

at posterior angles covered (Fig. 5B), pair of inner

lobes of aedeagus long, paramera not projecting in-

ner lobes (Fig. lA) G. Jlaveolus n. sp

- Pronotum not arched, in dorsal aspect, margin at pos-

terior angles visible (Fig. 5A), pair of inner lobes of

aedeagus short and transverse, paramera projecting in-

ner lobes (Fig. 2 A) G. pubescens n. sp.

Geomitopsis Scheerpeltz, 1931

Ashnaosorius Makhan, 2008: 1, new synonymy.

Libanotyphlus Coiffait, 1954: 155.

Remarks. Makhan (2008) described the new genus Ash-

naosorius on the basis of two new species from Suriname

that were formerly described under the genus Cubanoty-

phlus Coiffait & Decou (1972). He characterised the genus

by tarsi being composed of three tarsomeres and differ-

ences in the aedeagal structure to Geomitopsis Scheerpeltz,

1931. However, he compared his species only with G.

remilleti Orousset, 1985 that was placed in the subgenus

Pseudogeomitopsis Orousset, 1983. Fortunately, he pub-

lished photos of the legs. It can be derived from the pho-

tos that the tarsi are not composed of three tarsomeres, as

mentioned in the description, but composed of four tar-

someres as in the genus Geomitopsis. The first tarsomere

is very short and can be easily overlooked. Furthermore,

all other characters of Geomitopsis, i.e. sixth antennomere

narrower than fifth and seventh antennomeres, absence of

eyes, the specific structure of the aedeagus, and the over-

all habitus as derived from the photos of the original pub-

lication are equal 'm Ashnaosorius and Geomitopsis. Thus,

no generic difference between Ashnaosorius and Geomi-

topsis is found. Therefore, Ashnaosorius is regarded as

synonym to Geomitopsis.

Unfortunately, the type specimens of the species of Ash-

naosorius could not be studied. Requests to loan speci-

mens were not answered.

Geomitopsis amazonensis n. sp.

Figs 4A-F, 5F

Type material. Holotype, male: Brazil, Amazonas,

Reserva Ducke, 26 km NE Manaus, Plot A, leaf litter, Jan.

1996, leg. M.G.V. Barbosa (BMNH)

Paratypes: Brazil . 2 females from the same location as

holotype, but collected on April and Aug. 1995, leg.

M.G.V. Barbosa (BMNH); Peru. 1 female, Huanuco,

Yuypichis, Panguana (74°56.8’W, 9°37’S), manioca field,

22.9.1975, leg. W. Hanagarth (UIC).

Diagnosis. G. amazonensis is characterised by the deep

punctation of the pronotum and the elytra. G. remelleti

Orousset, 1985 has only free larger punctures in a longi-

tudinal row close to the midline. The Chilean species can

be distinguished from the Amazonian species by the

prominent shoulders.

Description. Length: 1.35 mm. Colouration: light brown,

legs and antennae yellow.

Head: 0.17 mm long, 0.22 mm wide; widest shortly in

front of posterior margin; clypeus slightly cheek-like ex-

tended; without distinctly narrower neck; without eyes;

base of antennae to anterior edge; in dorsal aspect; ante-

rior edge even; pair of setiferous punctures between base

of antennae and laterally on posterior vertex; punctation

weak; only few punctures larger; netlike micro sculpture

weak; surface shiny.

Antennae slightly longer than head and half of prono-

tum combined; first antennomere thick; second oval and

slightly narrower than first antennomere; third anten-

nomere conical and slightly shorter than second; follow-

ing antennomeres increasing in width except narrower

sixth and eighth antennomeres; antennomere four to ten

much wider than long; fourth antennomere twice as wide

as long; tenth antennomere slightly wider than twice as

wide as long.

Pronotum: 0.23 mm long, 0.25 mm wide; widest near

middle; sides evenly narrowed to anterior and posterior

angles, but posteriad more strongly narrowed than ante-

Bonn zoological Bulletin 64 (2): 109-116

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A new genus and new species of Neotropical Thoracophorini

115

riad; posterior angles obtuse; lateral margin continued to

posterior edge; anterior edge not margined; punctation

much deeper and coarser than on head; narrow midline

impunctate; on average, interstices between punctures half

as wide as diameter of punctures; in posterior half of mid-

line with oval impression; netlike micro sculpture weak,

but deeper than on head; surface moderately shiny.

Elytra: 0.21 mm long, 0.28 mm wide; widest near pos-

terior edge; narrowed to shortly rounded shoulders; punc-

tation weaker and sparser than on pronotum; on average,

interstices at least as wide as diameter of punctures; two

pairs of large circular impressions close to suture; poste-

rior impression larger than anterior impression; three to

four setiferous punctures in lateral margin; netlike mi-

crosculpture as deep as and as wide as on pronotum; sur-

face moderately shiny.

Abdomen as deeply, but more densely punctate than ely-

tra; netlike microsculpture deeper than on pronotum and

elytra; meshes distinctly wider than diameter of punctures;

surface less shiny than on pronotum.

Aedeagus with thick central lobe; central lobe with long

slender and straight digit ending in acute curved apex; dig-

it at ventral side with row of setae; paramera thick and sin-

uate; several sensillae on ventral and dorsal side.

Etymology. The specific name derived from the Brazil-

ian state Amazonas, where the species was collected.

Key to the species of Geomitopsis in the Neotropical re-

gion

1 . Pronotum distinctly and densely punctate, elytra with

round impressions close to suture (Fig. 5F)

G. amazonensis n. sp.

- Pronotum weakly and sparsely punctate, elytra with-

out impressions 2

2. Shoulders of elytra not carinate, elytra posteriad not

dilated (Fig. 5H) 4

- Shoulders of elytra carinate, elytra posteriad dilated

(Fig. 5G) 3

3. Pronotum with large impressions on the disc

G. campanae Saiz, 1973

- Pronotum with fine elongate impression in the pos-

terior half of the midline

G. chilensis Coiffait & Saiz, 1963

4. Aedeagus at ventral edge of central lobe with trian-

gular prominence G. amrishi (Makhan, 2007)

- Aedeagus at ventral edge of central lobe straight,

without prominence 5

5. Central lobe of aedeagus at ventral edge straight, with

short hook-like apex G rishwani (Makhan, 2007)

- Central lobe of aedeagus at ventral edge evenly

curved to acute apex G remilleti Orousset, 1985

DISCUSSION

The new genus Geotrochopsis must be certainly placed

to the tribe Thoracophorini, as the protibia has no inner

emargination with comb such as in the tribe Osoriini. Re-

garding the united gular sutures, it seems most closely re-

lated to the genus Clavilispinus Bernhauer, 1926. In con-

trast to the other blind genus, i.e. Geomitopsis, of the same

tribe that has four tarsomeres, Geotrochopsis has five tar-

someres. Furthermore, Geomitopsis was placed to the sub-

tribe Glyptomina by Herman (2001), because gular sutures

are separated. Irmler (2010) found that gular sutures can

be separated or united even in one genus, which makes

their constitution unsuitable for a generic differentiation.

Nevertheless, the generic characters, e.g. tarsi composed

of five tarsomeres and united gular sutures let suppose that

Geotrochopsis is closely related to Clavilispinus, although

the overall habitus is very different. Clavilispinus has well

developed eyes, a dorsoventrally depressed body, and no

pubescence, whereas Geotrochopsis is blind with cylin-

drical body and dense pubescence. It might be also relat-

ed to Allotrochus Fagel, 1955 regarding the overall habi-

tus and the slightly elongate procoxae. But, Allotrochus

has well developed eyes, no shortened elytra, and no pu-

bescence.

According to Herman (2001) the genus Geomitopsis is

recorded from Europe, Africa, and South America. Nine

species are loiown from the Mediterranean Europe includ-

ing northern Africa and Near East, six from tropical Africa,

and, together with the newly described species, six species

from the Neotropical region. Whereas at present several

blind genera are known in the tribe Osoriini, only this

genus with blind species was so far known from the tribe

Thoracophorini. The genus is characterised by the absence

of eyes, tarsi composed of four tarsomeres, shortened ely-

tra, a narrow sixth antennomere, and a characteristic struc-

ture of the aedeagus. A similar combination of characters

is found in the genus Arborilispinus Irmler, 2010. How-

ever, Arborilispinus has eyes while eyes are absent in Ge-

omitopsis.

Acknowledgements. I thank the curators of the museums (L.

Herman, R. Brooks, Z. Falin) as well as Jifi Janak for their steady

support and for the loan of the specimens. Heartily thanks are

also to A. Taghavian and J. Orousset from the Museum nation-

al d’Histoire naturelle, Paris, for the loan of G. remilleti and the

delivery of a paratype. I thank also D. Ahrens for the helpful

comments to improve the manuscript.

REFERENCES

Makhan D (2008) Ashnaosorius gen. nov. from Suriname

(Coleoptera: Staphylinidae: Osoriinae). Caloderma Suppl. 70:

1-8

Makhan D (2007) Two new species of Cubanotyphlus from Suri-

name (Coleoptera: Staphylinidae: Leptotyphlinae). Caloder-

ma 9: 5-8

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Ulrich Irmler

Scheerpeltz O (1931) Zoologische Forschungsreise nach den Jo-

nischen Inseln und dem Peleponnes. Sitzungsberichte Akade-

mie der Wissenschaften, Wien 140; 359-460

Orousset J (1985) Deux Osoriinae anophthalmes de Guyane

Frangaise (Coleoptera, Staphylinidae). Revue frangaise Ento-

mologie (N.S.) 7; 60-66

CoifFait FI (1954) Contribution a la connaissance de la faune cav-

emicole et endogee du Liban. (Mission H. Coiffait du Liban,

1951). IF Staphylinides nouveaux. Notes et Memoires sur le

Moyen-Orient 5: 155-162

Coiffait H, Saiz F (1965) Une espece du genre Geomitopsis (Col.

Staphylinidae). Revue Ecologie Biologie du Sol 4: 577-578

Saiz F (1973) Un nouveau Geomitopsis du Chile (Col. Staphy-

linidae). Nouveau Revue entomologique 1; 19-21

Herman E (2001) Catalogue of the Staphylinidae (Insecta;

Coleoptera). 1758 to the end of the second millennium. II.

Oxyteline group. Bulletin of the American Museum of Natu-

ral History 265: 1067-1806

Irmler U (2010) A new genus of Osoriinae in the Neotropical

region with a cladistic analysis of the tribe Thoracophorini (In-

secta: Coleoptera: Staphylinidae). Arthropod Systematics &

Phylogeny 68: 229-237

Bernhauer M (1926) 32. Beitrag zur Staphylinidenfauna Siid-

amerikas insbesondere Brasiliens. Memoire della Societa En-

tomologica Italiana 5; 152-169

Fagel G 0955) Osoriinae. Pp. 1-133 in: de Witte, G.F. (ed.) Ex-

ploration du Parc National due FUpemba. Institut du Parces

nationaux du Congo Beige 39

Bonn zoological Bulletin 64 (2): 109-116

©ZFMK

Bonn zoological Bulletin 64 (2): 117-138

March 2016

A critical review of Hoser’s writings on Draconinae,

Amphibolurinae, Laudakia and Uromastycinae

(Squamata: Agamidae)

Wolfgang Denzer* ^ Ulrich Manthey^ Philipp WagneU^ & Wolfgang Bohme^

^ Society for Southeast Asian Herpetology, Rubensstr. 90, D-12157 Berlin, Germany;

E-mail: wolfdenoxford@yahoo.co.uk

^ Society for Southeast Asian Herpetology, Kindelbergweg 15, D-12249 Berlin, Germany;

E-mail: manthey. sseah@t-online. de

^ Zoologische Staatssammlung Munchen, Munchhausenstr. 21, D-81247 Munchen, Germany;

E-mail: philipp. wagner.zfmk@uni-bonn. de

Department of Biology, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085, USA;

^ Zoologisches Forschungsmuseum Alexander Koenig, Leibniz-lnstitut fur Biodiversitat der Here, Adenauerallee

160, D-53113 Bonn, Germany; E-mail: w.boehme@zfmk.de

^ Corresponding author

Abstract. We analyzed four papers on agamid lizards by self-proclaimed Australian herpetologist Raymond Hoser with

respect to the presentation of diagnostic characters as well as their taxonomic and nomenclatural merits. In most cases

the taxonomic concepts were lifted from earlier phylogenetic publications and the diagnoses were copied from other au-

thors. Copied text in Hoser’s diagnostic section within the analyzed papers amounts to a staggering 83% for Draconinae,

82% for Amphibolurinae, 77% for Laudakia and 78% for Uromastycinae, respectively. We found a number of plagia-

rized paragraphs, sometimes half a page long. Hoser hardly ever makes any effort to attribute statements to the original

author and in some cases he even omitted to cite the relevant source. With respect to nomenclature, we found that Hoser

proposed names that were preoccupied or unavailable, that a nomen oblitum was resurrected incorrectly, nomina nuda

were produced, a type locality was restricted incorrectly and a questionable holotype was designated for a new species.

With respect to taxonomy, we found examples of wrong diagnoses, falsely attributed species, omission of taxa and a lack

of understanding or misinterpretation of previously published taxonomic studies on agamid lizards. Furthermore rele-

vant literature on taxonomy and nomenclature has been overlooked or disregarded.

Key words. Plagiarism, IZCN rules, nomina nuda, questionable type specimen designation, ambiguous diagnoses

INTRODUCTION

For the past few years now the Australasian Journal of

Herpetology (hereafter AJH) has been produced in print

and as an online journal where pdfs can be downloaded.

At the time of writing, 29 issues of the AJH have been

produced. The editor of and sole contributor to the jour-

nal appears to be Raymond Hoser who mainly writes about

reptile classification. These articles are an area of contro-

versy and most herpetologists as well as herpetological

journals and societies worldwide have recorded their ob-

jection to Hoser’s works (see Items for Action & Aclmowl-

edgments in Kaiser 2013); the scientific community cur-

rently appears almost unanimous in their approach not to

use Hoser’s nomenclature.

Albeit that the majority of herpetologists appears to be

in agreement on the suggested suppression of names pro-

posed by Hoser, it has to be noted that this action may not

be in agreement with The International Code of Zoolog-

ical Nomenclature (ICZN, 1999 & 2012; hereafter “the

Received: 11.08.2015

Accepted: 22.03.2016

Code"'), a set of regulation every zoologist is obligated to

follow and should wish to uphold. The Code is served by

the International Commission on Zoological Nomencla-

ture (hereafter, ICZN), which adjudicates instances where

taxon names may lead to confusion, are improperly pre-

sented or formed, or where published works threaten the

stability of the nomenclature in a given discipline. The

service of the ICZN includes a recently developed, for-

mal taxon name registration service in the form of

Zoobank (accessible at zoobank.org), where authors of

taxon names may formally establish a claim to their names

or other nomenclatural acts. Hoser registers all names pro-

posed by him with Zoobank and as a consequence the

names are available in the sense of the Code. However,

it must be noted that the Zoobank website does not have

any provision to prevent the registration of invalid nomen-

clatural acts, thus anyone can register and contribute pre-

sumed valid scientific names. In its current version.

Corresponding editor: F. Herder

118

Wolfgang Denzer et al.

Zoobank can only be considered as provisional until there

are rules implemented that prevent misuse of this data-

bank.

The Code has no provisions for the quality of publica-

tion in which taxonomic and nomenclatural acts are pro-

posed. In particular, there is no need for a journal to have

an editorial board or have a peer review process in place

to validate a proposed name. As has been noted, “the qual-

ity of taxonomic descriptions does not make a name un-

available there being no requirement as such in the

Code..."" (Thomson 2014), i.e. for the ICZN nomencla-

ture and taxonomy are not dependent upon each other. A

proposed taxonomy may be inconsistent, ambiguous or

even false and every herpetologist can choose to follow

it or not, but a proposed taxon name, if produced in ac-

cordance with the Code, becomes available immediately.

There exist only a few prerequisites for a journal to com-

ply with the Code in order to validate and make available

a proposed name. One such prerequisite (ICZN, Article

8) is that the journal is widely available (for example in

public libraries) “providing a public and permanent sci-

entific record” and “numerous identical and durable

copies” have to be assured. Typically 25 copies (Recom-

mendation 8b) constitute a sufficiently available edition.

In order to prove that sufficient copies have been printed

Hoser typically publishes a tax invoice in each issue of

AJH stating that 50 copies were printed. Distribution is,

however, not proven, but presumably at least some copies

are sent to libraries (all issues of the journal can be found

in the National Library of Australia online catalogue) and

distributed among subscribers to the journal. Additional-

ly, all issues or individual articles within a given issue are

presented online as downloadable pdfs a month after the

print version has been in circulation. Every nomenclatur-

al act is registered with Zoobank and hence the proposed

names may be considered published in accordance with

the Code and therefore available for the purposes of

nomenclature.

Editorial boards and high profile referees (reviewers) of

manuscripts are usually a measure for the quality of a jour-

nal and their names may even be published periodically

(e.g. Journal of Herpetology). The AJHdoQS not have an

editorial board to oversee standards of publication or for

undisclosed reasons has decided not to present that infor-

mation in any issues of AJH. However, according to Hoser

(2012: 41) manuscripts submitted to the journal are ref-

ereed by four independent reviewers. This extensive peer

review process should assure that all taxonomic and

nomenclatural decisions presented “[stand] up to the most

robust of scrutiny” (Hoser 2012: 41). Additionally this lev-

el of peer review should provide an assurance that the ar-

ticle adheres to commonly accepted editorial standards,

including ethical considerations such as avoidance of pla-

giarism or the inclusion of derogatory comments.

Plagiarism is generally defined as passing off ideas or

text from other publications as one’s own, whether or not

the source is cited (for definitions see plagiarism.org).

Copying text into one’s own work without citing its source

is the most flagrant form of plagiarism and in many coun-

tries is a violation of intellectual property rights and ille-

gal. Even copying a substantial part of a previous publi-

cation and citing the source is still a form of plagiarism,

if the copied text is not produced within quotation marks

or other means to make the reader aware that the original

research or text is not the work of the current author. Sim-

ilarly, minor modification of the original text such as re-

arrangement of phrases or the substitution of a few words

is still plagiarism, when the original author is not attrib-

uted in an appropriate manner.

Derogatory criticism of other authors in any scientific

publication must be avoided. Providing counterarguments

relating to scientific opinions of a certain author or a group

of authors is a well-established way in science to encour-

age discussion about the matter in question. However, per-

sonal attacks or defamations must be avoided by all means

and are not a part of a scientific (or other) publication.

In the following discussion we will analyze four of

Hoser’s (Hoser 2012a, 2013, 2014b & 2014c) publications

on agamid lizards and discuss our findings in taxonomic

and nomenclatural terms.

MATERIALS & METHODS

The papers were downloaded from the AJH website.

Hoser’s texts were analyzed with respect to their taxonom-

ic and nomenclatural decisions as well as to generally ac-

cepted editorial standards of scientific publications. Pre-

vious publications by other authors containing diagnos-

tic characters and descriptions were compared to the di-

agnoses used by Hoser. Any copied or plagiarized text was

marked and attributed to the original source including page

number. Hoser’s diagnoses do not follow Einnean tele-

graphic style and frequently contain long introductory sen-

tences that do not further the knowledge about a taxon.

We, therefore, accounted for any copied or plagiarized text

identified in Hoser’s diagnoses in two different ways: 1)

as a percentage of the whole diagnosis including introduc-

tory sentences and 2) as a percentage of the presented text

comprising diagnostic characters only. This was done by

accounting for lines of overall text vs. lines of copied text

in a way that favoured any originality in Hoser’s text, i.e.

a line, even if only half printed, was typically counted as

full, while in the case of copied text two half lines were

counted as one. Total lines in the publication about Am-

phibolurinae (Hoser 2013) were counted, those of the oth-

er publications discussed here were estimated as follows:

typically each page in the A J// contained about 140 lines

(70 lines per column). Abstracts and titles were printed in

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Critical review of Hoser’s publications on agamid lizards

119

full lines and the actual number of lines was therefore dou-

bled as if they had been in two columns. In the case of

the other papers the overall sum of lines was not count-

ed but calculated by assuming that each column contains

70 lines.

One of the sources referenced by Hoser (2013) is Cog-

ger (2000). Here we present the results in comparison to

Cogger (1983) in order to show that nearly all of the di-

agnostic characters used for the classification of amphi-

bolurine lizards are considerably older than claimed. Some

diagnostic characters could not be accounted for by com-

parison to earlier publications. Where the source was un-

clear an internet search was performed and if identified

(e.g. Wikipedia, Reptile Database etc.) parts were marked

accordingly. Obviously we do not know precisely which

sources were actually used by Hoser (original description,

review works, catalogues, web pages etc.) and therefore

we relate identified text passages to the publication where

we looked for and found identical phrases. As we cannot

reproduce every single character or paragraph for direct

comparison the respective pages where sets of characters

or a full description can be found are given together with

the number of copied lines and the respective source. At

the end of each section we give a summary of our find-

ings with informations on Hoser’s taxonomical approach

and sources used.

DISCLAIMER

As a general rule Hoser’s new taxon names are not used

in this paper and the respective taxon named by Hoser will

be mentioned as “new tribe / genus to accommodate / con-

tain the following XY” or by a similar phrase where the

placeholders are substituted by currently accepted names.

This is done to prevent accidental validation of Hoser’s

names, which subsequently could become available un-

der the rules of the Code. If, by accident, a new taxon

name proposed by Hoser is used herein that paragraph

shall be treated as not published and the name shall be con-

sidered as not available for the purposes of nomenclature.

This disclaimer is in compliance with Article 8.2 of the

Code.

RESULTS & DISCUSSION

A) Hoser (2014b) on Draconinae

As printed in the header of the paper, the Draconinae man-

uscript was received by the journal on 10 November 2013,

accepted on 1 June 2014 and published on 1 July 2014.

According to the tax invoice. Issue 22 of the AJH, which

includes the Draconinae paper, appears to have been

planned before October 2013, which is the date of the in-

voice (Hoser 2013: 36, Hoser 2014a: 5; invoice date 3 Oc-

tober 2013, several weeks before the publisher initially re-

ceived the manuscript). This could indicate that Hoser pays

in advance for the printing of issues, which would imply

that manuscripts may already be in hand, or that some of

the publication dates are otherwise manipulated.

The paper contains the following sections or headings:

Title, Abstract (including Keywords), Introduction, Unlaw-

ful Theft of Material and Data, and Notes on Taxa Named

Herein, followed by the actual taxonomic and nomenclat-

ural part, a Conflict of Interest section, and a References

Cited section. The publication additionally contains a table

depicting the proposed nomenclature.

The introduction to the paper is mainly concerned with

the phylogenetic and morphological data presented by ear-

lier authors, which serve as the basis for Hoser’s taxonom-

ic and nomenclatural decisions. As in most of his recent

papers, Hoser includes personal criticism of recent and

past herpetologists. Similarly, Hoser directly insults sev-

eral herpetologists in his Unlawful Theft of Material and

Data section of the paper. In this part we are also made

to believe that most of his research files had been confis-

cated and that his ideas were repeatedly used by recent

authors in order to rename taxa and produce junior syn-

onyms.

Overall, in this paper Hoser describes one new species,

proposes eight new genera, resurrects three names for sub-

genera, and erects 22 subgenera, ten new tribes and six

subtribes. His diagnosis of the genus Lyriocephalus Mer-

rem, 1820 may serve as an example how he defines a

genus and how we analyzed his statements. The follow-

ing is a true copy from Hoser (2014a: 38):

“Lyriocephalus Merrem, 1 820 is defined by the follow-

ing suite of characters: Mouth large; teeth erect in both

jaws. Incisors small and conical. No praeanal or femoral

pores (as opposed to the callous pore-like swelling of the

preanal scales of the males in the genera Agama Daudin,

1802, UromastixMQrrQm, \^20 andXenagamaBoulQngQY,

1895); tympanum hidden. Five toes. A dorsal crest; a V-

shaped gular fold; a bony supraorbital arch. Body com-

pressed, covered with small scales intermixed with en-

larged ones. A nuchal and a dorsal crest. A gular sac and

a V-shaped gular fold. Adult with a globular hump on the

nose. Pre and post-orbital bones forming an arch limiting

a supraorbital fossa.”

The first set of characters “Mouth large . . . arch” is a

copy from Boulenger’s synopsis leading to Lyriocephalus

(Boulenger, 1885: 251-252). The part in brackets “callous

. . . genera” is taken from a footnote in Boulenger (1885:

251), where it only refers to Agama and Aporoscelis [=

Xenagama]. The part containing Uromastix [sic] andXe-

nagama could not be identified, but is presumably taken

from another comparatively old publication as the genus

name Uromastyx is written in its historically used form.

The second set of characters “Body ... fossa” mirrors

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120

Wolfgang Denzer et al.

Boulenger’s (1885: 281) diagnosis of the genus. It is quite

obvious that copying has been done without giving it much

further consideration. The V-shaped gular fold appears

twice as does the dorsal crest. The characters “supraor-

bital arch” and “supraorbital fossa” are repeated without

comment; when a supraorbital arch is formed, this leads

to a supraorbital fossa between the arch and the dorsal out-

er ridge of the eye socket. Hoser cites Moody (1980) in

his bibliography. Had he looked at this publication he

would have found that as a matter of fact the supraorbital

arch in Lyriocephalus is formed by prefrontal and postor-

bital and not as claimed by Hoser (2014a: 38) by “pre and

post-orbital bones”.

Most taxonomic concepts proposed by Hoser have been

published by earlier authors (and cannot be repeated here

in full for comparison) but without taking the step of as-

signing genus names to species groups (e.g., Gono-

cephalus Kaup, 1825; Draco Linnaeus, 1758; Japalura

Gray, 1853) or to species where only insufficient materi-

al and/or data exist. In the following we will first provide

evidence that the taxonomic scheme proposed by Hoser

is either based on previously published concepts or con-

stitutes mere naming of more or less supported nodes in

phylogenetic publications concerned with Draconinae. In

the second part we will have a closer look at the diagnoses

of genera and compare those to previously published ma-

terial. We will discuss each group in the same sequence

as published by Hoser. In our analysis below we will not

discuss all of Hoser’s diagnoses in such great detail as the

one of Lyriocephalus and only point out inconsistencies

in taxonomy and nomenclature where we feel it should

be done for clarity.

The first genus Hoser deals with is Gonocephalus which

he proposes to divide into five subgenera along with the

erection of two new genera. His subgeneric classification

follows the species group assignment proposed by Man-

they & Denzer (1991) and Denzer & Manthey (2009, part).

Denzer & Manthey (l.c.) combined the Philippine species

with their bornensislbellii species group, which they had

considered a separate species group in the earlier publi-

cation, based on morphological similarities. Hoser elevates

two species to genus rank, namely G. robinsonii

(Boulenger, 1908) and G. mjobergi Smith, 1925. This had

already been suggested by Manthey (2010) where G

robinsonii was treated as (Gonocephalus incertae sedis)

robinsonii and by Denzer & Manthey (l.c.) where it was

suggested that G. mjobergi should be referred to as Genus

A within a Gonocephalus s. 1. complex. Owing to insuf-

ficient material (only a single female specimen has ever

been collected) Denzer & Manthey (l.c.) abstained from

proposing a genus name for G mjobergi until more ma-

terial will become available. They further stated that one

autapomorphic character in particular (longitudinal gular

folds) constituted a synapomorphy for the genus group G

mjobergi, Mantheyus Armyowdi & Stuart, 2001 and Ptyc-

tolaemus Peters, 1 864. With respect to G robinsonii Hoser

states in his introductory part to the genus Gonocephalus

that “no one has bothered to assign the taxon Gono-

cephalus robinsonii ... to a genus of its own”. As will be

discussed below Hoser has taken on this task but fails to

deliver as not a single of his characters is of any value to

diagnose his newly proposed genus (i.e., differentiate from

other genera or species groups).

Initially Hoser characterizes the genus Gonocephalus.

His diagnosis comprises seven lines and is copied from

Boulenger (1885: 282, 3.5 lines) and Denzer & Manthey

(2009: 255-256, 3.5 lines). His subgenus to accommodate

the chamaeleontinus group as defined by Manthey & Den-

zer (1991) is characterized by a single character copied

from Boulenger (1885) and separated from other proposed

subgenera by comparison in a way that their full diagnoses

are repeated. Hoser considers the chamaeleontinus group

as the nominate form. Next Hoser proposes a subgenus

to accommodate G grandis (Gray, 1845). For this

species he resurrects an available name proposed by Gray

(1845). The diagnostic character section amounts to ap-

proximately 25 lines, all of which are a copy of

Boulenger’s (1885: 298) description of the species. This

is followed by proposing a new subgenus for the Philip-

pine species group by using three characters (two lines)

taken from Boulenger (1885). The next new subgenus

comprises the bornensis group. The diagnostic characters

are mainly taken from Boulenger (1885) but rearranged

and slightly modified without copying directly. The name

he gives the subgenus is different from the name he uses

in the keywords to the paper, the latter of which therefore

becomes a nomen nudum. The last new subgenus proposed

contains the Sumatran megalepis species group and is

characterized initially by two lines copied from

Boulenger’s (1885) synopsis [key] to the genus followed

by Boulenger’s (1885: 291) full description of G tuber-

culatus (= G megalepis). This last part comprises 24

copied lines and ends with citing Boulenger (1885). How-

ever, Hoser does not make clear that the whole descrip-

tion is copied by, for example, using quotation marks.

Gonocephalus robinsonii is removed from its synonymy

with the genus and a new genus is proposed. This new

genus is diagnosed by three characters: karyotype, a

“greatly enlarged gular fold” and “a distinctive white low-

er jaw”. The karyotype section is a copy from Diong et

al. (2000: 74, 6 lines); the other two characters are sup-

posedly based on Hoser’s own research. We would like

to note that the karyotype can even vary within a species

(e.g., see Ota, 1988 for data on Japalura swinhonis Guen-

ther, 1 864), the enlarged gular fold is a false character as

G robinsonii is the only Gonocephalus species without

a gular fold, if one considers G mjobergi as not congener-

ic and the colour of the lower jaw constitutes a variable

character in G robinsonii which is dependant on age (see

photographs in Manthey 2010).

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121

Gonocephalus mjobergi is accommodated in a new

genus. This is done by copying the full description from

Denzer & Manthey (2009, 40 lines) including the above-

mentioned paragraph about the autapomorphy of longi-

tudinal gular folds. Despite using the complete character

set including that a large gular sac “partially conceals the

Gonocephalus-typQ typical gular fold” Hoser earlier

claims that G. robinsonii and G mjobergi have an “en-

larged gular fold.” Additionally this quote shows that

Hoser apparently intended to amend the original statement

but ended up doubling an adjective. The original phrase

reads: “partially conceals the Gonocephalus gular

fold” (Manthey & Denzer 2009: 257).

Hoser often uses brackets for the author of a taxon and

the year of description where he seems to interpret the

Code in his own way (e.g. he uses brackets for Gono-

cephalus robinsonii, Boulenger, 1908, G beyschlagi

Boettger, 1892, G doriae Peters, 1871). The use of brack-

ets for the author/year of a taxon is determined by the

ICZN rules (Article 51.3 [Use of parentheses], see ICZN

Code for an example). The Code prescribes brackets if the

allocation of a species changes with respect to a genus.

This is not the case here. Boulenger and the other authors

decided that the correct spelling should be Gonyocephalus

(an emendation introduced by Wagler [1830]) albeit that

Kaup originally used Gonocephalus and later amended it

to Goniocephalus, but the latter emendation and Gony-

ocephalus are not available under ICZN rules as the orig-

inal name has to be preserved.

The genus Japalura Gray, 1853 is broken up into three

genera, two of them divided additionally into two subgen-

era each. Japalura has for a long time been a matter of

taxonomic changes and only in recent years are we be-

ginning to understand their phylogenetic relationships. A

division into three genera can be derived from molecular

phylogenetic analyses, where results indicate that the

clades containing J. variegata Gray, 1 853 / J. tricarina-

ta (Blyth, 1853), J. polygonata (Hallowell, 1861), and J.

splendida Barbour & Dunn, 1919 / J. flaviceps Barbour

& Dunn, 1919 are only very remotely related (e.g., Py-

ron et al. 2013). Stuart-Fox & Owens (2003) considered

Japalura as comprised of “two widely divergent goups,”

named in their analysis as Japalura India / J. variegata-

group and Japalura SE Asia / J. splendida-growp (SE for

Southeast). They even mention that they consider both as

separate genera. Mahony (2009: 55) refers to the latter

species group as “eastern elade”. In an earlier publication

by Macey et al. (2000) they are referred to as Himalayan

and East Asian clades, respectively. Kastle & Schleich

(1998) proposed that the species of the Western clade with

a visible tympanum should be regarded as a separate

genus, for which the name Oriotiaris Gunther, 1 864 was

available. Hoser mostly follows these previously published

results to propose his taxonomic scheme.

Firstly he deals with Japalura species of the nominate

genus. Here he seems to accept Mahony’s view (2009) that

Japalura and Oriotiaris are congeneric. Japalura varie-

gata (type species of Japalura) and J. tricarinata (type

species of Oriotiaris) are phylogenetically sufficiently

close (Pyron et al, l.c., papers cited in Mahony, l.c.) that

Mahony (2009) already suggested to synonymize both

genera and treat Oriotiaris (resurrected by Kastle & Schle-

ich (1998)) as a Junior synonym of Japalura. Hoser treats

both as subgenera of Japalura.

The nominate genus Japalura is diagnosed in seven

lines which are copied from Boulenger (1885: 307) and

Mahony (2010: 4, definition of Japalura s.l.) with approx-

imately half of the text from each author. The genus is fur-

ther divided into a nominate subgenus and by resurrect-

ing an available name for the second subgenus. Hoser first

defines Oriotiaris. His diagnostic characters for this sub-

genus are copied from Gunther (1864: 1 50, five lines) and

Mahony (2009: 56, five lines). No other characters are giv-

en. In the case of one character taken from Mahony (2009)

Hoser even copies a typographic error, “. . .possession of

a small gular pouch in the later” [sic!].

The subgenus Japalura is diagnosed as follows: “The

diagnosis for the nominate subgenus Japalura is simply

a reversal of the diagnosis for Oriotiaris.'' Hoser distin-

guishes Japalura from his subgenus Oriotiaris as follows:

''Oriotiaris is further separated from the nominate sub-

genus Japalura by the absence (vs. presence) of dorsal

chevrons and presence (vs. absence) of a coloured gular

region, concealed tympanum, large crest spines in males

and erectile nuchal crest (roach), in members of Japalu-

ra." Japalura tricarinata is highly variable and eapable

of changing colour. There exist photographs of complete-

ly green individuals without any chevron pattern (see for

example Manthey 2010: 98, Fig RA02806-4). On the oth-

er hand, J. planidorsata Jerdon, 1870 does not have an

erectile nuchal crest nor does J. sagittifera Smith, 1 940

both of which are placed by Hoser in the nominate sub-

genus.

For Japalura polygonata Hoser resurrects its original

name Diploderma polygonatum Hallowell, 1861. Phylo-

genetic studies showed that J. polygonata is only remote-

ly related to other Japalura but seems to be the sister tax-

on of Gonocephalus robinsonii (Pyron et al. 2013). Diplo-

derma polygonatum was already suggested by Mahony

(2009: 55) in case the eastern clade (see below) should

turn out to be monophyletic. The genus is diagnosed with

four lines, all of which are copied from Boulenger (1885:

307).

Having already dealt with the variegata group, Hoser

proceeds to define a new genus for the eastern species

group, which he splits into two subgenera. The new genus

to accommodate all East Asian species is defined

in the space of approximately nine lines, seven of which

are directly copied from Boulenger (1885: 307-308). One

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Wolfgang Denzer et al.

character is the negation of a Boulenger character and the

rest are slightly amended but not identically copied from

Mahony (2009). No new characters are introduced by

Hoser. This genus is split into two subgenera on the ba-

sis of Boulenger’s synopsis (1885:308) the differences be-

ing the length of the tibia and presence or absence of a

longitudinal fold. For J. swinhonis, Hoser claims that the

“tibia is as long as the skull”. Already Stejneger (1907:

1 83) pointed out that in two old males he had studied “the

tibia is decidedly shorter than the skull”. With respect to

the second character we would like to note that J. chapaen-

sis Bourret, 1937, J. fasciata Mertens, 1926, J. grahami

(Stejneger, 1924) and J. micangshanensis Song, 1987 do

not have a longitudinal fold as claimed by Hoser. They

would therefore have to be transferred to his first sub-

genus.

The genus Calotes Daudin, 1 802 is divided into three

genera, of which two are further divided into subgenera

(the nominate genus into five and a newly proposed genus

into four subgenera). The basis for this taxonomic

scheme appears to be the result of the extensive molecu-

lar biological studies presented by Zug et al. (2006) and

Pyron et al. (2013). The proposed scheme clearly reflects

the nodes in previously published phylogenetic trees. A

division into three groups was already proposed by Smith

(1935) who differentiated between a C. versicolor group,

a C. liocephalus group and a group comprising C. ronxi

Dumeril & Bibron, 1837 and C. ellioti Gunther, 1864. The

first and last of Smith’s groups are elevated to genus lev-

el by Hoser, the liocephalus group is considered by Hoser

as a subgenus.

Initially the genus Calotes is diagnosed by a copy of

Boulenger’s diagnosis (1885:314, four lines) and subse-

quently compared to his newly erected genera (see below).

The first subgenus described within Calotes serves to ac-

commodate C. calotes (Linnaeus, 1758) and C. htunwini

Zug & Vindum, 2006. Their close relationship was dis-

covered in phylogenetic studies despite the fact that their

distribution is rather disjunct. The nominate subgenus is

defined by Hoser by initially repeating Boulenger’s key

(1885: 315-316, 3.5 lines) leading to C. ophiomachus (=

C. calotes) and subsequently by a complete copy of

Boulenger’s description of the species (Boulenger

1885:327, approximately 18 lines). By stating that all of

these characters define the subgenus Hoser renders his di-

agnosis false. C. htunwini does not have a nuchal crest

where the height “equals or exceeds the diameter of the

orbif ’ nor does it have a “dorso-nuchal crest composed

of closely set lanceolate spines” nor is this species green

above. Additionally we like to note that already

Boulenger’s description contains a mistake in stating that

in C. calotes a “gular sac is not developed”. This has been

copied by Hoser; however, male C. calotes actually have

a reasonably well developed gular sac during the breed-

ing season as have C htunwini but to a lesser extent.

Next Hoser proposes a new subgenus containing

species allied to Calotes versicolor (Daudin, 1802) by

copying 1.5 lines from Boulenger’s synopsis (Boulenger

1885: 314-315) followed by an entirely copied descrip-

tion of C versicolor from Boulenger (1885: 312, 20 lines).

Here also Hoser repeats Boulenger’s statement that in C.

versicolor the “gular pouch [is] not developed” which is

not true for male specimens during the breeding season

(Smith 1935; numerous photographs on the internet).

Hoser does not present any new characters for the C. ver-

sicolor group. Subsequently Hoser erects a new subgenus

containing two closely related species from the Western

Ghats, namely C. nemoricola Jerdon, 1853 and C. gran-

disquamis Gunther, 1875. To diagnose the genus he ini-

tially copies four lines from Boulenger’s synopsis (1885:

315) leading to these species followed by the reproduc-

tion of Boulenger’s description (1885: 326) of C. nemori-

cola (approximately 20 lines). No additional or new char-

acters are presented by Hoser.

Species allied to Calotes liolepis Boulenger, 1885 (C.

nigrilabris Peters, 1860 and C. desilvai Bahir &

Maduwage, 2005) are the content of a subgenus that is ini-

tially defined by repeating in full Hallermann’s key (2000:

161-162) leading to C. nigrilabris and C. liolepis, respec-

tively (3.5 lines each), followed by a copy of Boulenger’s

descriptions (1885: 327-328) of C. nigrilabris (approxi-

mately 22 lines) and C. liolepis (approximately 15 lines).

No other characters are presented in the diagnosis.

Species related to Calotes liocephalus Gunther, 1 872 are

placed by Hoser into a new subgenus which again is de-

fined by the characters given in Hallermann’s key (2000)

here for C. liocephalus and C. ceylonensis Muller, 1887

(3.5 lines each) followed by the respective descriptions

copied from Boulenger (1885: 329) for C. liocephalus (18

lines) and Boulenger (1890: 139-140) for C. ceylonensis

(13 lines) without presenting any further characters.

The last subgenus within Calotes proposed by Hoser is

monotypic and contains only C. aurantolabium Krishnan,

2008. Diagnostic characters are given in the space of 13

lines all of which are a copy of Krishnan (2008).

After having dealt with the species he considers

Calotes sensu stricto. Hoser proceeds to erect a new genus

to accommodate species related to C. mystaceus Dumeril

& Bibron, 1837. This genus is further divided into four

subgenera. The definition of the genus comprises approx-

imately 13 lines, which are a copy from Boulenger (1885:

315) or may partially have been taken from Hallermann

(2000: 162). Initially Hoser gives a short diagnosis for the

genus (2.5 lines) followed by the sentence: “In addition

to this, each of the relevant subgenera are further diag-

nosed and separated from the other genera by one or oth-

er of: A/ [diagnosis subgenus A] or B/ [diagnosis subgenus

B] or C/ [diagnosis subgenus C]”. This is followed by sep-

arating the genus from Calotes and another genus contain-

ing C. rouxii Dumeril & Bibron, 1 837 and C. ellioti Giin-

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Critical review of Hoser’s publications on agamid lizards

123

ther, 1864. The presentation of his diagnoses for the sub-

genera here is peculiar if not unique: the diagnostic char-

acters including comparisons presented for the nominate

subgenus and two other subgenera are absolutely identi-

cal to that of the genus!

The last subgenus is monotypic and erected for Calotes

nigriplicatus Hallermann, 2000. Here he repeats the full

description as given by Hallermann (2000: 156, 158, ap-

proximately 30 lines) only adapted in places where a com-

parison is made to one of his newly erected subgenera (i.e.

the name Calotes is replaced by Hoser’s new name). This

is followed by repeating again his diagnostic characters

for the already defined subgenera and genera. In the space

of two pages he uses the same 22 lines five times. In all

of this Hoser does not present a single new character.

Next he defines a new genus to accommodate Calotes

rouxii and C. ellioti. The diagnosis comprises two lines

and is copied from Hallermann’s key (2000: 162).

The last genus Hoser proposes is again monotypic and

only contains what he calls Calotes andamanensis, cur-

rently considered as Pseudocalotes andamanensis

(Boulenger, 1891). While Harikrishnan & Vasudevan

(2013: 11) state: “.. .these differences are not sufficiently

pronounced to justify the recognition of a new genus. In

the absence of a molecular phylogeny and based on ex-

ternal morphology alone, it is most appropriate to consid-

er this species as a member of Pseudocalotes...'' Hoser

opposes this by writing “is also sufficiently divergent to

warrant being placed in a separate genus”. Hoser’s initial

diagnosis is a complete copy (31 lines) from Krishnan’s

description (2008: 533) of the species, only substituted

with Hoser’s nomenclature in places where Krishnan made

comparisons with Calotes. This is followed by the descrip-

tion of Pseudocalotes andamanensis (14 lines) given by

Harikrishnan & Vasudevan (2013: 11) and subsequently

by yet another short description of this species including

comparisons with Calotes Daudin, 1802, Bronchocela

Kaup, 1827, Complicitus Manthey in Manthey and

Grossmann, 1997, Salea Gray, 1845, and Dendragama

Doria, 1888 (17 lines) as produced on the Reptile Data-

base website (original publication not identified). We note

that also the first two descriptions are available on the Rep-

tile Database website. Hence Hoser could have copied the

whole diagnosis from there without even consulting the

original publications. This assumption is viable as Harikr-

ishnan & Vasudevan (l.c.) are cited in an identical place

to that on the website and Krishan’s description stays with-

out a citation as this is also the case on the website. Al-

together he “describes” the species three times in 65 lines

of which 62 lines are copied from other sources and the

remaining lines are introductory sentences.

The genus Ceratophora Gray, 1835 is divided into three

genera including two subgenera reflecting the molecular

and morphological (rostral horn appendage) phylogeny of

Schulte et al. (2002). The nominate genus contains the

species related to C. stoddartii Gray, 1 834 which is divid-

ed subsequently into two subgenera. Hoser’s description

of the nominate genus is presented in 6.5 lines all of which

are taken from Boulenger (1885: 277) with only minor

changes. This is followed by a separation from his other

proposed subgenus and the other two proposed genera (12

lines). The complete text to describe the diagnostic char-

acter is copied from Boulenger (1885: 277) and Pethiyago-

da & Manamendra-Arachchi (1998: 1,4). The definition

of his subgenus to accommodate C tennentii Gunther,

1861 comprises approximately four lines all of which are

taken from Boulenger’s synopsis (1885: 277). The nom-

inate subgenus is defined by four lines again from

Boulenger (1885: 277).

Next Hoser erects a new genus for Ceratophora aspera

Gunther, 1 864, which is initially defined by two lines from

Boulenger (1885: 277) followed by characters taken from

Pethiyagoda & Manamendra-Arachchi (1998: 44, 46, six

lines, all copied) to separate it from the other proposed

genera by Hoser. Even the distributional data are copied

verbatim from Pethiyagoda & Manamendra-Arachchi

(1998: 44).

The last genus Hoser proposes for this group of lizards

only contains Ceratophora karu Pethiyagoda & Mana-

mendra-Arachchi, 1998. This is presented including

comparisons within approximately eight lines, all of which

are a copy from Pethiyagoda & Manamendra-Arachchi

(1998: 44) and can partially be found in an identical way

on the Reptile Database website.

Next Hoser deals with the lizards of the genus Bron-

chocela Kaup, 1827. He initially gives an introduction

where he seems to restrict the type locality of B.

cristatella (Kuhl, 1820) and to resurrect B. moluccana

(Lesson, 1830) (see discussion below). The genus is di-

vided into two subgenera the first of which contains B.ju-

bata (Dumeril & Bibron, 1 837) and B. orlovi Hallermann,

2004. The first three lines of the diagnosis are taken from

Boulenger (1885: 314 all copied) and a full description

(approximately 20 lines) of B. jubata is presented by a

copy of Hallermann’s description (2005: 171-172). Two

more lines of characters concerning the scales at the base

of the dorsal crest could have been taken from de Rooij

(1915: 123). One character cannot be retraced to earlier

publications and presumably comes from Hoser’s re-

search: “The dorsal crest gives the appearance as if it is

composed of tiny hairs as opposed to scales (as seen in

Bronchocela)" [sensu Hoser]. We note that adult males of

B. jubata have one of the most developed dorsal crests

among Bronchocela, consisting of lanceolate scales.

The only new species described by Hoser within the

Draconinae paper is a member of Bronchocela Kaup, 1 827

and refers to material collected on Halmahera Island,

Maluku Province, Indonesia. His description of this

species is purely based on colouration and an elongated

scale between the nasal and the rostral. We note that most

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Wolfgang Denzer et al.

- if not all - Bronchocela species are capable of extreme

colour changes. A typically brightly green coloured B.

cristatella (Kuhl, 1820) may become completely black

when disturbed or during copulation (WD pers. obs.).

Hoser’s choice of holotype (USNM 237431) is - to put

it mildly - slightly confusing. The specimen he chose ac-

tually has a bifurcated tail something that should have been

noted in the diagnosis (see collections. si.edu/search/re-

sults.htm?q=record_ID:nmnhvz_6091296). We further

note that the gender of Bronchocela is female but Hoser

creates a species name with a masculine ending. The de-

scription of his nQ^N Bronchocela species contains copied

sections from Boulenger (1885: 314, 316-317, approxi-

mately 26 lines) for B. cristatella.

In his comparison of the new species to other species

of the genus Bronchocela the author also often refers to

B. moluccana (Lesson, 1 830) which is currently consid-

ered a synonym of B. cristatella (Kuhl, 1820). Interest-

ingly he does not include B. moluccana in his species list

(table at the end of his taxonomic section) although it is

stated in his introduction to the genus that he regards B.

moluccana “as being a separate species”. We note that the

original name given would be Agama moluccana Lesson,

1 830 and the combination B. moluccana was only used

by Peters (1867 as Bronchocele), Stoliczka (1870) and Pe-

ters & Doria (1878), all of which were in later publica-

tions considered to be B. cristatella. Theobald (1876) used

the name B. moluccana in his Reptiles of British India for

a specimen from the Nicobars as a synonym of

Pseudocalotes archiducissae Fitzinger, 1 860, which again

turns out to be a synonym of B. cristatella. Bronchocela

moluccana constitutes a nomen oblitum and resurrection

should have been made clear with reference to the type

species and holotype.

Furthermore Hoser refers several times to Java as the

type locality for B. cristatella. In his original description

Kuhl (1820) never mentions a type locality and ever since

it has been unknown and never been restricted by any au-

thor (see for example Diong & Lim, 1998). One could ar-

gue that Hoser’s statement “West Java (herein treated as

terra typica)” is meant to be the newly defined type lo-

cality. This is an unfortunate choice under current condi-

tions, as the actual phylogenetic status of the Javanese pop-

ulations still needs further research as also pointed out by

Hoser. Additionally Hoser does not refer to a particular

specimen from his type locality and hence the restriction

is not valid.

The genus Phoxophrys Hubrecht, 1881 is divided into

three subgenera. In the introduction to the genus Hoser

claims that “as there has never been a definition or diag-

nosis of Pelturagonia” Mocquard, 1 890 he will “provide

one herein for the first time”. Hoser’s diagnosis only com-

prises two characters while that of Mocquard (1890) is

written in French, and longer with several characters. To

diagnose the genus Hoser uses approximately 20 lines, all

of which are copied from Inger (1960: 221) and include

a comparison to Japalura, the genus several Phoxophrys

species belonged to until Inger’s revision.

Hoser’s nominate subgenus is defined by a minimally

rephrased diagnosis of Phoxophrys tuberculata Hubrecht,

1881 again taken from Inger (1960: 225, seven lines). The

diagnosis of the subgenus to accommodate P. cephalum

(Mocquard, 1 890) only comprises two lines with two char-

acters (“presence of nuchal crest . . . and an absence of a

supraciliary spine”). The last subgenus only contains P.

spiniceps Smith, 1925. This is defined within seven lines,

all copied but slightly rearranged from Inger (1960:

224-225).

The next genus Hoser is concerned with comprises the

lizards of the genus Aphaniotis Peters, 1864. The genus

is divided into two subgenera on the basis of whether a

“protrusion on the snouf ’ is present or absent. The genus

and nominate subgenus diagnoses are identical and each

constitute a copy from Boulenger (1885: 274, four lines).

The other subgenus is defined by approximately four ad-

ditional lines that have been copied from the internet

(www.ecologyasia.com) or a source that we have not iden-

tified.

The genus Ptyctolaemus Peters, 1 864 currently consists

of two species, which Hoser considers to be two subgen-

era. The nominate subgenus containing P. gularis Peter,

1864 is initially defined within 15 lines copied from

Schulte et al. (2004: 230) followed by a comparison to P.

collicristatus Schulte & Vindum, 2004 (Schulte et al.

2004) taken from the same source (five lines). The defi-

nition of the subgenus for P. collicristatus is precisely the

other way round, i.e Hoser first uses the same five lines

from the comparison betwee P. gularis and P. collicrista-

tus to define the species and then the definition of the

genus (all copied from Schulte et al. 2004: 230).

The genus Salea Gray, 1 845 is currently considered to

contain two species (see below) and one highly question-

able species {S. gularis Blyth, 1854). In his introduction

to the genus Hoser states that “neither the genus ... or the

subgenus being properly defined to date . . . this is done

herein for the first time”. He does however not present a

single character to do so that has not been the result of a

copying process from Boulenger (1 885). Hoser breaks up

the genus into two subgenera based on the respective de-

scriptions of S. horsfieldii Gray, 1835 and S. anamallayana

(Beddome, 1878) taken from Boulenger (1885: 251-252,

312-314) with 36 lines (annotated as “modified from

Boulenger” but actually constituting a verbatim copy) and

22 lines, respectively. For the latter species he resurrects

its original name proposed by Beddome (1878).

The last genus Hoser deals with in this part of the pa-

per is Draco Linnaeus, 1758 which has been a matter of

intensive morphological studies in the 1980s by Inger

(1983) and Musters (1983). In recent years phylogenetic

studies by McGuire & Alcala (2000), McGuire & Kiew

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Critical review of Hoser’s publications on agamid lizards

125

(2001) and McGuire et al. (2007) completed the picture.

Hoser mainly uses these phylogenetic results and the tree

of Pyron et al. (2013) to divide the genus into nine sub-

genera and copies their respective diagnoses from

Boulenger (1885) or the morphology based publications

mentioned before. Not a single new character is introduced

by Hoser. Hoser’s general description of the genus is giv-

en in 4.5 lines all taken from Boulenger (1885: 253). On-

ly the phrase “much-produced” is replaced by “much-ex-

panded”.

The first new subgenus Hoser proposes serves to accom-

modate members of the Draco lineatus group (minus D.

lineatus which is placed in its own monotypic subgenus,

see below). Initially Hoser copies seven lines from

McGuire et al. (2007: 181) to define the group including

a statement related to a statistical analysis. However, Hoser

does neither use nor refer to a statistical method in his sec-

tion on methods. Subsequently he produces four lines form

the same source to define his subgenus further (McGuire

et al, 2007: 181) followed by a short description of D. bi-

maculatus Gunther, 1864 taken from Muster (1983: 40)

to distinguish this species from his subgenus. The last part

of Hoser’s diagnosis serves to separate D. lineatus Daudin,

1 802 from his proposed subgenus of the remaining linea-

tus group species. This is done by copying the diagnosis

comprising ten characters provided by McGuire et al.

(2007: 199). At the end of this paragraph Hoser annotates

“adapted from McGuire et al. (2007)” although he actu-

ally produces a complete verbatim copy from that source.

This goes so far that Hoser even has the typographical er-

ror “posnuchal” [sic!] in the same place.

The new monotypic subgenus to accommodate Draco

bimaculatus initially repeats the four lines taken from

Musters (s. above) followed by a copy (ten lines) from

McGuire et al. (l.c.) as given under the previously defined

subgenus. Next Hoser uses again the “adapted” diagno-

sis for Z). lineatus provided by McGuire et al. (2007: 199,

16 lines including typographical error, see above) and fi-

nally describes the species by copying Boulenger

(1885:263, 19 lines) which again is annotated as having

been “adapted” albeit constituting a word-for-word copy.

Draco modiglianii Vinciguerra, 1 892 is placed by Hoser

into its own new subgenus on the basis of a short diag-

nosis (3.5 lines) that has been copied from Musters (1983:

45).

Species related to Draco blanfordii Blanford, 1878 are

combined in yet another new subgenus which he defines

by copying three sets of characters originally from

Boulenger (1885: 255, synopsis to the species, approxi-

mately nine lines). No other characters are presented.

Species related to Draco maculatus (Gray, 1845) are

contained in a new subgenus that is entirely defined by

18 lines coming from Boulenger (1885: 262). The nom-

inate subgenus is diagnosed in approximately three lines

copied from Inger (1983: 17).

Then Hoser defines Draco lineatus Daudin, 1802 in

pretty much the same way he did to diagnose the linea-

group (s. above). Initially he uses McGuire et al. (l.c.)

to define the lineatus-group (approximately nine lines

copied); this is followed by separating D. bimaculatus

from that group and the proposed subgenus by copying

Musters (1983: 40, 4.5 lines). Finally Hoser reproduces

the full set of characters as given by McGuire et al. (2007:

199) for the species annotated as adapted but actually

copied. For this subgenus Hoser resurrects an old avail-

able name from Fitzinger (1843).

Species related to Draco fimbriatus Kuhl, 1820 are

placed into a subgenus for which another name proposed

by Fitzinger (l.c.) is resurrected. The subgenus is defined

in approximately three lines and subsequently separated

from D. maculatus (again three lines), all copied from

Boulenger (1885: 254-255).

The last subgenus Hoser erects serves to accommodate

the Indian species Draco dussumieri Dumeril & Bibron,

1837. To name the subgenus Hoser resurrects another of

Fitzinger’s names (l.c.). The diagnosis consists of four

lines taken from Boulenger ’s synopsis (1885: 255) fol-

lowed by approximately 17 lines of description copied

from the same source (Boulenger 1885: 268).

After having defined his genera and subgenera Hoser

endeavours to divide the subfamily into tribes and sub-

tribes. Hoser proposes ten tribes and six subtribes, which

will be numbered numerically in the following in order

to prevent accidental validation; genus names are given

here in their currently accepted form.

Tribe 1 only contains lizards of the genus Draco. Tribe

2 contains the gmeraJapahira [in part] and Pseudocalotes

(subtribe 2.1), Sitana and Otocryptis (subtribe 2.2), Acan-

thosaura and Oriocalotes (subtribe 2.3) and Salea (sub-

tribe 2.4). Tribe 3 only contains Calotes. Tribe 4 is rep-

resented by Gonocephalus robinsonii and Japalura

polygonata. Tribe 5 consists of Ceratophora, Cophotis,

Pseudocophotis and Lyriocephalus (subtribe 5.1), Gono-

cephalus mjobergi (subtribe 5.2), Gonocephalus, Bron-

chocela, Complicitus, Hypsicalotes, Coryphophylax and

Aphaniotis (subtribe 5.3). Tribe 6 comprises Japalura [in

part, including Oriotiaris] and Ptyctolaemus. The remain-

ing tribes contain a single genus each: Tribe 7

Lophocalotes, Tribe 8 Phoxophrys, Tribe 9 Mantheyus and

Tribe 1 0 Dendragama.

The nodes produced in Pyron et al. (2013) are given in

the following as A-H with corresponding tribe numbers

(as given above) from Hoser in brackets: A(l) - Draco,

B(2) - Japalura Eastern clade, Pseudocalotes, Sitana,

Otocryptis, Acanthosaura, Salea, C(3) - Calotes, D(4) -

Japalura polygonata & Gonocephalus robinsonii, E(5) -

Ceratophora, Cophotis, Lyriocephalus, Gonocephalus,

Bronchocela, Coryphophylax, Aphaniotis, F(6) - Ptycto-

laemus and Japalura variegata clade, G(8) - Phoxophrys

and H(9) - Mantheyus.

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Wolfgang Denzer et al.

As can be seen Hoser’s taxonomic scheme essentially

reproduces the clades resulting from the phylogenetic

analysis by Pyron et al. (2013). His subtribes can be de-

rived in a similar way. Lophocalotes (Hoser’s Tribe 7) and

Dendragama (Hoser’s Tribe 10) are not included in Py-

ron’s analysis nor are Harpesaurus, Thaumatorhynchus,

and Psammophilus. The first two are recognized by Hoser

in their own monotypic tribes, the other three are not dealt

with at all. Other genera also not included in Pyron et al.

(l.c.) such as Oriocalotes, Hypsicalotes, and CompUcitus

are assigned to a tribe but without giving a reason for do-

ing so. However, with a bit of nomenclatural research one

could find possible reasons for his groupings: Oriocalotes

paulus was considered by Boulenger as Acanthosaura mi-

nor, hence Hoser’s pairing of these two genera. In his in-

troductory sentence to the genus CompUcitus he states that

it was “formerly placed in Bronchoceld\ In the same pub-

lication (Malkmus 1994, a paper written in German) Hyp-

sicalotes is also considered to be a member of the genus

Bronchocela. This is presumably Hoser’s reasoning be-

hind grouping these two genera in the same tribe along

with Bronchocela and several other species from the same

node in Pyron et al. (l.c.). Had Hoser decided to follow

the majority of earlier publications, all of which are cit-

ed by him, he would likely have included these two gen-

era in the tribe containing Calotes, their original genus

name.

Hoser’s division of the subfamily Draconinae can on-

ly be understood and followed if Pyron’s paper is at hand

for comparison. His classification scheme is poor-quali-

ty if not worthless as most tribes and subtribes are not di-

agnosed by shared characters but only through their con-

tent. In Hoser’s words: ’’...tribe is best defined by diag-

nosis of the component genera” or a similar wording. Such

a definition may comprise the character sets of 13 genera

as given in the first section of his paper where the genera

are defined. All diagnoses are copied and no additional da-

ta or characters are given. Some genera were not defined

in their own right in Hoser’s first section of the paper.

These genera are therefore diagnosed by him as a char-

acter set defining a tribe or subtribe. In the following we

will briefly analyze these additional diagnoses:

Hypsicalotes Denzer & Manthey, 2000 is diagnosed by

repeating entirely the diagnosis including comparisons to

other genera as given in Denzer & Manthey (2000, ap-

proximately 60 lines). Coryphophylax Fitzinger, 1869 is

defined in the spaee of six lines copied from Boulenger

(1885: 282). Cophotis Peters, 1861 is diagnosed by copy-

ing Boulenger (1885: 251-252, three lines; 275, three

lines). Pseudocophotis Manthey in Manthey & Gross-

mann, 1997 is a copy of Boulenger ’s description of

Cophotis sumatrana (= Pseudocophotis sumatrana).

CompUcitus Manthey in Manthey & Grossmann, 1997 is

defined in approximately two lines taken from the Rep-

tile Database website (primary source not identified).

Lophocalotes Gunther, 1 872 is diagnosed in approximate-

ly eight lines of which four lines each are from Boulenger

(1885: 251) and de RooiJ (1915: 116) or partially from

Hallermann (2004). The genus Phoxophrys Hubrecht,

1881 is defined for a second time but this time using

Boulenger (1885: 251, 280, six lines copied) instead of

using Inger (1960). The definition of Dendragama Doha,

1 888 has been copied (approximately three lines) from de

RooiJ (1915: 117-118).

Summary Section A

Apart from some minor alterations the phylogentic tree

published by Pyron et al. (2013) serves as Hoser’s main

basis for his classification scheme of the Draconinae.

In our analysis of Hoser’s proposed taxonomy for the

subfamily Draconinae most characters were identified and

can be attributed to other sources. Hoser gives his diag-

noses in the space of 2430 lines where 1884 lines consti-

tute the actual characters of which 1560 lines have been

identified as identical copies. If only diagnostic charac-

ters are taken into account approximately 83% are a copy

and if the full diagnoses are considered this percentage still

comes to approximately 64%. If the full paper is taken in-

to account (7140 lines, estimated) the copied text still

amounts to approximately 22%. At the end of his paper

Hoser cites several hundred references in a space of close

to 16 pages. However, according to our analysis he only

used approximately 30 of those to produce his proposed

taxonomy, the bibliography of which could probably have

been printed in the space of two pages. If this is taken in-

to account the percentage of copied text in relation to the

full paper rises by yet another 5%.

Hoser’s main source for descriptive characters was

Boulenger (1885). Additionally he copied sections from

Inger (1960) for Phoxophrys, Denzer & Manthey (2000)

for Hypsicalotes, Hallermann (2004) for Bronchocela,

McGuire et al. (2007) for Draco, Denzer & Manthey

(2009) for Gonocephalus, Mahony (2009) for Japalura,

Zug et al. (2006), Krishnan (2008) and Hallermann (2000)

for Calotes, Harikrishnan & Vasudevan (2013) for

Pseudocalotes, Pethiyagoda & Manamendra-Arachchi

(1998) for Ceratophora and Schulte et al. (2004) fox Ptyc-

tolaemus. Several genera such as Harpesaurus, Thauma-

torhynchus, and Psammophilus are not treated at all.

B) Hoser (2013) on Amphibolurinae

We note that the manuscript on Amphibolurinae was re-

ceived hy AJHoxv 20 July 2013, accepted for publication

on 4 October 2013, and published on 20 October 2013.

However, a tax invoice printed at the end of the publica-

tion (p. 36) states that the journal was printed on 3 Octo-

ber 2013, implying printed copies may have existed be-

fore the paper was accepted.

Bonn zoological Bulletin 64 (2): 117-138

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Critical review of Hoser’s publications on agamid lizards

127

The paper comprises the following sections or headings:

Title, Abstract (including Keywords), Introduction, fol-

lowed by the description of two genera and seven tribes,

References Cited, and a statement about Conflict of Inter-

est. An explicit section for Materials is missing (but see

below) and one has to assume that the description of the

newtaxa constitutes a combined Results/Discussion/Con-

clusion section.

The introductory part of a publication typically includes

a brief overview and often the author’s motivation for writ-

ing the paper, as well as his ideas about the subject. In stan-

dard practice, reviewers of manuscripts in mainstream

journals would not spend much effort in correcting this

part unless false statements are presented. However, in the

case of this particular introduction, it is instructive for a

better understanding of the broader picture of Hoser’s

works to mention several paragraphs.

Hoser begins by providing a reason for why the Aus-

tralasian Amphibolurinae Wagler, 1830 are so well stud-

ied. According to Hoser this “has arisen due to a combi-

nation of circumstances relatively unusual to Australia”

(emphasis added). The two factors alluded to are a “sta-

ble political and economic situation” including a transport

infrastructure that facilitates access to even the most re-

mote parts and “well-funded government paid herpetolo-

gists and relatively wealthy . . . private herpetologists . . .

able to travel to the most remote parts of the continent. . .

A significant portion of the Introduction deals with the

publications by Wells and Wellington (1983, 1985), which

are considered highly controversial papers in their own

right and still do not find full acceptance within the her-

petological community. One part of a paragraph reads as

follows (Hoser 2013: 34): “I have found myself resurrect-

ing names proposed by earlier authors. This includes a

number of effectively unused Wells and Wellington names

such as IntellagamaWQWs & Wellington, 1985, Gowidon

Wells & Wellington, 1 983 ...” However, at the time of this

paper’s publication (October 2013) the genus name Intel-

lagama had already been validated by Amey et al. (2012)

and the genus name Gowidon in the combination G. lon-

girostris (Boulenger, 1883) was made available by

Melville et al. (2011). Both publications were not cited by

Hoser (2013).

One of the paragraphs in Hoser’s Introduction would

never pass standard review of any formal publication in

science and would be removed by editors as it is against

ethical standards of publication. A group of herpetologists

(one named in particular) that is highly critical of Hoser’s

papers is called “a mob of criminals and ratbags” (Hoser

2013: 34).

At the end of the Introduction, Hoser uncritically lists

14 publications, five of which are his own, “and sources

cited therein” that apparently constitute the source mate-

rial for his research. However, major publications on Aus-

tralian agamids relevant to taxonomy and nomenclature.

such as Melville et al. (2011), Hugall & Lee (2004), Hugall

et al. (2008), Schulte et al. (2003), or Macey et al.

(2000a,b) are not mentioned at all and the reader would

need to refer to the few cited papers and their bibliogra-

phies to determine how Hoser derived some of his ideas.

While this paragraph could be regarded as a Materials sec-

tion, it should be noted that not a single museum speci-

men is referenced, nor is there any mention that museum

material was examined. Earlier in the Introduction Hoser

(2013: 33) notes: “In terms of the materials and methods,

this was based on my own field and lab work involving

most species as well as a review of the relevant literature

spanning the last 200 years.” With respect to “lab work,”

the reader does not get any further explanation of what

this entailed, making the process non-transparent and non-

reproducible. The list of references given in the References

Cited section of the paper comprises only 16 citations, five

of which are Hoser’s own publications. Of those, at least

Hoser (1998) on Acanthophis and Hoser (2012) on Afron-

aja are entirely irrelevant to agamid lizard taxonomy.

Only Joger (1991) and Pyron et al. (2013) are referenced

as publications that include original molecular phyloge-

netic research that is indispensible to Hoser’s arguments.

However, Joger ’s paper (l.c.) only includes Amphibolu-

rus vitticeps (Ahl, 1926) (= Pogona vitticeps) and

Physignathus temporalis (Gunther, 1867) (= Lophog-

nathus temporalis) in the analysis, making it of only pe-

ripheral interest for a detailed phylogenetic analysis of

Australian taxa (Joger 1991, Material Examined). Joger

(1991: 619) even notes: “Because of the lack of antisera

for East Asian and Australian agamids, the position of their

three lineages - AmphibolurusI Physignathus,

CaloteslAcanthosaura, and Gonocephalus - relative to

each other could not be determined.” It is important to note

that Joger did not study the genera Physignathus Cuvier,

1829 mdAmphibolurus Waglor, 1830, but two specimens

of populations that were considered members of these gen-

era at that time, but are assigned to different genera to-

day. Additionally, no nomenclatural decisions were pro-

posed by Joger (l.c.).

It therefore stands to reason that Pyron et al. (2013)

serves as the basis for Hoser’s taxonomic and nomenclat-

ural proposals. This becomes particularly obvious in the

grouping of Moloch Gray, 1841 and Chelosania Gray,

1 845 within a single tribe. Without recent molecular phy-

logenies it is unlikely that any morphologically-oriented

herpetologist would group a thorny devil (whose vernac-

ular name illustrates a key aspect of the species’ scale mor-

phology) with a lizard that has a completely homogeneous

dorsal scalation. The phylogeny of Pyron et al. (2013) is

also reflected in acknowledging the difference between

Physignathus and Intellagama, the splitting of Hypsilu-

rus into several genera, the erection of a tribe for the genus

Ctenophorus, and combining Amphibolurus, Chlamy-

dosaurus, Diporiphora, Gowidon (Lophognathus), Pog-

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Wolfgang Denzer et al.

ona, Rankinia, and Tympanocryptis within one tribe, al-

beit using a different nomenclature.

The Introduction is followed by a section that serves to

erect two new genera and seven tribes. New tribes are pre-

sented in random order without presenting necessary in-

formation on possible phylogenetic relationships. Our

analysis will mainly follow Hoser’s order but for reasons

of clarity the tribe containing the genera related to Am-

phibolurus and the tribe containing all Ctenophorus will

be dealt with last.

The first genus described in the paper is meant to ac-

commodate only Diporiphora superba Storr, 1974. The

actual diagnostic characters account for approximately

nine lines of text, of which eight can be accounted for in

Cogger (1983: 238, key leading to D. superba, 1983: 243,

description of D. superba). Differences include changing

“gular fold absent” to “no gular fold” and replacing the

% sign by “percent,” as well as replacing the numeric “4”

by the word “four.” The unaccounted text consists main-

ly of introduced verbs and a slightly modified description

of the colouration.

The second genus described deals with Hypsilurus

spinipes (Dumeril & Bibron, 1851). The diagnostic char-

acter section accounts for approximately 12 lines and is

annotated as “adapted from Cogger, 2000.” About 11 lines

are the result of directly repeating Cogger’s description

of H. spinipes (Cogger 1983: 245-46, as Gonocephalus

spinipes) and the only differences are the introduction of

a few verbs and conjunctions.

The new genus containing Hypsilurus spinipes is sub-

sequently placed into a newly erected tribe that addition-

ally contains Tiaris Dumeril & Bibron, 1837 (see below).

The diagnosis for the tribe contains two sets of charac-

ters. The first part (ca. seven lines) is a copy of Cogger’s

key leading to Hypsilurus (Cogger 1983: 217, as Gono-

cephalus) apart from a few introduced verbs and conjunc-

tions (one line). The second part (eight lines) states the

characters shared by Hypsilurus species and is a copy (six

lines) of the genus diagnosis given by Manthey & Denz-

er (2006). The latter paper is not cited, and the diagnosis

was most probably retrieved from the Reptile Database

(Uetz & Hosek 2015), where it is publicly available (cit-

ed, and with approval of the authors). The genus Tiaris is

not characterized at all in Hoser’s paper, nor is the read-

er informed which species it contains. In Hoser’s paper

the name stands on its own and is therefore a nomen

nudum according to the Code. It should also be noted that

Tiaris Dumeril & Bibron, 1837 is not available for any

agamid genus as it is preoccupied by Tiaris Swainson,

1 827 (Aves, Passeriformes) [see Manthey & Denzer (2006

)]•

According to Hoser an agamid genus Tiaris is of Aus-

tralian origin and “the only genus it is likely to be con-

fused with” is the one newly erected for Hypsilurus

spinipes. The only other Hypsilurus species in Australia

that Hoser could refer to is H. boydii (MacLeay, 1884),

albeit that it is actually quite difficult to confuse these two

species. Hypsilurus boydii is known from older literature

as Tiaris boydii (e.g., MacLeay 1884). Hence, we assume

that Hoser meant to include this species in the same tribe

as H. spinipes. However, H. boydii is morphologically

(Manthey & Denzer 2006) and genetically (Pyron et al.

2013) closely related to H. dilophus (Dumeril & Bibron,

1837), which Hoser places into a different tribe.

The next tribe is erected to accommodate the genus Hyp-

silurus, assigning Lophura (Hypsilurus) godeffroyi Peters,

1867 as the terminal taxon. Hoser’s introduction to the

tribe starts with the sentence “Currently most widely

known as Hypsilurus dilophus (Dumeril & Bibron,

1837).” It is not clear whether Hoser here expresses his

view that H. godeffroyi (a well-defined and valid species)

is identical to (conspecific with) H dilophus. His state-

ment is even more confusing considering that H. dilophus

is actually the type species of the genus Tiaris Dumeril

& Bibron, 1 837, a genus he assigned a paragraph earlier

to a different tribe (for synonymy of Hypsilurus see Man-

they & Denzer 2006). Additionally there exist no objec-

tive reasons to combine H dilophus and H. godeffroyi in-

to one group. Morphologically, they are very different

species that were even placed into different species groups

by Manthey & Denzer (2006). In his genus description to

accommodate H. spinipes, Hoser states that Tiaris (nomen

nudum, see above) and all Hypsilurus species have a “lon-

gitudinal row of grossly enlarged scales on the throat.”

This is actually a character used by Cogger (1983) to dif-

ferentiate between H spinipes and H boydii, which should

read, “median longitudinal line of . . . similar to those in

the nuchal crest.” The statement holds true if only Aus-

tralian species are considered, as is the case with Cogger

(I.C.), but when including taxa outside of Australia, as

Hoser’s analysis does, it is false, as most species of Hyp-

silurus outside Australia actually lack a median line of en-

larged scales on the gular pouch. These are only well de-

veloped in H. boydii and H. dilophus, and to a lesser ex-

tent in H. hikidanus Manthey & Denzer, 2006. As already

pointed out by Manthey & Denzer (2006), H dilophus,

H boydii, and H. spinipes may be considered as a species

group, and if considered as a separate genus only the name

Lophosaurus Fitzinger, 1843 would be available but not

Tiaris. The diagnostic character section for Hypsilurus

comprises 15 lines of which 13 lines are copied from Cog-

ger (l.c.) and Manthey & Denzer (2006).

Subsequently, Hoser deals with the water dragons from

Australia and Southeast Asia. The relationship, biogeog-

raphy, and nomenclature of Physignathus cocincinus Cu-

vier, 1 829 and Intellagama lesueurii (Gray, 1831) has been

a matter of intense discussion, and since the advent of mo-

lecular phylogeny there have been several publications to

address the issues (e.g., Schulte et al. 2003, Macey et al.

2000a,b). Still Hoser makes no mention of this and erects

Bonn zoological Bulletin 64 (2): 117-138

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Critical review of Hoser’s publications on agamid lizards

129

two new tribes. His claim to have resurrected the genus

name Intellagama has already been dealt with (see above).

The diagnostic characters to define the tribe for 7. lesueurii

account for ca. five lines of text, three of which are copied

from Cogger’s key (Cogger 1983: 217). The remaining

part of the description deals with colouration, but not as

one would expect, with a description of the colouration

of 7 lesueurii. Hoser instead merely states that the coloura-

tion is not that of P. cocincinus.

The tribe containing the latter species is again initially

diagnosed via Cogger’s key (f c.), with three lines out of

four being copied. The remaining part of the diagnosis

deals with the colouration of Physignathus cocincinus.

Apart from the dorsal ground colour (one line) the pat-

terns and colourations (three lines) were copied from

Wikipedia, with no primary source identifiable. The

Wikipedia page already provided this description of the

colouration in 2012 (accessed August 2014, file history

checked for December 2012), i.e. before Hoser’s paper

was published.

Next Hoser introduces a tribe to accommodate the gen-

era Moloch and Chelosania. The two diagnostic sections

consist of two lines and four lines, respectively, of which

approximately four lines are copied text (Cogger 1983:

217).

Another tribe is erected to accommodate the genus

Ctenophorus. The diagnostic characters are presented in

roughly sixteen lines, ten of which are copied from Cog-

ger (1983) and three from Cogger (1993). One set of char-

acters regarding the supralabial scales could not be ac-

counted for and is potentially the only part of an original

description in the entire paper. From the lack of method-

ology, it is not possible to determine how these observa-

tions were made or which specimens were used, render-

ing the data non-reproducible. Furthermore, the copied

part contains several mistakes that need to be addressed.

The diagnosis for the new tribe containing the genus

Ctenophorus is at best confusing, and perhaps of no tax-

onomic value entirely. Part of Hoser’s definition reads as

follows (emphasis added): “nuchal crest and/or series of

enlarged keeled vertebral scales present or absent and if

absent present along at least the anterior two thirds of

the body; enlarged strongly keeled or spinose scales are

present elsewhere on the dorsum.” The diagnostic char-

acters are identical those used in Cogger’s key to the gen-

era (1983: 217), apart from the conflicting phrase “absent

present” and the placement of the semicolon. The use of

both “absent” and “presenf ’ in close combination makes

it unclear how this character is to be scored. In common

usage, placing a semicolon will not change a diagnosis sig-

nificantly. However, in this case only the part directly pre-

ceding the semicolon relates to the character of “enlarged

keeled vertebral scales present or absenf’. The character

after the semicolon “enlarged strongly keeled scales . . .

present ... on the dorsum” stands on its own. This way

all Ctenophorus without this character are excluded from

the genus! Cogger (1983: 217) included this particular set

of characters as a full statement in the diagnosis for the

genus Amphibolurus. There it reads, ’’nuchal crest and/or

vertebral keel may be present, but if the latter is present

on at least the anterior two-thirds of the body then en-

larged, strongly-keeled or spinose scales are present else-

where on the dorsum”, giving it a completely different

meaning. It should be noted that a key matching this part

of Hoser’s diagnosis, including the (wrong) placement of

the semicolon, can be found in another earlier publication

(Cogger 1993: 163, or in the online version on page 10,

character 9a). A further character to define the tribe pre-

sented by Hoser is described on the basis of the online pub-

lication (page 11) but introducing yet another mistake.

Hoser’s character reads: “a row of enlarged scales from

below the eye to above the eye” instead of “to above the

ear”!

The genus- and species-richest tribe introduced in the

paper contains the genera Amphibolurus, Chlamy-

dosaurus, Caimanops, Cryptagama, Diporiphora,

Gowidon (Lophognathus), Pogona, Rankinia, and Tymap-

nocryptis, as currently accepted by most Australian her-

petologists (here listed according to Cogger 2014; it should

be noted that Gowidon is not yet generally accepted).

However, according to Hoser’s compilation the genus

Lophognathus no longer exists. Hoser instead uses

Gowidon, a name available for 7. longirostris, but ignores

7. Wells & Wellington, 1985, 7. gilberti Gray, 1842,

and 7. temporalis (Gunther, 1867). For this reason the

reader has to assume that Hoser considers these species

as congeneric or even conspecific. However, in this case

the name Gowidon would not be available since the genus

name Lophognathus has nomenclatural priority over it,

with 7. gilberti being the type species of the genus.

If Hoser had been consistent in following the data of Py-

ron et al. (2013), then Tympanocryptis Peters, 1 863 should

also be a member of this tribe. However, Hoser does not

include it here or in any other tribe, nor does he use a dif-

ferent taxonomy to pinpoint where the species of this

genus might be grouped, perhaps as part of one of the oth-

er genera used in the revised classification scheme. It ap-

pears that, just as some members of Lophognathus, the

genus Tympanocryptis was simply disregarded or forgot-

ten. Tympanocryptis is an available name that should be

used, the type species being T lineata Peters, 1863.

Hoser recognizes Caimanops Storr, 1974 and two gen-

era proposed by Wells & Wellington (1983, 1985) con-

taining Diporiphora species. One of these genera was

erected to accommodate D. albilabris albilabris Storr,

1974 and 7). albilabris sorbia Storr, 1974. The second was

erected for 7). /zwga Houston, 1977 and 7). winneckei Lu-

cas & Frost, 1 896. We assume that Hoser resurrects these

genera from their synonymy with Diporiphora, albeit

without mentioning it specifically or giving a reason for

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130

Wolfgang Denzer et al.

doing so. These genera were formally synonymized with

Diporiphora by Doughty et al. (2012), owing to their close

phylogenetic relationship, and were included by Pyron et

al. (l.c.) under Diporiphora. It appears that Hoser over-

looked the publication by Doughty et al. (2012).

Caimanops has so far been considered a monotypic genus,

with C. amphiboluroides as the sole species. If Hoser had

properly followed Pyron et al. (l.c.), he should also have

assigned D. australis to Caimanops as these two species

form a clade.

The diagnostic characters employed by Hoser to group

the above genera into a new tribe are as follows: Initial-

ly he singles out Chlamydosaurus by repeating Cogger

(1983 two out of two lines copied) but introducing a mis-

take. Hoser ’s character reads, “a large loose frill or skin

around the neck” but it should read, “. . . frill of skin . . . ”.

Hoser then proceeds to define the general characters of

this diverse group (Cogger 1983; four out of four lines

copied). In his last part of defining the tribe he excludes

the genus Ctenophorus in an identical way as he defined

the tribe containing the genus, including all the mistakes

discussed above. Overall Hoser’s diagnostic character sec-

tion of this tribe comes to 22 lines of which 16 lines are

a result of copying.

Summary Section B

Hoser’s classification scheme for amphibolurine lizards

mostly reflects the nodes in the phylogenetic tree pub-

lished by Pyron et al. (2013). Additionally several genera

proposed in the highly controversial papers by Wells &

Wellington (1983, 1985) are accepted as valid.

Hoser gives his diagnoses in the space of 153 lines, of

which 121 lines constitute the actual characters, with 100

lines clearly identifiable as copied. If only diagnostic char-

acters are taken into account this amounts to 82% copied

material, with the full diagnoses included it is still 65%.

If the full paper is taken into account (438 lines) the copied

text amounts to 23%.

All but a single character can be identified and attrib-

uted to secondary sources (Cogger 1983, 2000; Manthey

& Denzer 2006; Anonymous on Wikipedia; Uetz & Hosek

2015).

Most characters used to describe genera are taken di-

rectly from Cogger (1983 or subsequent editions). Sever-

al important publications on Australian agamid lizards

such as Melville et al. (2011), Hugall & Lee (2004), Hugall

et al. (2008), Schulte et al. (2003), or Macey et al.

(2000a,b) have been omitted. The genera Lophognathus

and Tympanocryptis are not treated at all.

C) Hoser (2012a) on Laudakia Gray, 1845

The manuscript was received on 13 March 2012, accept-

ed on 8 April 2012 and the paper was published on 30 June

2012. Hoser’s paper is presented in the following way: Ab-

stract (including Keywords), Introduction, description of

taxa, and References Cited. The second to fifth paragraphs

of the Introduction, describing the general appearance and

behaviour of the group, contain 13 lines of copied mate-

rial from an online source (www.sauria.co.uk). The rest

of the Introduction deals with the nomenclature and phy-

logeny of Laudakia Gray, 1 845 and Phrynocephalus Kaup,

1825. His nomenclatural arguments refer mainly to Hen-

le (1995) and are misinterpreted (see comments for the

stellio group).

Hoser’s “five- way division” of Laudakia Gray, 1845

mainly reflects the phylogenetic schemes published by

Macey et al. (1998, 2000b, 2006), who identified nodes

supporting a L. tuberciilata group, a L. caucasia group

with L. lehmanni as the sister taxon (proposed as a new

subgenus by Hoser), as well as nodes supporting the

monophyly of L. stellio (Linnaeus, 1758) and L. sacra

(Smith, 1935). Hoser’s fifth group comprises Phryno-

cephalus Kaup, 1825, a genus of lizards that has never

been in the synonymy of Laudakia. Macey et al. (2000b)

found Phrynocephalus to be a sister taxon to both the clade

containing the L. caucasia group and L. stellio. Howev-

er, in a later publication by Melville et al. (2009), the

monophyly of the genus Laudakia was confirmed and

Phrynocephalus emerged as the sister taxon to the whole

clade. Apparently, Hoser and his supposed reviewers over-

looked this important publication, which is not cited in his

bibliography.

The first genus Hoser deals with is that of Phryno-

cephalus Kaup, 1 825, which he considers “similar in most

respects to Laudakia sensu lato” (Hoser 2012: 1 8), a state-

ment most herpetologists would disagree with. Hoser does

not present a meaningful definition of the genus apart from

“lacking of an obvious tympanum” to distinguish Phryno-

cephalus from Laudakia and a “dorsoventrally de-

pressed” body to distinguish it “from all other other

Agamids in the region where these groups of lizards oc-

cur.” This entire diagnosis holds no definitive value as

there are other agamid genera in the area under consider-

ation that have a dorsoventrally depressed body shape

(e.g., Brachysaura Blyth, 1856, Bufoniceps Arnold,

1992, and Trapelus Cuvier, 1829). Another interesting fact

is that Hoser only recognizes 26 species within the genus

Phrynocephalus while the actual number had already sur-

passed 40 species by the time his paper was published.

This is certainly something any expert reviewer would

have been able to point out, even by a simple search of

the Reptile Database (Uetz & Hosek 2015).

Hoser moves on to define what he considers to be the

actual genus Laudakia. This is represented by the tuber-

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Critical review of Hoser’s publications on agamid lizards

131

culata group and hence L. tuberculata (Gray, 1 827) be-

comes the type species. Initially he uses tympanum pres-

ence and body shape to distinguish it from Phryno-

cephalus and other genera of agamid lizards in the region.

This is followed by a short description (eight lines) most-

ly copied from a key to the species of agamid lizards of

Pakistan (six lines; Khan 2002: 100 & 101). The same key

is available on the Internet as part of Khan’s undated

eBook Herpetology of Pakistan. Neither of these two pub-

lications are cited by Hoser! Based on the idiosyncratic

character “fifth toe extends beyond second” we have no

doubts that Khan’s publication is the source; other authors

make a comparison to the first toe. Hoser’s further char-

acters include the dentition and the number of whorls in

each tail segment. Both characters are from Baig (1992)

but the writing has been sufficiently altered that they do

not constitute copied material. The entire diagnosis of the

genus comprises 32 lines (18 lines of diagnostic charac-

ters) of which six lines are copied, presumably from Khan

(2002).

The next genus proposed by Hoser serves to accommo-

date the Laudakia caucasia (now Paralaudakia caucasia,

see below) group. Up to this point we have mostly ab-

stained from judging Hoser’s diagnoses but the poor qual-

ity of this one requires analysis. It begins with an essen-

tially copied general diagnosis from earlier in the paper,

but “excluding those genera formerly placed within Lau-

dakia sensu lato” by which Hoser means the other gen-

era he proposed (see above). Subsequently, he copies from

Khan (2002): “tympanum is large,. . . fifth toe extends be-

yond second; caudal scales in distinct annuli,” which is

unfortunately a character set that still defines the entire

genus. Up to this point, no character has been listed that

could be used to define the new genus. Next, Hoser states

that “the scales of dorsal rows are smooth,” “the premax-

illa has two teeth in the [new] genus versus three in Lau-

dakia [= L. tuberculata group],” and “lizards in this [new]

genus have 14-15 molars, versus 14-15 [sic!] in Lau-

dakia.'" The author then once more repeats the general

paragraph to differentiate Phrynocephalus and other

agamids. He then separates his new genus from L. sacra

by providing a full description of this species that is iden-

tical to the one found in Ananjeva et al. (1990). To sum-

marize this for clarity: the only diagnostic characters pre-

sented to define his new genus -other than those charac-

ters which are common to all genera concerned- are

“scales of dorsal rows are smooth,” “premaxilla has two

teeth,” and presumably the number of molar teeth.

The definition for the new genus is, unfortunately for

Hoser, not cohesive because the vertebral scales of Par-

alaudakia caucasia (Eichwald, 1831), the proposed type

species of the new genus, are actually keeled and those

of P. himalayana (Steindachner, 1867) and P.

badakhshana (Anderson & Leviton, 1969) are smooth

(Boulenger 1885; Khan 2002; Baig et al. 2012). We are

not aware - and it is outside the scope of this paper to in-

vestigate further - how many teeth are present in the pre-

maxilla and how many molars the other genera possess,

in order to verify or falsify these two characters, nor is

Hoser apparently. To our knowledge there are no publi-

cations dealing with the dentition of all genera in ques-

tion. In total his diagnosis of this genus comprises about

56 lines, of which 27 are copied from Ananjeva et al.

(1990). Three lines describing diagnostic characters are

taken from Baig (1992) and Khan (2002), but not copied

directly. Baig et al. (2012) established the genus Paralau-

dakia Baig, Wagner, Ananjeva & Bohme, 2012 to accom-

modate species related to caucasia and himalayana as well

as lehmanni and stoliczkana.

For the Laudakia stellio (now Stellagama stellio, see be-

low) species group, which Hoser considers to be mono-

typic, the author resurrects Plocederma Blyth, 1 854. This

can only be explained by misinterpreting Henle (1995)

who proposed to use this genus name for the stellio group,

which he considered to comprise L. stellio, L. caucasia,

L. erythrogastra, L. himalayana, L. lehmanni, L. nupta,

and L. melanura. The type species for the genus Ploced-

erma is L. melanura. Only if this species were included

in Hoser’s stellio group (which it is not) would the name

be available for the group. Because the stellio group as it

is considered nowadays (i.e. monotypic) did not have any

previous available name disposable, Baig et al. (2012) es-

tablished the name Stellagama Baig, Wagner, Ananjeva

& Bohme, 2012.

The new genus to accommodate Stellagama stellio is

initially only defined by repeating his general description

(two characters: “distinct tympanum” and “dorsoventral-

ly depressed” body) followed by two lines taken from

Khan (2002) and a description of ''Laudakia stellio" (ap-

proximately 14 lines, all copied) taken from a website

(Go 9 men, www.bayramgocmen.com/album/picture.php?

/1012/category/345, accessed September 2014). The full

diagnosis comprises 26 lines (18 lines of diagnostic char-

acters), of which 16 are copied.

To define his newly proposed monotypic subgenus to

accommodate Laudakia lehmanni (Nikolsky, 1896),

Hoser presents diagnostic characters in the space of ap-

proximately 38 lines, all of which come straight from

Baig’s description (1992) of L. lehmanni. The order of

characters is slightly different from the original and in a

few places verbs or conjunctions have been added. The

diagnosis is followed by distributional data and habitat de-

scription, which constitutes (apart from one sentence) a

copy of the text produced on the lUCN RedList webpage

(six lines out of seven copied). The whole diagnosis com-

prises 42 lines of which 38 lines constitute diagnostic char-

acters all of which are a copy from Baig (1992).

Hoser lists four papers by Baig and co-authors in the

References Cited section, but none of these contains a de-

scription of L. lehmanni (now Paralaudakia lehmanni, see

Bonn zoological Bulletin 64 (2): 117-138

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132

Wolfgang Denzer et al.

below). A detailed description of P. lehmanni that is very

similar to the one used by Hoser - but with characters in

a different order - was first given in Baig’s PhD thesis

(Baig, 1992: 130 & 132, not cited by Hoser) which was

completed under the supervision of WB. Since Baig’s the-

sis has to be considered unpublished, Bohme and cowork-

ers (Baig et al. 2012) posthumously published a paper

based on the thesis to preserve Baig’s extensive taxonom-

ic work and to make it available for scientists working on

this subject. In the latter publication Baig’s description is

repeated with minor changes and with pretty much the

same wording seen in Hoser (2012a). Baig et al. (2012)

was published in print on 1 8 July 2012 and Hoser (2012a)

was published in print 30 June 2012. Both papers were

accepted for publication by the respective journals in April

2012. We also note that Baig et al. (2012) was made avail-

able in advance online on the publisher’s website on 6 Ju-

ly 2012, appearing a week after Hoser’s publication. The

most likely way by which Hoser would have been able to

retrieve Baig’s text would have been by downloading the

thesis from a governmental website in Pakistan (Pakistan

Research Repository, http://eprints.hec. gov.pk/2407/1/

2262.htm). Although it is difficult to proof, but based on

the exact wording, we are convinced that Baig’s thesis was

available to Hoser, who did not consider it necessary to

reference it. However, even if not published a PhD the-

sis constitutes intellectual property belonging to the can-

didate and his thesis supervisor. In any case the precise

repetition of wording from a thesis without appropriate

clarification, attribution and referencing constitutes a vi-

olation of authorship rights. Any use of a verbatim copy

of excerpts from a thesis needs permission by either the

author, his thesis supervisor or the university department

where the candidate studied for the degree. However, a

reader who does not know about Baig’s thesis might sup-

pose that Hoser’s diagnosis has precedence, with Baig et

al. (2012) copying Hoser’s ideas and wording when the

opposite is the case. In this instance, Hoser clearly uses

the intellectual property of another and passes it off as his

own. Such behaviour would even be seen as plagiarism

if Hoser obtained the description from a third source,

which we have not identified. In a recent paper Hoser

(2015) even claims priority and that “they [Baig et al.

(2012)] did however remanufacture theirs [morphologi-

cal evidence] as “new” data, which in itself is fraudulenf ’.

Hoser (l.c.) clearly states that data were available from ear-

lier studies but again does not disclose or cite the source.

Not only did Hoser plagiarize Baig (1992), he even con-

siders his actions as justifiable and additionally accuses

the true original author of fraudulent behaviour!

The last genus Hoser proposes is monotypic for Lau-

dakia sacra (Smith, 1935). His diagnosis is given within

27 lines, of which 25 are a direct copy from the descrip-

tion of L. sacra by Ananjeva et al. (1990; see also Uetz

& Hosek 2015) and two lines are copied from Khan

(2002). The only two other characters are those used pre-

viously (“a distinct tympanum” and “the body is dorso-

laterally depressed”), repeating nearly the entire general

paragraph for the fifth time. The diagnosis of this genus

is given in 33 lines, with 30 lines presenting the actual di-

agnostic characters, of which 27 lines are copied.

Summary Section C

The taxonomic basis for Hoser’s proposals on Laudakia

can be found in their entirety in Macey et al. (1998, 2000b,

2006). Most of Hoser’s proposed classification addition-

ally reflects nodes in the phylogeny published by Pyron

etal. (2013).

In total, Hoser’s paper on Laudakia comprises an esti-

mated 980 lines, of which 420 lines constitute his Refer-

ences Cited section (560 lines pure text including Title and

Abstract). We would like to mention that already his in-

troductory part contains at least 13 lines that can be found

on websites (not taken into account here as copied text)

and that we further identified several diagnostic charac-

ters Hoser used but without copying directly. His diag-

noses come to 180 lines of which 148 lines constitute di-

agnostic characters. With respect to the latter we found that

114 lines (77%) were copied from previously published

research papers or reviews. Hoser’s main sources are

Ananjeva et al. (1990), Baig (1992), Khan (2002) and Baig

et al. (2012). Hoser (2015) even claims priority with re-

spect to the data albeit that his taxonomic scheme and all

his characters have been copied from Baig (1992) and sub-

sequent publications.

D) Hoser (2014c) on Uromastycinae

The manuscript of this paper was received by the journal

on 2 November 2013, accepted on 15 May 2014, and fi-

nally published on 30 August 2014. It is presented in the

following way: Abstract (including Keywords), Introduc-

tion, Notes on the taxa named herein followed by the de-

scription of taxa. Conflict of Interest, and References Cit-

ed. In total Hoser newly describes or resurrects within this

publication two tribes, five genera, and four subgenera.

Five of these taxa are monotypic.

The Introduction is relatively short and summarizes the

taxonomic history of Uromastyx and gives Hoser’s view

on taxonomy, without any identified copied parts. How-

ever, two extraordinary statements should be discussed

here. In terms of material used for his study, Hoser refers

to the “inspection of live specimens at various facilities

since 1993.” Many Saara or Uromastyx species inhabit

political unstable areas and it is very unlikely that there

are live specimens of many important species, such as S.

asmussi (Strauch, 1863), available at any facility Hoser

might have visited since 1993. Therefore it is very prob-

lable that most of the data he presents are not from ex-

Bonn zoological Bulletin 64 (2): 117-138

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Critical review of Hoser’s publications on agamid lizards

133

amined material, but from published sources. These

sources are cited in the Introduction as “Significant stud-

ies relevant to the taxonomy of the Uromastycinae ...” and

include for example Hall (1999) and Swofford (2002), two

general publications about statistical phylogenetic meth-

ods without any significance to the taxonomy of the group

at all.

In his Abstract and Introduction, Hoser gives the im-

pression that he is the first to use the approach by Pyron

et al. (2013) to distinguish between Uromastycinae

Theobald, 1868 and Leiolepidinae Fitzinger, 1843 on the

subfamily level, but mentioning in passing that “some au-

thors have already taken this step.” This concept was used

more than a decade ago by Macey et al. (2000a), in a pa-

per not cited by Hoser. Instead, Hoser cites Macey et al.

(2000b) on the trans-Tethys migration, which has hardly

any relevance to uromasticine / leiolepidine taxonomy (on-

ly one species of Uromastyx and two species of Leiolepis

were included in the study). We also note that the terms

Uromastycinae and Leiolepidinae were used synonymous-

ly by different authors (e.g., Wilms & Bohme 2007: 436).

The entire first definition of Uromastycinae used by

Hoser is identically phrased to Wilms et al. (2009:67; four

lines), followed by the definitions of his two new tribes

copied from the same source. The further detailed defini-

tion for this subfamily is not fully copied, but obviously

taken from Boulenger’s synopsis (Boulenger 1885: 405;

14 lines). All diagnosing parts of the subsequent defini-

tion of Uromastyx are entirely taken from various parts

(text and key) of Wilms et al. (2009).

The first taxon Hoser describes is the subgenus Uro-

mastyx (within Uromastyx) and the given diagnosis is tak-

en from Wilms et al. (2009: 67 & 82; 35 lines copied for

the genus Uromastyx, seven for the subgenus Uromastyx,

of which 3.5 lines are within quotation marks). For his sec-

ond, monotypic subgenus diagnosis, erected to accommo-

date U. occidentalis Mateo, Geniez, Lopez-Jurado &

Bons, 1999, he uses eight lines directly copied from Wilms

et al. (I.C.).

Even though Hoser quotes Wilms et al. (l.c., 3.5 lines)

in the following diagnosis of Aporoscelis, the parts not di-

rectly quoted are also copied from that reference (two

lines).

For his first new genus description Hoser is using a dif-

ferential diagnosis separating his new taxon by the diag-

nosis of other taxa. Here, Hoser is summarizing the in-

formation given in the diagnostic key by Wilms et al.

(2009: 82), followed by a description taken from

Boulenger (1885: 405) with 26 lines copied. Also the di-

agnosis of the same taxon at a different rank (subgenus)

is taken from Wilms et al. (2009, 11 lines copied). With-

in his new genus, Hoser describes two additional new sub-

genera. Even here all mentioned characters diagnosing

these taxa are identical to Wilms et al. (2009: 82; 22 lines

copied).

Diagnosing his second new genus, Hoser is following

the same scheme of presenting a differential diagnosis.

And again, all given characters, especially meristic char-

acters (e.g., scale or whorl counts) are exactly the same

as given in Wilms et al. (l.c.) and no other additional char-

acters are provided. This description is followed by the

subgenera to be included in the previously described

genus. Again, all mentioned characters are exclusively tak-

en from Wilms et al. (l.c.). The new genus is described

by copying 14 lines from Wilms et al. (2009: 82-83), and

subsequently three new subgenera are proposed using 44

lines from the same source.

While redescribing the genus Saara according to his

new taxonomy, Hoser provides several characters to dis-

tinguish his new tribe, including Saara, from the tribe that

includes Uromastyx. Here he is exclusively using the char-

acters provided by Wilms et al. (2009: 81-82; 15 lines)

for the three species forming the genus Saara. However,

Hoser is splitting this genus into three distinct monotyp-

ic genera, including in addition to Saara the genus Cen-

trotrachelus Strauch, 1 863, which he resurrects to accom-

modate S. asmussi, and a new genus that only contains S.

loricatus Blanford, 1874. In order to describe these two

genera Hoser again uses Wilms et al. (2009: 81-82; 32

lines copied).

Finally, Hoser erects two new tribes to accommodate his

proposed genera. The first tribe is described using three

lines from Wilms et al. (2009:67) and 13 lines from

Boulenger (1885:405). The second tribe is solely defined

by characters given by Wilms et al. (2009: 67, 82 & 83;

15 lines).

In the References Cited section Hoser lists 154 refer-

ences (three-and-a-half pages), giving the impression of

a well-conducted, literature-based study. However, none

of the references is cited in the running text (other than a

lengthy list of general references as part of the Introduc-

tion), 78 of the references do not refer to Uromastyx tax-

onomy or distribution (several are concerned with

Leiolepis, others with maintenance of Uromastyx), and 48

references do not refer to Uromastyx at all (including de-

scription of statistical methods, herpetofaunal lists outside

the distribution of Uromastyx). The only references Hoser

appears to actually use are those by Boulenger (1885) and

Wilms et al. (2009), from which many lines are copied ver-

batim without appropriate attribution.

Summary Section D

The taxonomic basis for Hoser’s proposals on Uromastyci-

nae is a representation of nodes taken fron the phyloge-

ny published by Pyron et al. (2013).

In total, Hoser’s paper on Uromastyx contains an esti-

mated 1490 lines, of which 490 lines are referenced pub-

lications and 1000 lines are text (inclusive of title and ab-

stract). His diagnoses contribute 556 lines, of which 326

Bonn zoological Bulletin 64 (2): 117-138

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134

Wolfgang Denzer et al.

lines are diagnostic characters. With respect to the latter,

of all diagnostic characters mentioned, 255 lines (78%)

were identically phrased or copied from previously pub-

lished research papers or reviews, with Wilms et al. (2009)

and Boulenger (1885) being the main sources.

SUMMARY & CONCLUSIONS

We analyzed four of Hoser’s publications on agamid

lizards and found in all cases significant amounts of copied

or plagiarized text to present the diagnostic characters

(83% for Draconinae, 82% for Amphibolurinae, 77% for

Laudakia, 78% for Uromastycinae). There is no harm per

se in repeating diagnostic characters from the older liter-

ature and using them in order to define a genus. A species

of Draco has a certain number of elongated ribs in the

patagium and the nostrils are directed sidewards or up-

wards. Similarly, species of Gonocephalus have a gular

fold and Chlamydosaurus kingi possess a frill. There ex-

ist only a limited number of different expressions to pres-

ent certain character sets. However, we think that Hoser’s

approach is on a different level that most scientists and

editors would consider a sort of plagiarism. We found

paragraphs that clearly show that Hoser’s presentation of

the diagnosis is a result of a copy-and-paste procedure with

typographical errors in exactly the same place in his text

where they occurred in the original publication. Further-

more, the direct uses of statements from the older litera-

ture lack attribution; merely including titles in a bibliog-

raphy is not attribution. Meristic characters or statistical

values tend to be given with the identical numbers of a

source paper (for examples, see the sections on Draco or

Uromastyx) although it is clear that Hoser neither took any

measurements nor conducted a statistical analysis, as he

would not have had access to the same specimens (or any

specimens for that matter).

By pure repetition of character sets, which are often as

old as 125 years, several of Hoser’s diagnoses are rendered

inaccurate, inconsistent, or even false. Often a diagnosis

consists of more than one character set taken verbatim

from two different publications; sometimes as much as

half a page is copied in full, or long descriptions are tak-

en directly from a previous publication. In at least three

cases {Laudakia, Paralaudakia lehmanni, Hypsilurus),

sets of characters that were copied by Hoser had been pub-

lished in an identical or near-identical manner before, but

the original sources are not cited at all!

We were able to identify the sources of most (~ 90%)

of the diagnostic characters used by Hoser (2012a, 2013,

2014b, c). If the percentage of word-for-word copying of

Hoser’s diagnostic characters section is evaluated, this

amounts to approximately 80% of his presentation. Even

if the whole diagnoses are taken into account for which

Hoser typically uses long sentences that have nothing to

do with the actual definition of the taxon, the percentage

still stands at over 60%. With respect to Hoser’s full pub-

lications considered here, approximately 20% {Laudakia

paper 11%) of the text constitutes a verbatim copy from

other sources. In several cases copied sections exceed 100

lines of identical text, and often full descriptions of species

or excerpts from publications concerned with the phyloge-

ny or taxonomy of agamid lizards are repeated word-for-

word.

Hoser’s papers often contain an exhaustive bibliogra-

phy which gives the impression of a properly performed

literature search. However, we found that actually fewer

than 50 publications (out of several hundred referenced)

were used. Three publications used are not referenced at

all, and none of the publications by his fellow Australian

Jane Melville (authored or co-authored) was cited or used,

although these contain phylogenies of Laudakia and Di-

poriphora as well as nomenclatural proposals preceding

those of Hoser. The use of unreferenced material is a clear

breach of commonly accepted editorial standards and

should be avoided by all means. Hoser’s papers should not

have passed any peer review based on the amount of

copied text and in our opinion his work constitutes in sev-

eral cases a form of plagiarism.

Analysing Hoser’s proposed nomenclature we can iden-

tify cases where a name is preoccupied and unavailable,

where a name is being resurrected that was resurrected be-

fore, where names are used that had been very recently

synonymized with other genera but for which the litera-

ture was overlooked or disregarded. In one case Hoser as-

signs a name to a genus that does not include the type

species which he places into another genus. In other cas-

es, he produces nomina nuda or resurrects a nomen obli-

tum incorrectly. He further restricts a type locality with-

out identifying a type specimen from that area and selects

a holotype for a newly described species that has a bifur-

cated tail without any mentioning of this particular fea-

ture.

With respect to taxonomy, in each of the four papers we

find examples of wrong diagnoses, falsely attributed

species, and misinterpretation of previously published tax-

onomic studies. Furthermore, Hoser omitted several gen-

era in his classification schemes (e.g. Harpesaurus, Thau-

matorhynchus, Psammophilus and Tympanocryptis) as

well as many species (e.g. species of Phrynocephalus and

Lophognathus). Presented as they are, Hoser’s taxonom-

ic schemes for the subfamilies Amphibolurinae and Dra-

coninae, as well as his division of the genera Laudakia and

Uromastyx, just constitute a grouping and naming exer-

cise within the confines of a particular published phytoge-

ny he chooses to follow. The slightest changes in these

phylogenies will render them false, in particular as mo-

lecular data have not been used to study all genera and

species under consideration. Hence, Hoser’s taxonomies

and nomenclature acts are highly unstable and little help-

Bonn zoological Bulletin 64 (2): 117-138

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Critical review of Hoser’s publications on agamid lizards

135

ful for species assigment. A herpetologist trying to assign

a newly collected specimen or an existing museum vouch-

er to a specific taxon will still have to look for the origi-

nal or subsequent publications where keys are available,

and would have to revert to Hoser’s paper(s) only to de-

termine his proposed nomenclature - if those names were

available. Kaiser et al. (2013) and Kaiser (2013) suggest-

ed a suppression of all Hoser names to prevent such case

and produced a list with recommended appropriate

names. With respect to the taxa dealt with in the present

paper we propose to suppress Hoser’s names completely

and recommend the usage of generally accepted names

which can be found in the Reptile Database (Uetz & Hosek

2015).

While available names have to be used according to the

Code, a taxonomy does not necessarily have to be accept-

ed. If we were to work in the same way as Hoser does we

could claim here that Gonocephalus mjobergi and Ptyc-

tolaemus share a common character, namely longitudinal

gular folds. We could further claim to consider this a

synapomorphy not shared by the Japalura variegata / Ori-

otiaris group, propose a new tribe excluding Japalura /

Oriotiaris with Ptyctolaemus gularis (Peters, 1 864) as the

type species and name it accordingly. While our tribe

would have a common character to define it, two of

Hoser’s tribe diagnoses, which are defined by their con-

tent rather than common characters, would become in-

valid. Equally we could claim that Hypsicalotes kinabalu-

ensis (de Grijs, 1937) has a unique set of characters (which

it has) that distinguishes the genus from all other Dracon-

inae and remove it from Hoser’s tribe, only to name a new

tribe.

Although this paper is mainly meant to analyze Hoser’s

taxonomy and nomenclature we have to address some is-

sues with regard to the Code. There is no requirement for

a publication to be peer-reviewed or to comply with any

other commonly accepted editorial standards. The ethics

recommended by the Code do not have to be adhered to.

Even a photograph and short description of the coloura-

tion followed by a new name published in a daily news-

paper would qualify as valid and therefore the name would

be considered available. Zoobank is the official registry

of the ICZN. Everyone can register with Zoobank (a vi-

able approach and we hope it will stay like this) and sub-

sequently register nomenclatural acts. However, Zoobank

is not curated and there is no review process in place to

check the correctness of submitted data. This literally in-

vites pure naming exercises by “harvesting” nodes

(nomenclatural vandalism) from a previously published

phylogenetic tree. At the end of November 2015, Hoser

had 873 nomenclatural acts registered with Zoobank,

which on the face of it leads to two different nomencla-

tures for many reptilian taxa as his names are not accept-

ed by the overwhelming majority of the herpetological

community.

The ICZN needs to implement provisions to prevent un-

scientific and unethical publication of nomenclatural pro-

posals to become available. We are convinced that Hoser

is abusing the system. The preceding examples provide

sufficient evidence to demonstrate his abuses. We strong-

ly recommend that the ICZN uses their plenary power to

suppress all of Hoser’s nomenclatural acts published in the

Australasian Journal of Herpetology. We feel that, if this

step is not taken, a large part of the herpetological com-

munity will - with great respect for the ICZN and with

great regret - continue to use the alternative nomenclat-

ural system of the Reptile Database as a reference for

available names.

CONFLICT OF INTEREST

All four authors have submitted taxonomic papers on

agamid lizards including nomenclatural changes without

acknowledging Hoser’s nomenclature or referencing his

publications. Our own publications have been extensive-

ly used verbatim by Hoser without asking for permission

to do so. The authors therefore have a personal interest to

put this on public record.

Acknowledgements. The present publication benefited from

valuable input provided by colleagues in the field of herpetol-

ogy. We are grateful to George Zug and Hinrich Kaiser for a pre-

review of the manuscript and for their comments, corrections and

suggestions.

BIBLIOGRAPHY

Amey AP, Couper PJ, Shea GM (2012) Intellagama lesueurii

(Gray, 1831), the correct binomial combination for the Aus-

tralian Eastern Water Dragon (Sauria, Agamidae). Zootaxa

3390; 65-67

Ananjeva NB, Peters G, Macey JR, Papenfuss TJ (1990) Stel-

lio sacra (Smith 1935) - a distinct species of Asiatic Rock

Agamid from Tibet. Asiatic Herpetological Research 3:

104-115

Baig KJ (1992) Systematic studies of the 5're///o-Group of Aga-

ma (Sauria; Agamidae). PhD Dissertation Quaid-i-Azam Uni-

versity, Islamabad, 287 pp

Baig KJ, Wagner P, Ananjeva NB, Bohme W (2012) A morphol-

ogy-based taxonomic revision of Laudakia Gray, 1845

(Squamata; Agamidae). Vertebrate Zoology 62; 213-260

Beddome RH (1878) Description of a new genus of Tree-Lizards

from the higher ranges of the Anamallays. Proceedings of the

Zoological Society of London 1878; 153-154, pi. XIV

Boulenger GA(1885) Catalogue of the lizards in the British Mu-

seum (Natural History). 1. Geckonidae, Eublepharidae, Uro-

platidae, Pygopodidae, Agamidae. 575 pp. Trustees British

Museum, London

Boulenger GA (1 890) The fauna of British India, including Cey-

lon and Burma. Reptilia and Batrachia. xviii+541pp. Taylor

& Francis, London

Cogger HG (1983) Reptiles and Amphibians of Australia. 660

pp, 3'‘^ ed. Ralph Curtis Publishing, Sanibel, USA

Bonn zoological Bulletin 64 (2); 117-138

©ZFMK

136

Wolfgang Denzer et al.

Cogger HG (1993) Amphibia & Reptilia. 439 pp in; Glasby CG,

Ross GJB, Beesley PL (eds) Fauna of Australia Volume 2A

Part 1,; AGPS Publishing, Canberra. Online (Volume. 2A,

Chapter 23) at www.environment.gov.au/node/13758 (last ac-

cessed 15* July 2015)

Cogger HG (2000) Reptiles and Amphibians of Australia. 808

pp, 6* ed. Ralph Curtis Publishing, Sanibel, USA

Cogger HG (2014) Reptiles and Amphibians of Australia. 1064

pp, 7* ed. CSIRO Publishing, Canberra, Australia

Denzer W, Manthey U (2009) Remarks on the type specimen of

Gonocephalus mjobergi Smith, 1925 (Sauria; Agamidae).

Bonner zoologische Beitrage 56; 255-258

Diong CH, Lim SS (1998) Taxonomic review and morphome-

tric description of Bronchocela cristatella (Kuhl, 1 829) (Squa-

mata; Agamidae) with notes on other species in the genus. The

Raffles Bulletin of Zoology 46; 345-359

Diong CH, Low M-H, Tan E-C, Yong H-S, Hikida T, Ota H

(2000) On the monophyly of the Agamid genus Gonocephalus

Kaup, 1825 (Reptilia; Squamata) -A Chromosomal Perspec-

tive. Current Herpetology 19; 71-79

Doughty P, Kealley L, Melville J (2012) Taxonomic assessment

of Diporiphora (Reptilia; Agamidae) dragon lizards from the

western arid zone of Australia. Zootaxa 3518; 1-24

Dumeril AM, Bibron G (1837) Erpetologie generale ou Histoire

Naturelle complete des Reptiles. ii+572pp. Eibrarie Encyclo-

pedique de Roret, Paris

Fitzinger EJ (1843) Systema Reptilium. Fasciculus Primus. Am-

blyglossae. 106 pp. Braumilller & Seidel Bibliopolas, Wien

Gray JE (1845) Catalogue of the specimens of Eizards in the col-

lection of the British Museum, xxviii+289. British Museum

Natural History, Eondon

Gunther ACEG (1864) The Reptiles of British India, xxvii+452,

pi. 1-26. The Ray Society, London

Hall TA (1999) BioEdit; a user-friendly biological sequence

alignment editor and analysis program for Windows 95/98/NT.

Nucleic Acids Symposium Series 41; 95-98

Hallermann J (2000) A new species of Calotes from the Moluc-

cas (Indonesia), with notes on the biogeography of the genus

(Sauria; Agamidae). Bonner Zoologische Beitrage 49;

155-163

Hallermann J (2004) A new species of the genus Bronchocela

from the tropical Rain Forest of Southern Vietnam. Russian

Journal of Herpetology 11; 30-34

Hallermann J (2005) A taxonomic review of the genus Bron-

chocela (Squamata; Agamidae), with description of a new

species from Vietnam. Russian Journal of Herpetology 12;

168-183

Hallermann J, Bohme W (2000) A review of the genus

Pseudocalotes (Squamata; Agamidae), with description of a

new species from West Malaysia and comments on the gener-

ic concept for South East Asian arboreal agamids. Reptilia-

Amphibia21; 193-210

Hallowell E (1860) Report upon the Reptilia of the North Pa-

cific Exploring Expedition, under command of Capt. John

Rogers, U.S.N. Proceedings of the Academy of Natural Sci-

ence of Philadelphia; 480-509

Harikrishnan S, Vasudevan K (2013) Rediscovery of Calotes an-

damanensis Boulenger, 1891, and assessment of its generic

allocation (Squamata; Sauria; Agamidae). Herpetozoa 26;

3-13

Henle K (1995) A brief review of the origin and use of 'stellio'

in herpetology and a comment on the nomenclature and tax-

onomy of agamids of the gQmxsAgama (sensu lato). Herpeto-

zoa 8; 3-9

Bonn zoological Bulletin 64 (2); 117-138

Hoser RT (2012a) A five- way division of the agamid genus Lau-

dakia Gray, 1845 (Squamata; Sauria; Agamidae). Aus-

tralasian Journal of Herpetology 14; 17-23

Hoser RT (2012b) Robust taxonomy and nomenclature based on

good science escapes harsh factbased criticism, but remains

unable to escape an attack of lies and deception. - Australasian

Journal of Herpetology 14. 37-64

Hoser RT (2013) A seven way division of the Amphibolurinae

(Squamata; Sauria; Agamidae). Australasian Journal of Her-

petology 21; 33-36

Hoser RT (2014a) A break up of the genus Acrochordus Hom-

stedt, 1787, into two tribes, three genera and the description

of two new species (Serpentes; Acrochordidae). Australasian

Journal of Herpetology 22; 2-8

Hoser RT (2014b) A logical new taxonomy for the Asian sub-

family Draconinae based on obvious phylogenetic relation-

ships and morphology of species (Squamata; Sauria; Agami-

dae; Draconinae). Australasian Journal of Herpetology 22;

9-59

Hoser RT (2014c) A long overdue taxonomic rearrangement of

the Uromastycinae (Squamata; Sauria; Agamidae). Aus-

tralasian Journal of Herpetology 23; 54-64

Hoser RT (2015) PRINO (Peer reviewed in name only) journals;

When quality control in scientific publication fails. Aus-

tralasian Journal of Herpetology 26; 3-64

Hugall AF, Lee SY (2004) Molecular Claims of Gondwanan Age

for Australian Agamid Lizards are Untenable. Molecular Bi-

ology and Evolution 21; 2102-2110

Hugall AF, Foster R, Hutchinson M, Lee MS (2008) Phyloge-

ny of Australasian agamid lizards based on nuclear and mi-

tochondrial genes; implications for morphological evolution

and biogeography. Biological Journal of the Linnean Socie-

ty 93; 343-358

ICZN (1999 & 2012) International code of Zoological Nomen-

clature. The Natural History Museum, London

Inger RF (1960) A review of the agamid lizards of the genus

Phoxophrys Hubrecht. Copeia (3); 221-225

Inger RF (1983) Morphological and ecological variation in the

flying lizards (Genus Draco). Fieldiana Zoology New Ser. 18;

1-35

JogerU (1991) A molecular phylogeny of agamid lizards. Copeia

(3); 616-622

Kastle W, Schleich HH (1998) Notes on Comparative Etholo-

gy and Taxonomy of the Genus Japalura, Pp 233-246 in;

Schleich HH, Kastle W (eds.) Contributions to the Herpetol-

ogy of South-Asia (Nepal India). Veroffentlichungen aus dem

Fuhlrott-Museum, Wuppertal

Kaiser H (2013) The Taxon Filter, a novel mechanism designed

to facilitate the relationship between taxonomy and nomen-

clature, vis-a-vis the utility of the Code’s AxUcXq 8 1 (the Com-

mission’s plenary power). Bulletin of Zoological Nomencla-

ture 70; 293-302

Kaiser H, Crother BI, Kelly CMR, Luiselli L, O’Shea M, Ota

H, Passo P, Schleip WD, Wuester W (2013) Best Practises;

In the 2L' Century, Taxonomic decisions in herpetology are

acceptable only when supported by a body of evidence and

published via peer-review. Herpetological Review 44; 8-23

Khan MS (2002) Key and checklist to the lizards of Pakistan

(Reptilia; Squamata; Sauria). Herpetozoa 15; 99-119

Krishnan S (2008) New species of Calotes (Reptilia; Squama-

ta; Agamidae) from the Southern Western Ghats, India. Jour-

nal of Herpetology 42; 530-535

Kuhl H (1820) Beitrage zur Kenntnis der Amphibien, Pp 75-

132 in; Kuhl H (ed.); Beitrage zur Zoologie und vergleichen-

den Anatomic. Hermannsche Buchhandlung, Frankfurt a. M.

©ZFMK

Critical review of Hoser’s publications on agamid lizards

137

Macey JR, AnanjevaNB, Larson A, Rastegar-Pouyani N, Sham-

makov SM, Papenfuss TJ (1998) Phylogenetic Relationships

among Agamid Lizards of the Laudakia caucasia Species

Group; Testing Hypotheses of Biogeographic Fragmentation

and an Area Cladogram for the Iranian Plateau. Molecular

Phylogenetics and Evolution 10: 118-131

Macey JR, Schulte II JA, Larson A, Ananjeva NB, Wang Y,

Pethiyagoda R, Rastegar-Pouyani N, Papenfuss TJ (2000a)

Evaluating Trans-Tethys migration; An example using

acrodont lizard phylogenetics. Systematic Biology 49:

233-256

Macey JR, Schulte II JA, Earson A (2000b) Evolution and phy-

logenetic information content of mitochondrial genomic struc-

tural features illustrated with acrodont lizards. Systematic Bi-

ology 49: 257-277

Macey JR, Schulte II JA, Fong JJ, Das I, Papenfuss TJ (2006)

The complete mitochondrial genome of an agamid lizard from

the Afro-Asian subfamily agaminae and the phylogenetic po-

sition of Bufoniceps and Xenagama. Molecular Phylogenet-

ics and Evolution 39: 881-886

Macleay W (1 884) Notes on some reptiles from the Herbert Riv-

er, Queensland. Proceedings of the Einnean Society New S.

Wales 8: 432-436

Mahony S (2009) A new species of Japalura (Reptilia; Agami-

dae) from northeast India with a discussion of the similar

species Japalura sagittifera Smith, 1940 and Japalura

planidorsata ^QTdon, 1870. Zootaxa2212; 41-61

Mahony S (2010) Systematic and taxonomic revaluation of four

little known Asian agamid species, Calotes kingdonwardi

Smith, 1935, Japalura kaulbacki Smith, 1937, Salea

kakhienensis Anderson, 1879 and the monotypic genus Mic-

topholis Smith, 1935 (Reptilia: Agamidae). Zootaxa2514: 1-

Malkmus R (1994) Herpetologische Beobachtungen am Mt. Ki-

nabalu, Nord-Bomeo. IV. Mitteilungen des Zoologischen Mu-

seums Berlin 70; 217-249

Manthey U, Denzer W (1991) Die Echten Winkelkopfagamen

der Gattung Gonocephalus Kaup (Sauria; Agamidae) 1. Die

megalepis-Gm^pe mit Gonocephalus lacunosus sp. n. aus

Nord-Sumatra. Sauria 13: 3-10

Manthey U, Denzer W (1991) Die Echten Winkelkopfagamen

der Gattung Gonocephalus Kaup (Sauria: Agamidae) IT All-

gemeine Angaben zur Biologie und Terraristik. Sauria 13: 19-

Manthey U, Denzer W (1991a) Die Echten Winkelkopfagamen

der Gattung Gonocephalus Kaup (Sauria: Agamidae) III. Go-

nocephalus grandis {Grny, 1845). Sauria, 13: 3-10

Manthey U, Denzer W (1992) Die Echten Winkelkopfagamen

der Gattung Gonocephalus Kaup (Sauria; Agamidae) IV. Go-

nocephalus mjoebergi Smith, 1925 und Gonocephalus robin-

50W/7 Boulenger, 1908. Sauria 14: 15-19

Manthey U, Denzer W (1993) Die Echten Winkelkopfagamen

der Gattung Gonocephalus Kaup (Sauria: Agamidae) VI. Die

chamaeleontinus-GruTppe. Sauria 15; 23-28

Manthey U, Denzer W (2000) Description of a new genus, Hyp-

sicalotes gen nov. (Sauria; Agamidae) from Mount Kinabalu,

North Borneo, with remarks on the generic identity of Gono-

cephalus schultzewestrumi Urban, 1999. Hamadryad 25;

13-20

Manthey U, Denzer W (2006) A revision of the melanesian-aus-

tralian Angle Head lizards of the genus Hypsilurus (Sauria;

Agamidae: Amphibolurinae), with description of four new

species and one new subspecies. Hamadryad 30: 1-40

Manthey U, Grossmann W (1997) Amphibien & Reptilien Sud-

ostasiens. 512 pp. Natur und Tierverlag, Munster

Bonn zoological Bulletin 64 (2); 117-138

Manthey U (20 1 0) Agamid Eizards of Southern Asia - Dracon-

inae 2, Eeiolepidinae. 168 pp, Chimaira, Frankfurt/M

McGuire JA, Alcala AC (2000) A taxonomic revision of the fly-

ing lizards (Iguania: Agamidae: Draco) of the Philippine Is-

lands, with a description of a new species. Herpetological

Monographs 14: 81-138

McGuire JA, Kiew BH (2001) Phylogenetic systematics of

Southeast Asian flying lizards (Iguania: Agamidae; Draco) as

inferred from mitochondrial DNA sequence data. Biological

Journal of the Einnaean Society 72: 203-229

McGuire JA, Dudley R (2005) The Cost of Eiving Earge; Com-

parative Gliding Performance in Flying Eizards (Agamidae:

Draco). The American Naturalist 166: 93-106

McGuire JA, Brown R, Mump ini RA, Andayani N (2007) The

Flying Eizards of the Draco lineatus group (Squamata: Igua-

nia: Agamidae): A taxonomic revision with descriptions of two

new species. Herpetological Monographs 21: 179-212

Melville J, Hale J, Mantziou G, Ananjeva NB, Milto K (2009)

Historical biogeopgraphy, phylogenetic relationships and in-

traspecific diversity of agamid lizards in the Central Asian

deserts of Kazakhstan and Uzbekistan. Molecular Phylogenet-

ics and Evolution 53: 99-112

Melville J, Ritchie EG, Chappie SN, Glor RE, Schulte II JA

(2011) Evolutionary origins and diversification of dragon

lizards in Australia’s tropical savannas. Molecular Phyloge-

netics and Evolution 58; 257-270

Mocquard F (1890) Recherches sur la faune herpetologique des

lies de Borneo et de Palawan. Nouvelles Archives du Museum

National d’Histoire Naturelle, Ser. 3: 115-168, 5pls

Moody SM (1980) Phylogenetic and historical biogeographical

relationship of the genera in the family Agamidae (Reptilia:

Eacertilia). Ph.D. Thesis, University of Michigan, 373 pp

Musters CJ (1983) Taxonomy of the genus Draco E. (Agami-

dae, Eacertilia, Reptilia). Zoologische Verhandelingen 199;

1-120

Ota H (1988) Karyotypic differentiation in an agamid lizard,

Japalura swinhonis swinhonis. Experientia 44: 66-68

Peters W, Doria G (1878) Catalogo die rettili e die Batraci ra-

coolti da O. Beccari, E.M. D’Albertis e A.A. Bruijn nella sot-

to-regione Austro-Malese. Annali del Museo Civico di Sto-

riaNaturale di Genova ser. 1: 323-450, pis 1-7

Pethiyagoda R, Manamendra-Arachchi K (1998) A revision of

the endemic Sri Eankan agamid lizard genus Ceratophora

Gray, 1835, with description of two new species. Journal of

South Asian natural History 3: 1-50

Pyron RA, Burbrink FT, Wiens JJ (2013) A phylogeny and re-

vised classification of Squamata, including 4161 species of

lizards and snakes. BMC Evolutionary Biology 13; 1-53

Rooij N de (1915) The reptiles of the Indo - Australian Archi-

pelago. 1. Eacertilia, Chelonia, Emydosauria. xiv+384 pp, Tay-

lor & Francis, Eeiden

Schulte II JA, Macey JR, Pethiyagoda R, Earson A (2002) Ros-

tral horn evolution among Agamid Eizards of the genus Cer-

atophora endemic to Sri Eanka. Molecular Phylogenetics and

Evolution 22; 111-117

Schulte II JA, Melville J, Earson A (2003) Molecular phyloge-

netic of ancient divergence of lizard taxa on either side of the

Wallace’s Eine. Proceedings of the Royal Society of Eondon

B270: 597-603

Schulte II JA, Vindum JV, Win H, Thin T, Twin KS, Shein AK

(2004) Phylogenetic relationships of the genus Ptyctolaemus

(Squamata; Agamidae), with a description of a new species

from the Chin Hills of Western Myanmar. Proceedings of the

California Academy of Sciences 55; 222-247

©ZFMK

138

Wolfgang Denzer et al.

Smith, MA (1935) The fauna of British India, Ceylon and Bur-

ma. Reptilia and Amphibia. Vol. II Sauria. xiii+440 pp, Tay-

lor & Francis, London

Stejneger, L (1907) Herpetology of Japan and adjacent territo-

ry. Bulletin of the United States National Museum 58: xx+577

Stoliczka, F (1870) Observations on some Indian and Malayan

Amphibia and Reptilia. Journal of the Asiatic Society of Ben-

gal 39: 124-228

Stuart-Fox DM, Owens IFF (2003) Species richness in agamid

lizards: chance, body size, sexual selection or ecology? Jour-

nal of Evolutionary Biology 16: 659-669

Swofford DL (2002) PAUP*. Phylogenetic Analysis Using Par-

simony (*and Other Methods). Version 4.0b 10. Sinauer Asso-

ciates, Sunderland, Massachusetts, USA

Theobald W (1876) Descriptive catalogue of the reptiles of

British India, xiii+238 pp + xxxviii+xii. Thacker Spink & Co,

Calcutta (Kolkata)

Thomson S (2014) Comment on Spracklandus Hoser, 2009

(Reptilia, Serpentes, Elapidae): request for confirmation of the

availability of the generic name and for nomenclatural vali-

dation of the journal in which it was published (Case 3601).

Bulletin of Zoological Nomenclature 71: 133-135

Uetz P, Hosek J (eds): The Reptile Database. Online at

http://www.reptile-database.org last accessed July 30, 2015

Wells RW, Wellington CR (1983) A synopsis of the class Rep-

tilia in Australia. Agamidae. Australian Journal of Herpetol-

ogy 1: 73-129

Wells RW, Wellington CR (1985) A classification of the Amphib-

ia and Reptilia of Australia. Australian Journal of Herpetol-

ogy, Suppl. Ser. (1): 64 pp

Wilms TM, Bohme W (2007) Review of the taxonomy of the

spiny-tailed lizards of Arabia (Reptilia: Agamidae: Eeiolepid-

inae: Uromastyx. Fauna of Arabia 23: 435-468

Wilms TM, Bohme W, Wagner P, Eutzmann N, Schmitz A (2009

“2007”) On the Phytogeny and Taxonomy of the Genus Uro-

mastyx Merrem, 1820 (Reptilia: Squamata: Agamidae: Uro-

mastycinae) - Resurrection of the Genus Saara Gray, 1845.

Bonner zoologische Beitrage 56: 55-99

Zug GR, Brown HH, Schulte II JA, Vindum JV (2006) System-

atics of the Garden Eizards, CaJotes versicolor Group (Rep-

tilia, Squamata, Agamidae), in Myanmar: Central Dry Zone

Populations. Proceedings of the California Academy of Sci-

ences, Ser. 4, 57: 35-68

Bonn zoological Bulletin 64 (2): 117-138

©ZFMK

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Book Review

Lanza B, Funaioli U & Riccucci M (2015) The Bats of

Somalia and Neighbouring Areas. Frankfurt Contributions

to Natural History 60, Edition Chimaira, Frankfurt/M., 566

pp. ISBN 978-3-89973-447-8. Price: € 78,-.

In this voluminous book the authors cover every aspect

of the 79 bat species they found associated with Somalia

and adjacent areas. There are introductory chapters to the

order Chiroptera (with key to the two suborders), subor-

der Megachiroptera, followed by suborder Microchi-

roptera with a key to the families, then family with key

to the genera, and genus with keys to the species, before

each species is discussed individually. Each of these chap-

ters follows a similar structure with “derivatio nominis”

(name derivations and etymology), “composition and dis-

tribution”, and “distinctive characters”. Family and genus

chapters add paragraphs on “type genus [species]”,

“iconography” (= a listing of the figures associated with

that chapter), “synonymy” and “common names”, “tax-

onomy”, “echolocation calls,” where available, “biology”,

“Somali records” and “Personal observations.” Within this

pattern the authors provide a great amount of detail on the

species, all illustrated with a color drawing of the bat with

one wing outstretched (sometimes replaced by color pho-

tographs of mounted specimens), up to three (!) sets of

pen-and-ink skull drawings per species, nicely showing

individual variation, and variously photographs or draw-

ings of penis, bacula, palatal ridges, or details of denti-

tion, skull, tragi or noseleaves depending on the family’s

special characteristics. Quite a bit of information is con-

tained in the name derivations, from the Greek and Eat-

in roots to interesting details on naturalists or hunter-col-

lectors the bat may be named after, and - where available

- the bat’s biology. Common names listed include Eng-

lish and Italian names but no local Somali names. Well-

organized tables identify many of the individual measure-

ments of the 3650 museum specimens examined, all giv-

en with museum acronyms and numbers, and summary

statistics.

At the end of the book, under “Addenda,” two species,

Kerivoula smithii and Laephotis wintoni are added as “oc-

curring in areas nearby Somalia”, followed by 32 detailed

distribution maps, some of which covering more than one

species, with numbered point localities, all of which iden-

tified by name (!) in separate legends. This is then fol-

lowed by the Acknowledgements, a detailed six-page

Gazetteer with many useful spelling variants and 16 pages

of References including literature as recent as some of the

species entries from Happold and Happold’s 2013 bat vol-

ume in the new Mammals of Africa series, which is great-

ly complemented by this new book.

As if this wasn’t enough, the book concludes with a sep-

arate section by Funaioli and Eanza entitled “An outline

of the geography of Somalia” that provides the smaller dis-

tribution maps in the main section with much more detail

Benedetto [.anza

Ugo Funaioli

Marco Riccucci

The Bats of Somalia

and Neighbouring Areas

regarding political subdivisions, average yearly rains, and

a “Sketch map of the vegetation [zones] of the Somali

Democratic Republic,” the latter nicely illustrated with one

black & white and 17 color photographs and some more

general information on flora and fauna of Somalia. A sep-

arate reference section concludes this chapter, which is es-

pecially valuable, as it is next to impossible to travel safe-

ly in Somalia at this time.

If there was ever a modem “Rolls Royce”- equivalent

of a “Bats of ...’’-book, with all the possible bells and whis-

tles, this must be it. Yet, the publisher. Edition Chimaira,

somehow managed to accommodate every lavish aspect

of this book and seemingly all special wishes of the au-

thors in a still portable size, printed on durable glossy pa-

per, which does justice to all the different types of illus-

trations, but keeps the price for this rather specialized

books within reasonable limits. May this volume find its

way to all those interested in Somalia or the fauna of the

Horn of Africa in general and bats in particular, and may

it serve as an exemplarily illustrated and detailed account

for this part of Africa, both areas where the bat-volume

of Kingdon’s Mammals of Africa falls short.

Jan Decher, Section of Mammalogy, ZFMK

Bonn zoological Bulletin 64 (2); 139