Bonn zoological Bulletin 68 (2): 183-187

2019 - Breitling R.

https://do1.org/10.20363/BZB-2019.68.2.183

ISSN 2190-7307

http://www.zoologicalbulletin.de

Research article

urn:|sid:zoobank.org: pub: FO246630-F65F-4FF 1-BF4E-008D483BDA2C

Euophrys petrensis C. L. Koch, 1837

is a genuine member of the genus 7alavera (Araneae: Salticidae)

Rainer Breitling

Faculty of Science and Engineering, University of Manchester, Manchester MI 7DN, UK

* Corresponding author: Email: rainer.breitling@manchester.ac.uk

urn:|sid:zoobank.org:author: 1 7A3B585-0E06-436C-A99A-1C7F24DC88D7

Abstract. The small jumping spider Euophrys petrensis C. L. Koch, 1837 combines morphological characters of both

Euophrys s. str. and Talavera, and its generic placement has consequently been contentious. After many years of being

placed in Talavera, the species has recently been transferred back to Euophrys. Here, public DNA barcoding data are used

to confirm that the species should be placed in the genus 7a/avera, as T: petrensis, stat. rev., as is also indicated by several

putative morphological synapomorphies identified earlier.

Key words. Araneae, DNA barcoding, phylogenetic systematics.

INTRODUCTION

The taxonomic placement of Euophrys petrensis has been

problematic for some time, since the revision and major

expansion of the genus 7Zalavera Peckham & Peckham,

1909, by Logunov (1992). Logunov (1992) transferred

four Palaearctic members of Euophrys s. lat. to Talavera

(T. aequipes, T: monticola, T. thorelli, and T: trivittata).

However, he delayed the transfer of their close relative,

E. petrensis, as the latter showed ambiguous characters

implying a possible closer affinity to Euophrys s. str. (e.g.,

chitinous rings in the epigynum and twisted insemination

ducts, as well as a pronounced sexual dimorphism). Lo-

gunov (1992) also remarked that 7’ aequipes occupies a

similar morphologically intermediate position between

Talavera and Euophrys s. str.

Logunov et al. (1993) then indicated that all species

of their petrensis group of Euophrys s. lat. (E. petrensis,

E. aequipes, and E. thorelli) should be included in the

genus Jalavera, without, however, providing additional

arguments regarding EF. petrensis itself.

The transfer of the latter was formalized by Zabka

(1997; see also Zabka & Proszynski 1998), and Talavera

petrensis was generally accepted as the valid combination

by subsequent authors. Logunov & Kronestedt (2003) re-

viewed Zalavera s. lat. and list the diagnostic characters

of the expanded genus: absence of a tibial apophysis; en-

dite tooth on the male maxilla; long white/red hairs on

the base of the cymbium; clearly exposed embolus-tegu-

lum membrane; thin, thread-like insemination ducts; and

scales with a well-marked keel on carapace and abdomen.

However, most recently, Proszynski et al. (2018) trans-

ferred T. petrensis back to Euophrys, but maintained

Received: 29.05.2019

Accepted: 30.07.2019

other members of the petrensis group (sensu Logunov

et al. 1993), such as 7) aequipes and T: thorelli, within

Talavera. This re-transfer was based on Proszynski’s

non-cladistic approach combined with a different relative

weighting of the various characters already highlighted as

ambiguous by Logunov (1992): the coiled embolus and

colourful frontal hairs of the male. The absence of a tib-

ial apophysis, which Logunov & Kronestedt (2003) de-

scribe as one of the most important diagnostic characters

of Zalavera, was considered as non-informative, as the

apophysis in Euophrys s. str. is typically highly reduced,

and the thin copulatory ducts were considered an artefact

of observation by optical microscopy, while “they appear

much broader” when observed by scanning electron mi-

croscopy (Proészynski et al. 2018); this latter argument is

not quite convincing, being based on a rather subjective

comparison of published figures. The other characters

discussed by Logunov & Kronestedt (2003) received no

further attention.

Given this controversial history, it was interesting to

examine if the publicly available barcoding sequences

could be used to supplement the morphological data to

resolve the placement of Euophrys petrensis, in analogy

to the approach taken in Breitling (2017, 2019). In a bar-

coding study of German spiders, the species had already

been shown as sister of Zalavera aequipes, rather than

Euophrys frontalis (Astrin et al. 2016: Supplementary

Figures S1 and S82): as these results were based on a lim-

ited set of species and were not further discussed in the

article, the robustness of this relationship remained un-

clear, but the results certainly indicated that the barcode

data should contain relevant phylogenetic information.

Corresponding editor: B. Huber

Published: 02.09.2019

184 Rainer Breitling

MATERIAL AND METHODS

All public Cytochrome Oxidase Subunit 1 5’ region bar-

codes (COI-5P) for members of Euophrys, Talavera and

Pseudeuophrys (i.e. Euophrys s. \at.), and the closely re-

lated genus Chalcoscirtus were downloaded in FASTA

format from the BOLD database (www.boldsystems.org:

Ratnasingham & Hebert 2007) in April 2018. Sequences

were aligned in BioEdit v7.2.5 (Hall 1999), and trailing

gaps and particularly short sequences were removed to

maximize the amount of sequence positions for which all

specimens in the analysis had data available. The result-

ing dataset included 540 residues of the COI-5P barcode

for 59 specimens. The BOLD and GenBank accession

numbers of all sequences in the final dataset are shown

in Figure 1.

The data included barcode sequences for four species

of Talavera (13 specimens of the type species, 7? minuta,

two specimens of 7! petrensis and one specimen each of

T: thorelli and T: aequipes). Together, these four species

represent all four species groups of 7alavera s. lat. de-

fined by Logunov & Kronestedt (2003). Also included

in the dataset are representatives of Euophrys s. str. (17

specimens of the type species, EF. frontalis, and five of

E. monadnock), of Pseudeuophrys (seven specimens of

the type species, P. erratica, six specimens of P. lanig-

era, and two of P. obsoleta), and two species of Chal-

coscirtus (two specimens of C. alpicola, and three of C.

carbonarius), which in preliminary analyses of salticid

barcodes were consistently nested within Euophrys (Bre-

itling 2019).

The plausibility of the identification of the 7’ petrensis

specimens critical for this analysis was confirmed using

the data deposited in BOLD: the material was collected in

Saxony-Anhalt, well within the known range of the spe-

cies (which extends from Ireland to Central Asia and from

Finland to the Mediterranean), identified by Karl-Hinrich

Kielhorn, an experienced arachnologist, and is deposit-

ed in the Zoologische Staatssammlung Munchen. Both

specimens are adult males, which are particularly easy to

identify reliably, and the habitus photographs available in

BOLD further support the identification.

Phylogenetic trees were reconstructed using phylog-

eny.fr (Dereeper et al. 2008) as described in Breitling

(2017), using the default workflow and parameters to in-

fer Maximum Likelihood (PhyML 3.1; Guindon & Gas-

cuel 2003), Maximum Parsimony (TNT 1.1; Goloboff et

al. 2008) and Bayesian trees (MrBayes 3.2.6; Ronquist

& Huelsenbeck 2003). Sequences were not partitioned

by codon for the analysis. For the Bayesian analysis, the

standard (4by4) model of nucleotide substitution was

used, while the rate variation across sites was set to “in-

vgamma”. Four Markov Chain Monte Carlo chains were

run for 100 000 generations, sampling every 100 gen-

erations, with the first 1000 sampled trees discarded as

“burn-in”. Finally, a 50% majority rule consensus tree

Bonn zoological Bulletin 68 (2): 183-187

was constructed. For the Maximum Likelihood analysis,

the default substitution model was selected assuming an

estimated proportion of invariant sites (of 0.577) and 4

gamma-distributed rate categories to account for rate het-

erogeneity across sites. The gamma shape parameter was

estimated directly from the data (gamma=1.268). Branch

support values are based on an Approximate Likeli-

hood-Ratio Test (aLRT; Anisimova & Gascuel 2006) for

the Maximum Likelihood results, on 1000 bootstrap rep-

licates for the Maximum Parsimony and Neighbour Join-

ing results, and on posterior probabilities for the Bayes-

ian analysis. Trees were visualized using 1TOL (Letunic

& Bork 2016) and annotated in Adobe Illustrator.

RESULTS & DISCUSSION

The barcoding data provide unambiguous support for the

placement of Euophrys petrensis C. L. Koch, 1837, in the

genus Talavera, as T. petrensis, stat. rev. The maximum

interspecies distance between barcodes within the genus

Talavera is 8.5% (between one of the 7! petrensis speci-

mens and 7! aequipes). In contrast, the minimum distance

between 7alavera and any member of Euophrys is 11.4%

(between one of the 7? petrensis specimens and one spec-

imen of E. monadnock). The minimum distance between

T: petrensis and the type species of Euophrys, E. frontalis,

is 12.2%, and the closest member of Pseudeuophrys (P.

obsoleta) has a distance of 12.9%. This is clearly a much

larger barcoding distance, especially when considering

that, on average, barcode sequences for species assigned

to the same genus differ by 10.5% among Salticidae in

the BOLD database, and the numbers are even lower

for other families (e.g., 10.0% for Thomisidae, 8.5% for

Theridiidae, and 6.2% for Lycosidae; Breitling, unpubl.

data). To give a sense of the scale of this difference: of

the 72 base pairs shared between 7? petrensis and either

7. minuta or E. frontalis (but not both), 16 are shared with

E. frontalis, and 56 with T. minuta. Assuming a uniform

mutation rate (as would appear reasonable for such a

closely related group), this finding is obviously not com-

patible with a closer relationship between 7: petrensis

and E. frontalis. Consequently, all the tree reconstruction

methods recover a monophyletic genus 7alavera, includ-

ing T. petrensis, with strong bootstrap and posterior prob-

ability support (99% bootstrap support in the Maximum

Parsimony tree, 100% bootstrap support in the Neigh-

bour Joining analysis, 100% posterior probability in the

Bayesian analysis, and 100% approximate LRT in the

Maximum Likelihood tree), and almost always with the

same internal topology (7 petrensis as sister to the other

three species), as shown in Figure 1. The corresponding

node is the most strongly supported interspecific node in

the entire tree, with higher support values than the also

highly supported node joining the three Pseuwdeuophrys

species. None of the trees indicates a closer relationship

©ZFMK

Tree scale: 0.01 —— 4

Talavera petrensis 185

-=FBARB291-11 Talavera petrensis COI-5P KX537369

= FBARB290-11 Talavera petrensis COI-5P KX537012

SMNKAO25-16 Talavera aequipes COI-5P

NOARAO49-11 Talavera thorelli COI-5P

SPITO297-14 Talavera minuta COI-5P KP656616

CNGBE785-14 Talavera minuta COI-5P KP645979

UAMIC1793-14 Talavera minuta COI-5P KU876297

SPITO298-14 Talavera minuta COI-5P KP652374

RRSSC7097-15 Talavera minuta COI-5P MG044579

RRSSC7092-15 Talavera minuta COI-5P MG043543

SSGBC553-14 Talavera minuta COI-5P MF814838

CNTNE1835-14 Talavera minuta COI-5P KP651965

SSBAD5950-13 Talavera minuta COI-5P KM838862

SSBAD5946-13 Talavera minuta COI-5P KM837257

SSBAD5936-13 Talavera minuta COI-5P KM826638

ARONT803-10 Talavera minuta COI-5P HQ924648

ARBCM173-13 Talavera minuta COI-5P KP649108

SPSLO316-13 Pseudeuophrys lanigera COI-5P KX039358

GBBSP748-15 Pseudeuophrys lanigera COI-5P KY270088

GBBSP907-15 Pseudeuophrys lanigera COI-5P KY268896

GBBSP1726-15 Pseudeuophrys lanigera COI-5P KY268631

GBBSP014-15 Pseudeuophrys lanigera COI-5P KY268633

FBARB577-12 Pseudeuophrys lanigera COI-5P KX537344

SPIEU1281-11 Pseudeuophrys obsoleta COI-5P

GBBSP816-15 Pseudeuophrys obsoleta COI-5P KY270439

FBARB544-11 Pseudeuophrys erratica COI-5P KX537310

GBBSP210-15 Pseudeuophrys erratica COI-5P KY269279

FBARB845-14 Pseudeuophrys erratica COI-5P KX537290

FBARB075-11 Pseudeuophrys erratica COI-5P JN299225

SMNSAO06-16 Pseudeuophrys erratica COI-5P

NOARAO65-11 Pseudeuophrys erratica COI-5P

FBARB914-14 Pseudeuophrys erratica COI-5P KX537007

SSJAB2041-13 Chalcoscirtus alpicola COI-5P KM824738

SPIRU1282-11 Chalcoscirtus alpicola COI-5P KF367929

ARBCM176-13 Chalcoscirtus carbonarius COI-5P KP653096

ARBCM2083-14 Chalcoscirtus carbonarius COI-5P KP649199

ARBCM2080-14 Chalcoscirtus carbonarius COI-5P KP653975

PRMA988-12 Euophrys monadnock COI-5P KP655713

SPRMA776-12 Euophrys monadnock COI-5P KP647526

ARBCM2082-14 Euophrys monadnock COI-5P KP647452

CNGRD1300-12 Euophrys monadnock COI-5P KM824291

CNGRD1299-12 Euophrys monadnock COI-5P KM839479

10010) —_—

r 0099 Pes

SMNSAO056-16 Euophrys frontalis COI-5P

SMNKA103-16 Euophrys frontalis COI-5P

SMNKA0O21-16 Euophrys frontalis COI-5P

SMNKA0O20-16 Euophrys frontalis COI-5P

NLARA023-12 Euophrys frontalis COI-5P

GBBSP1283-15 Euophrys frontalis COI-5P KY270204

FBARBO76-11 Euophrys frontalis COI-5P JN299226

FBARB256-11 Euophrys frontalis COI-5P KX536888

MNKA105-16 Euophrys frontalis COI-5P

SMNKA104-16 Euophrys frontalis COI-5P

SMNKA022-16 Euophrys frontalis COI-5P

GBBSP381-15 Euophrys frontalis COI-5P KY269856

GBBSP240-15 Euophrys frontalis COI-5P KY268958

GBBSP1800-15 Euophrys frontalis COI-5P KY268665

GBBSP067-15 Euophrys frontalis COI-5P KY268626

FBARB393-11 Euophrys frontalis COI-5P KX537345

GBBSP976-15 Euophrys frontalis COI-5P KY268541

Fig. 1. DNA-Barcode-based tree of Zalavera and its relatives. The topology and branch lengths are based on the Neighbour Joining

tree. The genus 7alavera forms a very strongly supported monophyletic group, including 7: petrensis, in all four tree reconstruc-

tions, but its relationships to the other members of Euvophrys s. lat. are not convincingly resolved in the dataset. Black circles indi-

cate more than 95% posterior probability (MB), approximate likelihood (ML), or bootstrap support (MP, NJ); grey circles indicate

support values between 50% and 95%. White circles indicate branches that were either not recovered or had a support below 50%.

Support values for intraspecific branches are not shown (all species were recovered as monophyletic in all analyses).

of 7: petrensis to Euophrys s. str. or Pseudeuophrys with

any degree of support. This result is consistent with the

barcode-based trees of German spider species shown in

the Supplementary Material of Astrin et al. (2016).

As expected, the details of the relationship between

Talavera, Euophrys s. str. and Pseudeuophrys are not un-

ambiguously resolved by the barcode data. In the Bayes-

ian and Maximum Likelihood analyses there is some

Bonn zoological Bulletin 68 (2): 183-187

evidence that the three genera are part of a (monophy-

letic) clade (Euophrys s. lat.), as would be expected on

the basis of their morphological similarity, and Pseu-

doeuophrys is consistently recovered as monophyletic as

well in all analyses. The diversity of Euophrys s. str. 1s

not sufficiently represented to assess its monophyly, and

its relationship with Chalcoscirtus is not convincingly

resolved. In several of the reconstructed trees, Chalco-

©ZFMK

186 Rainer Breitling

scirtus is nested within a paraphyletic Euophrys s. str.

As Chalcoscirtus appears to be represented only by fe-

male specimens in the database, it is not impossible that

this finding is due to a misidentification. But, as the case

involves multiple specimens from several locations, in-

cluding material identified by an experienced arachnol-

ogist, Gergin Blagoev, the possibility that Chalcoscirtus

forms a highly derived monophyletic subgroup within a

paraphyletic Euophrys s. str. as presently defined cannot

be discarded outright. The genus Chalcoscirtus is defined

by a number of characteristic synapomorphies, such as

the dark, strongly sclerotized, glabrous prosoma, the

male opisthosomal scutum and the absence of retromar-

ginal cheliceral teeth (Cutler 1990). Its key diagnostic

character is the presence of a ventral tibial apophysis, but

the absence of this apophysis in Euophrys is a symplesio-

morphy, shared with Pseudeuophrys and Talavera. Thus,

there are currently no convincing synapomorphies for

Euophrys s. str., and the genitalia of the two genera are

obviously very similar in both sexes. Of course, it would

also be possible that Chalcoscirtus is polyphyletic, and

that the two species in the dataset are not closely relat-

ed to Chalcoscirtus s. str., 1.e., C. infimus and its mostly

Central Asian close relatives. However, the large num-

ber of convincing synapomorphies supporting the genus

makes this hypothesis much less plausible.

It is striking that the barcoding data allow such a clear

assignment of “Euophrys”’ petrensis to the genus Tala-

vera. The ambiguous morphological data had indicated

a more “intermediate” position, 1.e., a trichotomy or very

shallow branching between Euophrys, Talavera s. str. and

T: petrensis, which could have been reflected in a much

smaller (or non-existent) difference in barcoding distance

and possibly an unresolved or ambiguous placement of 7°

petrensis in the various phylogenetic reconstructions. The

molecular evidence can stimulate a renewed look at the

morphological data as well. It appears that the absence

of a tibial apophysis, presence of an endite tooth on the

maxilla, exposed embolus—tegulum membrane, long red

and white hairs at the base of the cymbium, and keeled

scales are indeed synapomorphies of 7alavera (and, in

the case of the missing tibial apophysis, its supposed

sister group Zanzania, Logunov & Kronestedt 2003, sub

Lilliput), while the coiled embolus, relatively thick and

twisted insemination duct, and pronounced sexual dimor-

phism, including colourful cymbial setae, are possible

symplesiomorphies maintained in the basal 7’ petrensis,

and shared with, e.g., Euophrys, but lost in the other 7a-

lavera species, as already suggested by Logunov (1992).

More derived 7alavera species (beyond the petrensis and

aequipes group) are then additionally characterized by a

number of derived characters of the male pedipalp (e.g.,

a more complex sperm duct, a chitinous ligament con-

necting the embolus and the tegulum, and a distal tegular

sclerite; Logunov & Kronestedt 2003).

Bonn zoological Bulletin 68 (2): 183-187

Of course, the fact that we have been able to refute

one of the two alternative hypotheses that we began with

does not necessarily mean that the other hypothesis is

correct. Thus, while the barcode data do not provide any

reason to doubt the taxonomic placement suggested by

Logunov & Kronestedt (2003), future studies may well

change this assessment. Additional analyses, using larger

datasets, will also be required to determine the precise

relationships between the various Euophryine genera,

including Chalcoscirtus. The success of barcode infor-

mation in suggesting an unambiguous solution to the

taxonomic conundrum regarding the generic placement

of Talavera petrensis should encourage the further use

of public barcoding databases as a valuable resource to

complement morphological approaches to spider taxono-

my, especially in cases where morphological analysis has

resulted in ambiguous or controversial placements.

Acknowledgements. I thank Jonas Astrin, Dmitri Logunovy,

Wayne Maddison, Junxia Zhang and an anonymous referee for

their detailed and critical comments on the manuscript, which

have helped improving the argument considerably.

REFERENCES

Anisimova M, Gascuel O (2006) Approximate likelihood-ratio

test for branches: A fast, accurate, and powerful alternative.

Systematic Biology 55(4): 539-552

Astrin JJ, Hofer H, Spelda J, Holstein J, Bayer S, Hendrich L,

Huber BA, Kielhorn KH, Krammer HJ, Lemke M, Monje

JC, Moriniere J, Rulik B, Petersen M, Janssen H, Muster C

(2016) Towards a DNA barcode reference database for spi-

ders and harvestmen of Germany. PLoS ONE 11 (e€0162624):

1-24. https://journals.plos.org/plosone/article?id=10.1371/

journal.pone.0162624

Breitling R (2017) Public DNA barcoding data resolve the

status of the genus Arboricaria (Araneae: Gnaphosidae).

Arachnologische Mitteilungen 54: 24-27. https://arages.

de/105431/aramit5405

Breitling R (2019) Barcode taxonomy at the genus level. Eco-

logica Montenegrina 21: 17-37

Cutler B 1990 A revision of the western hemisphere Chalcos-

cirtus (Araneae: Salticidae). Bulletin of the British Arachno-

logical Society 8: 105-108

Dereeper A, Guignon V, Blanc G, Audic S, Buffet S, Chevenet

F, Dufayard JF, Guindon S, Lefort V, Lescot M, Claverie JM,

Gascuel O (2008) Phylogeny.fr: robust phylogenetic analy-

sis for the non-specialist. Nucleic Acids Research 36 (Web

Server issue): W465—W469. https://www.ncbi.nlm.nih.gov/

pubmed/1 8424797

Goloboff PA, Farris JS, Nixon KC (2008) TNT, a free pro-

gram for phylogenetic analysis. Cladistics 24: 774-786.

https://onlinelibrary.wiley.com/doi/abs/10.1111/j.1096-

0031.2008.00217.x

Guindon S, Gascuel O (2003) A simple, fast and accurate al-

gorithm to estimate large phylogenies by maximum like-

lihood. Systematic Biology 52: 696-704. doi: https://doi.

org/10.1080/10635150390235520

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

©ZFMK

Talavera petrensis 187

Letunic I, Bork P (2016) Interactive tree of life ((TOL) v3: an

online tool for the display and annotation of phylogenetic

and other trees. Nucleic Acids Research 44: W242—W245,

https://www.ncbi.nlm.nih.gov/pubmed/27095 192

Logunov DV (1992) Definition of the spider genus Talavera

(Araneae, Salticidae), with a description of a new species.

Bulletin de l’Institut Royal des Sciences Naturelles de Bel-

gique (Ent.) 62: 75-82

Logunov DV, Cutler B, Marusik YM (1993) A review of the

genus Euophrys C. L. Koch in Siberia and the Russian Far

East (Araneae: Salticidae). Annales Zoologici Fennici 30:

101-124

Logunov DV, Kronestedt T (2003) A review of the genus Za-

lavera Peckham and Peckham, 1909 (Arareae, Salticidae).

Journal of Natural History 37: 1091-1154

Proszynski J, Lissner J, Schafer M (2018) Taxonomic survey

of the genera Euophrys, Pseudeuophrys and Talavera, with

description of Euochin gen. n. (Araneae: Salticidae) and with

proposals of a new research protocol. Ecologica Montenegri-

na 18: 26-74

Bonn zoological Bulletin 68 (2): 183-187

Ratnasingham S, Hebert PDN (2007) BOLD: The Barcode of

Life Data system (www. barcodinglife.org). Molecular Ecol-

ogy Notes 7: 355-364. https://onlinelibrary.wiley.com/do1/

full/10.1111/j.1471-8286.2007.01678.x

R Core Team (2018) R: A language and environment for sta-

tistical computing. R Foundation for Statistical Computing,

Vienna, Austria. https://www.R-project.org/

Ronquist F, Huelsenbeck JP (2003) MrBayes 3: Bayesian phy-

logenetic inference under mixed models. Bioinformatics

19 (12): 1572-1574

Wright ES (2016) Using DECIPHER v2.0 to Analyze Big Bi-

ological Sequence Data in R. The R Journal 8 (1): 352-359

Zabka M (1997) Salticidae: Pajaki skaczace (Arachnida: Ara-

neae). Fauna Polski 19: 1-188

Zabka M, Proszyhski J (1998) Middle European Euophrys C.

L. Koch, 1834 (Araneae: Salticidae) — one, two or three gen-

era? In: Selden PA (ed.) Proceedings of the 17th European

Colloquium of Arachnology, Edinburgh 1997. Edinburgh,

pp. 115-120

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BHL

Blank Page Digitally Inserted

Bonn zoological Bulletin 68 (2): 189-203

2019 -Kanturski M. & Bezdék A.

https://do1.org/10.20363/BZB-2019.68.2.189

ISSN 2190-7307

http://www.zoologicalbulletin.de

Research article

urn:|sid:zoobank.org:pub: 7EF7A263-F535-43F7-B6E5-94012D1CE048

The Knautia feeding species of Aphis (Insecta: Hemiptera: Aphididae)

with notes on Aphis knautiae Holman nomen nudum

Mariusz Kanturski!* & AleS Bezdék?

' Department of Zoology, Faculty of Biology and Environmental Protection, University of Silesia in Katowice,

Bankowa 9, PL-40-007 Katowice, Poland

* Biology Centre of the Czech Academy of Sciences, Institute of Entomology, Branisovska 1160/31,

CZ-370 05 Ceské Budéjovice, Czech Republic

* Corresponding author: Email: mariusz.kanturski@us.edu.pl

'urn:lsid:zoobank.org:author:78C290A3-D07B-4A F9-9358-ED8C05A702BF

2urn:Isid:zoobank.org:author:E7 A39AC2-A E8C-4CF 1-8594-795 1 FFS5A 7058

Abstract. Here we present a review of Knautia feeding aphid species of the genus Aphis Linnaeus, 1758 (Insecta: Hemip-

tera: Aphididae). A new European species Aphis holmani sp. nov. 1s described from specimens collected in the Czech Re-

public, Bulgaria and Romania by the late Jaroslav Holman, who already recognized this new species, but never published

its description. It has previously been referred to as Aphis knautiae Holman nomen nudum. Morphological characters of

apterous and alate viviparous females as well as the sexual generation (oviparous females and males) are described and

figured. The new species is associated with Knautia drymeia and its affinities with other related species living on species

of Knautia are presented. Morphological characteristics of known morphs of other Knautia feeding species A. confusa

Walker, 1849, A. Jongini Huculak, 1968 and A. thomasi (Borner, 1950) are given and figured. Additionally hitherto un-

known sexual forms of A. thomasi are described from specimens collected in the Czech Republic and Poland. A key to

known aphid species living on species of Knautia is also provided.

Key words. Aphidini, Aphidoidea, description, Europe, taxonomy, Aphis holmani sp. nov.

INTRODUCTION

Aphis Linnaeus, 1758 with 589 valid species is the most

speciose genus within the Aphidoidea as a whole (Favret

2019; Blackman & Eastop 2019). Species belonging to

this genus are known mostly from the northern hemi-

sphere with small number of taxa which are native for the

southern part of the globe. Most of species form groups

of similar or very similar species whose elucidation may

be difficult, but mostly all they are characterized by ter-

minal process which is no longer that the fourfold of the

basal part length, relatively short siphunculi and a subtri-

angular or tongue-shaped cauda. Viviparous and sexual

generations of Aphis are furthermore characterized by un-

developed or low antennal tubercles with a little convex

frons, marginal tubercles on prothorax, abdominal seg-

ments I and VII and occasionally on other segments of

the abdomen, which although lack in some species (Heie

1986).

The plant genus Knautia L. (Dipsacaceae) comprises

about 50—55 species distributed in western Eurasia and

northwestern Africa, with the highest species diversity is

in southern and southeastern Europe, especially the Alps

and the Balkan Peninsula. Species from this genus can be

found dry grasslands, wet meadows, alpine grasslands,

Received: 19.08.2019

Accepted: 04.11.2019

forests, and ruderal communities (Ehrendorfer 1976;

ReSetnik et al. 2014). So far 11 aphids species have been

described or recorded from ten Knautia species of which

three species belong to genus Aphis: A. confusa Walker,

1849, A. longini Huculak, 1968 and A. thomasi (Borner,

1950) are associated with Knautia arvensis (L.) Coult.,

K. dinarica (Murb.) Borbas, K. dipsacifolia Kreutzer and

K. integrifolia Ehrend. Only A. confusa feeds on all four

species (both, A. Jongini and A. thomasi are known only

from K. arvensis) (Holman 2009; Blackman & Eastop

2019).

Knautia drymeia Heuff. has a native distribution range

from South-eastern Germany to Northern Greece (Alba-

nia, Austria, Bulgaria, Czech Republic, Croatia, Greece,

Hungary, Montenegro, Romania, Serbia and Slovakia)

(ReSetnik et al. 2016). It is a perennial herb, usually with

a monopodial stalk bearing a terminal leaf rosette and

lateral flowering stems. Its habitats are forests and forest

margins, especially deciduous and hard-wood floodplain

forests, but it also reaches into subalpine habitats (Ehren-

dorfer 1962; Fischer et al. 2008). Only four aphid species

from the tribe Macrosiphini have been reported from this

species to date: Aulacorthum knautiae Heie, 1960, Mac-

rosiphum knautiae Holman, 1972, M. rosae (Linnaeus,

Corresponding editor: R. Peters

Published: 11.12.2019

190 Mariusz Kanturski & Ales Bezdék

1758) and Ovatomyzus boraginacearum Eastop, 1952

(Holman 2009).

Another nominal species associated with K. drymeia

is Aphis knautiae Holman which was for the first time

mentioned in 1981 in the list of aphids of Romania (Hol-

man & Pintera 1981) but recognized as a nomen nudum

(ICZN 1999; Remaudiere & Remaudiere 1997; Favret

2019). During the work in the Aphidoidea collection of

the late Jaroslav Holman (now deposited in the Biology

Centre of the Czech Academy of Sciences, Institute of

Entomology, Ceské Budéjovice, Czech Republic) speci-

mens collected in the Czech Republic, Bulgaria and Ro-

mania from K. drymeia and labeled as “Aphis knautiae

Holman” have been found and recognized as A. knautiae

Holman nomen nudum. A careful examination and com-

parison with other Aphis species associated with Knau-

tia revealed that the specimens represent an undescribed

species; its diagnosis and detailed description is given in

this paper which makes this name available.

MATERIAL AND METHODS

The specimens were examined using light microscope

Leica DM 3000 led with Leica MC 190 HD camera and

Nikon Eclipse E600 with Nikon DS—Fi camera. The mea-

surements were done according to Ilharco & van Harten

(1987) and Blackman & Eastop (2006). Measurements

are given in millimeters (Supplementary tables 1-4).

Abbreviations

ANT = antennae or their lengths

ANT I-VI = antennal segments J, II, HI, IV, V,

VI or their lengths (ratios between

antennal segments are simply

given as e.g. “VI: III’)

BASE = basal part of last antennal

segment or its length

BD II = basal articular diameter of ANT III

BL = body length (from anterior border

of the head to the end of cauda)

FEMORA III = hind femora length

FEMORA HI LS = longest setae on hind femora

GP = genital plate

PHT = posterior seta (hair) of hind

trochanter

HW = greatest head width across com-

pound eyes

HT II = second segment of hind tarsus or

its length

LS ANT III = length of longest setae of ANT III

MAX W = maximal width of abdomen

PT = processus terminalis of last anten-

nal segment or its length

MTu = marginal tubercles

Bonn zoological Bulletin 68 (2): 189-203

SIPH L = siphunculi length

SIPH W = maximum width of siphunculus

TIBIAE III = hind tibiae length

URS = ultimate segments of rostrum

(IV + V) or their length

apt. = apterous viviparous female

al. = alate viviparous female

Q = oviparous female

J = male

Depositories of the material examined

DZUS = Hemiptera Collection of the Department of

Zoology, University of Silesia in Katowice

Poland

IECA = Biology Centre of the Czech Academy of

Sciences, Institute of Entomology, Ceské

Budéjovice, Czech Republic;

ZMPA = Zoological Institute, Polish Academy of

Sciences, Warsaw, Poland

RESULTS

Shared characters of Aphis species feeding on Knautia

spp.

Body egg or pear-shaped. Head slightly sclerotized with

very small and slightly visible ANT tubercles and sub-

convex frons. Head and body sclerotization often with

microsculpture. ANT 5 or 6-segmented. ANT HI without

secondary rhinaria, ANT V with small ciliated rhinarium.

ANT VI with small ciliated major rhinarium, 5—6 visi-

ble additional rhinaria and 4 apical setae. Ultimate ros-

tral segment with 2 (sometimes 4) accessory setae. First

segments of tarsi with 3—3—2 ventral setae. Dorsum of

thorax membranous, sometimes with more or less visible

sclerotic plates on marginal areas. Dorsum of thorax and

abdomen covered by a few short, rigid setae with blunt

apices. Abdomen membranous. ABD I and VII always

with MTu. Siphunculi short, cylindrical, slightly taper-

ing. Cauda short, tongue-shaped. Alate viviparous fe-

males with small number of secondary rhinaria on ANT

If] and IV. Abdomen with marginal sclerotic plates on

ABD II-IV. ABD VI with wide postsiphuncular sclerites.

Oviparous females with more or less swollen hind tibiae

with rounded or slightly oval pseudosensoria on almost

whole length and genital plate divided into two separated

sclerotic parts. Males apterous with small number of sec-

ondary rhinaria on ANT ITI—V. They are small, rounded

with sclerotic rings. Male genitalia with wide and lobate

parameres with a large number of long and pointed setae.

Basal part of the phallus short and hooked-shaped.

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A new Aphis species from Europe 19]

Morphological characteristics of species

Aphis (Aphis) confusa Walker, 1849

Walker, 1849: XLV

Figs 1-7; Supplementary tables 1-4; Tables 1—2

Apterous viviparous female (n=27).

Colour in life: apterae are pale yellow, yellowish

green, green or dark green, depending on location on host

(Blackman & Eastop 2019); pigmentation on slide: head

sclerotized, brown. Antennae light brown with lighter

ANT and basal part of ANT IV or ANT HI, and basal part

of ANT IV and V. Legs yellow to pale brown with brown

distal parts of tibiae and tarsi (sometimes only slightly

visible). Abdomen yellow with brown to dark brown

SIPH and light brown cauda and anal plate (Fig. 1a).

0.50 mm

HW 0.36—-0.40 x ANT. Head setae 0.015—0.025 mm

long, 0.75—0.90 x BD III. ANT 0.51-0.58 x BL. ANT IV

slightly shorter, as long as or slightly longer than ANT V.

ANT VI with PT 2.45—3.00 x BASE. Other antennal ra-

tios: VI:I 1.28-1.52, V-III 0.56—0.69, IV:II 0.56-0.76,

PT:HI 0.87-0.91, PT:IV 1.42-1.71, PT:V_ 1.55—1.80.

ANT bearing very short and blunt setae (Fig. 5a). ANT

III setae shorter than the width of the segment, 0.010—

0.015 mm long, LS III 0.50-0.75 x BD III. ANT I with

4—5, ANT II with 3-5, ANT III with 4-9, ANT IV with

2-4, ANT V with 2-5, ANT VI with 2-3 basal setae.

Rostrum reaching hind coxae. URS 0.37-0.50 x ANT

I, 0.27-0.34 x ANT VI, 0.37—0.46 x PT, 1.09-1.37 x

BASE and 1.00—1.22 x HT II. Mesosternal furca wide,

fused basally. HI FEMORA bearing long, slightly rigid

setae with blunt or slightly expanded apices (Fig. 5b),

0.015—0.032 mm long, IIT FEMORA LS 0.63—0.76 x

0.50 mm

Fig. 1. Apterous viviparous females of Knautia feeding Aphis species. (a) A. confusa; (b) A. holmani sp. nov.; (¢) A. longini;

(d) A. thomasi.

Bonn zoological Bulletin 68 (2): 189-203

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192 Mariusz Kanturski & Ales Bezdék

Fig. 2. Known alate viviparous females of Knautia feed-

ing Aphis species. (a) A. confusa;, (b) A. holmani sp. nov.;

trochantero-femoral suture length. Posterior seta on

hind trochanter 0.76—1.00 x trochantero-femoral suture

length (Fig. 5c). III TIBIAE bearing long and rigid setae

with slightly blunt apices, 0.017—0.030 mm long. HT II

0.37-0.44 x ANT III, 0.27-0.30 x ANT VI, 0.37-0.42 x

PT and 1.00—1.12 x BASE. Abdomen with very small

MTu on ABD IV. SIPH 1.60-1.88 x cauda, 0.13-0.18 x

BL, and 1.09-1.30 x ANT HI. Setae on ABD I-V 0.015—

0.020 mm long, about 0.85 x BD III. Setae on ABD VI-—

VIII 0.017—0.025 mm long, 0.85—1.13 x BD II. Genital

plate anterior setae 0.035—0.042 mm long, 1.75—2.00 x

BD III. Cauda with 5-8 setae (Fig. 6a, b).

Alate viviparous female (n=A4).

Colour in life: unknown; pigmentation on slide: head

and thorax sclerotized, brown to dark brown. Antennae

brown to light brown with paler basal half of ANT III,

IV and V. Wings light brown with darker venation. Fore

Bonn zoological Bulletin 68 (2): 189-203

0.50 mm

Fig. 3. Known oviparous females of Knautia feeding Aphis

species. (a) A. confusa; (b) A. holmani sp. nov.; (¢) A. thomasi.

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A new Aphis species from Europe 193

and middle femora uniformly yellow to light brown, hind

femora yellow to light brown with darker distal half. Tib-

iae yellow to light brown with brown distal ends and tar-

si. Abdomen pale with brown sclerite, plates, SIPH, cau-

da and anal plate (Fig. 2a). HW 0.34—0.36 x ANT. Head

setae 0.012-0.015 mm long, 0.76—-0.88 x BD III. ANT

0.62—0.67 x BL. ANT II with 4—5 secondary rhinaria,

ANT IV slightly shorter or slightly longer than ANT V.

ANT VI with PT 2.28—2.90 x BASE. Other antennal ra-

tios: VIII 1.32—1.70, V:II 0.57—0.77, IV:II 0.62-0.69,

PT: 0.92-1.20, PT:IV 1.33-1.93, PT:V_ 1.55—1.70.

ANT bearing very short and blunt setae. ANT III setae

shorter than the width of the segment, 0.010—-0.012 mm

long, LS II 0.58—-0.83 x BD I. ANT I with 4-5, ANT II

with 4-5, ANT III with 5—7, ANT IV with 4-6, ANT V

with 4-6, ANT VI with 2-3 basal setae. Rostrum reach-

ing mesosternum. URS 0.42-0.45 x ANT III, 0.26—0.31

x ANT VI, 0.37-0.45 x PT, 0.90-1.10 x BASE and 1.10—

1.17 x HT II. It] FEMORA bearing medium sized to long,

slightly rigid setae with blunt apices, 0.017—0.027 mm

long, III FEMORA LS 0.59-0.67 = trochantero-femoral

suture length. Posterior seta on hind trochanter 0.40—0.81

x trochantero-femoral suture length. I] TIBIAE bearing

long and rigid setae with blunt apices, 0.017—0.027 mm

long. HT II 0.38-0.39 x ANT III, 0.22—0.28 x ANT VI,

0.32—0.41 x PT and 0.77-0.95 x BASE. Abdomen with

solid sclerotic bar on ABD I, small spinal and spinopleu-

ral sclerites on ABD ABD II-V. ABD V without presi-

phuncular sclerites. ABD VI with large spinal plate. ABD

VII with small spinal plate (Fig. 7a). SIPH 1.50—1.61 x

cauda, about 0.13 <x BL, and 0.87—0.95 x ANT III. Setae

on ABD I-V 0.010—0.015 mm long, 0.66—0.88 x BD III.

Setae on ABD VI-VIII 0.015—0.020 mm long, 0.88-—1.17

x BD III. Genital plate anterior setae 0.020—0.040 mm

long, 1.47—2.35 x BD HI. Cauda with 5-7 setae.

Oviparous female (n=A4).

Colour in life: unknown; pigmentation on slide: head

slightly sclerotized, yellow to light brown. Antennae yel-

low or pale with ANT I, ANT II, apical part of ANT V,

BASE and PT light brown. Legs yellow with light brown

distal parts of tibiae and tarsi. Hind tibiae uniformly light

brown. Abdomen yellow with brown SIPH, cauda and

anal plate (Fig. 3a). HW 0.47-0.50 x ANT. Head setae

0.010-0.012 mm long, 0.58—0.62 x BD III. ANT about

0.48 x BL. ANT IV slightly shorter than ANT V. ANT

VI with PT about 2.50 x BASE. Other antennal ratios:

VII = 2.00-2.33, V:IIl 0.85-0.91, IV:III 0.64—-0.83,

PT: 1.42-1.66, PT:IV 2.20-2.22, PT:V 1.66—1.81.

ANT bearing very short and blunt setae. ANT III setae

shorter than the width of the segment, 0.007-0.010 mm

long, LS III 0.58—0.62 x BD III. ANT I with 5, ANT II

with 3, ANT II with 5-6, ANT IV with 2, ANT V with

3-4, ANT VI with 2-3 basal setae. Rostrum reaching

hind coxae. URS 0.67-0.83 x ANT III, 0.33—0.35 x ANT

VI, 0.47-0.50 x PT, 1.18-1.25 x BASE and 1.18-1.33

Bonn zoological Bulletin 68 (2): 189-203

0.50 mm

Fig. 4. Known males of Knautia feeding Aphis species. (a)

A. confusa; (b) A. holmani sp. nov.; (¢) A. thomasi.

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194 Mariusz Kanturski & Ales Bezdék

Table 1. Morphological differences between apterous and alate viviparous females of Aphis holmani sp. nov. and A. confusa.

Apterous viviparous females

eer Aphis holmani sp. nov. Aphis confusa

Femora setae pointed with blunt apices

Head setae pointed with blunt apices

URS 0.13-0.16 0.10-0.12

I FEMUR LS 0.045—0.050 0.030-0.045

Head LS 0.030-0.035 0.015-0.025

Frontal setae 0.030-0.035 0.017—0.025

ABD VIII setae 0.025—0.030 0.017-0.025

GP anterior setae L 0.045—0.050 0.035-0.042

PT/BASE 1.75—2.20 2.45-3.00

URS/HT II 1.40-1.52 1.00-1.22

URS/ANT VI 0.40—0.43 0.27-0.34

URS/PT 0.59-0.66 0.37-0.45

HT IV/ANT II 0.26-0.35 0.37-0.44

HT I/BASE 0.83—0.90 1.00-1.25

SIPH/CAUDA 1.05—-1.50 1.60-1.88

SIPH/ANT III 0.59-0.91 1.09-1.30

IIT FEMUR LS/TFS 0.90-1.11 0.63—-0.76

GP anterior seta/ BD III 2.25—2.94 1.75—2.00

ABD VIII setae/BD II 1.11-1.76 0.85—0.90

Alate viviparous females

Aphis holmani sp. nov. Aphis confusa

URS 0.125—0.135 0.100—-0.110

Il FEMUR LS 0.032-0.035 0.022-—0.025

ABD I-V setae 0.020—0.025 0.010-0.015

ABD VI-VIII setae 0.025—0.040 0.015-0.020

HW/ANT 0.32-0.33 0.34—0.36

URS/HT II 1.50-1.56 1.10-1.17

URS/ANT II 0.51-0.59 0.42-0.45

URS/ANT VI 0.36-0.37 0.26-0.31

URS/PT 0.51-0.52 0.37-0.45

URS/BASE 1.25-1.28 0.90-1.10

SIPH/CAUDA 1.14-1.36 1.50-1.61

SIPH/BL 0.090-0.010 0.13

SIPH/ANT III 057-0.58 0.87-0.95

I FEMU LS/TFS 0.80—0.86 0.59-0.67

Head setae/BD III 0.11-0.14 0.08-0.09

Frontal setae/BD III 0.13-0.16 0.07-0.10

ABD FLV setae/BD III 0.13 0.006—0.008

ABD VI-VIII setae/BD III 0.16 0.08-0.11

x HT II. Mesosternal furca separated or slightly fused,

wide. Posterior seta on hind trochanter 0.64—1.00 x tro-

chantero-femoral suture length. II] FEMORA with 60-89

pseudosensoria, bearing medium sized to long, slightly

rigid setae with slightly blunt apices which are mostly

pointed on the dorsal side, 0.010-0.075 mm long, III

FEMORA LS 0.59-0.77 x trochantero-femoral suture

length. III TIBIAE with 60-89 mostly circular, different

in size pseudosensoria and with long and rigid setae with

blunt apices, 0.010—0.030 mm long. HT I 0.57-0.62 x

ANT IL, 0.26—-0.28 x ANT VI, 0.37—0.40 x PT and 0.93—

Bonn zoological Bulletin 68 (2): 189-203

1.00 x BASE. Abdomen with small marginal tubercles

on ABD I, V and VU. SIPH 1.23—1.28 x cauda, about

0.11 x BL, and 1.28—-1.33 x ANT III. Setae on ABD I-V

0.010—0.015 mm long, about 0.62—0.88 x BD III. Setae

on ABD VI-VIII 0.015—0.025 mm long, 0.93—1.00 x BD

II. Genital plate anterior setae 0.026—0.035 mm long,

1.52—2.18 x BD III. Cauda with 6-7 setae.

Male (n=5).

Colour in life: unknown; pigmentation on slide: head

sclerotized, light brown. Antennae light brown with

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A new Aphis species from Europe 195

Table 2. Morphological differences between sexual morphs of Aphis holmani sp. nov. and A. confusa.

Oviparous females

Character Aphis holmani sp.nov. Aphis confusa

Tibiae setae pointed with blunt apices

Pseudosensoria 25-42 60-89

ABD VIII setae 8-10 3-5

ANT 0.77-0.85 0.70-0.73

II FEMUR LS 0.04—0.05 0.025—0.035

TIBIAE LS 0.035—0.045 0.01—0.03

Head setae 0.017—0.040 0.010—0.012

Frontal setae 0.030—0.040 0.015—0.025

ABD F[-V setae 0.032—0.075 0.010—0.015

ABD VIII setae 0.025—0.030 0.015—0.025

ANT/BL 0.51-0.56 0.48

HW/ANT 0.44—0.46 0.47—0.50

PT/BASE 2.68—3.06 2.50

URS/BASE 1.37-1.43 1.18-1.25

SIPH/CAUDA 1.07-1.12 1.23-1.28

SIPH/BL 0.08—-0.09 0.11

SIPH/ANT II 0.87-0.96 1.28-1.33

II FEMUR LS/TFS 0.84—1.00 0.57-0.77

GP anterior seta/ BD III 2.50-3.23 1.52-2.18

Head setae/BD II 0.85—-1.47 0.58-0.62

ABD [-V setae/BD III 1.10-1.29 0.62-0.88

ABD VI-VIII setae/BD II 1.88—2.00 0.93-1.00

Males

Aphis holmani sp. nov. Aphis confusa

ANT IV rhinaria 12-18 2-12

ANT V rhinaria 11-13 4-10

TFS 0.032—0.037 0.040—0.045

ABD [-V setae 0.022—0.027 0.010—0.013

ABD VI-VIII setae 0.030—0.052 0.015—0.030

ANT/BL 0.87—1.06 0.76—0.80

HW/ANT 0.31-0.36 0.39-0.40

PT/BASE 2.94—3.28 2.64—2.73

ANT V/ANT III 0.82-0.83 0.76—0.80

ANT IV/ANT HI 0.82-0.88 0.61-0.71

ANT IV/ANT V 1.00—1.06 0.80—0.89

URS/HT II 1.33-1.35 1.21-1.30

URS/ANT II 0.55 0.48—0.50

HT I/BASE 0.88—1.00 0.82-0.86

SIPH/CAUDA 0.88—-0.90 1.00—1.20

SIPH/ANT II 0.47-0.50 0.57—0.64

Il] FEMUR LS/TFS 1.06—-1.08 0.78—0.80

Head setae/BD III 1.11-1.33 0.50—0.76

Frontal setae/BD III 1.33 0.60—0.96

ABD L[-V setae/BD III 1.46—1.80 0.50—0.076

ABD VI-VIII setae/BD III 2.00—2.46 0.75-1.15

slightly paler basal part of ANT II. Legs with yellow or

light brown femora and pale to yellow tibiae with brown

distal parts and tarsi. Abdomen pale with light brown

SIPH, cauda and genitalia (Fig. 4a). HW 0.39-0.40 x

Bonn zoological Bulletin 68 (2): 189-203

ANT. Head setae 0.010—0.017 mm long, 0.50-0.76 x

BD III. ANT 0.76-0.80 x BL. ANT III with 0 (on one

segment)-14, ANT IV shorter, than ANT V with 2-12,

ANT V with 1-11 secondary rhinaria. ANT VI with PT

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196 Mariusz Kanturski & AleS Bezdék

0.01 mm

0.025 mm

Fig. 5. Key morphological differences between apterous viviparous females of Knautia feeding Aphis species. A. confusa. (a) ANT

III seta with blunt apex; (b) hind femora inner setae with blunt apices; (c) long hind trochanter seta; A. holmani sp. nov.: (d) ANT

III seta with pointed apex; (e) hind femora inner setae with pointed apices; (f) very long hind trochanter seta; A. Jongini: (g) ANT

III long and pointed setae; (h) hind femora inner setae with pointed apices; (i) long hind trochanter seta; A. thomasi: (j) ANT II and

IV very short setae with blunt apices; (k) very short hind trochanter inner setae with blunt apices; (1) very short hind trochanter seta.

2.64—2.73 <x BASE. Other antennal ratios: VI:III 1.64—

1.77, V:III 0.76—-0.80, IV:II 0.61-0.71, PT:IIl 1.20—

1.28, PT:IV 1.80-1.95, PT:V 1.57—1.60. ANT bearing

very short and blunt setae. ANT III setae shorter than

the width of the segment, about 0.010 mm long, LS HI

0.50-0.76 x BD III. ANT I with 4-5, ANT I with 3, ANT

II with 4-5, ANT IV with 3, ANT V with 1-3, ANT VI

with 1—2 basal setae. Rostrum reaching from hind coxae

to ABD I. URS 0.48-0.50 x ANT III, 0.27-0.30 x ANT

VI, 0.37-0.41 x PT, 1.00-1.33 x BASE and 1.21—1.30

x HT II. Mesosternal furca separated. Posterior seta on

hind trochanter 1.00—1.40 x trochantero-femoral suture

Bonn zoological Bulletin 68 (2): 189-203

length. III FEMORA bearing short to medium sized and

rigid setae with mostly blunt apices which are pointed on

the dorsal side, 0.015—0.032 mm long, IIT FEMORA LS

0.78—0.80 x trochantero-femoral suture length. III TIB-

IAE bearing short to medium sized and rigid setae with

blunt apices, 0.010-0.025 mm long. HT I 0.38—0.40 x

ANT HI, 0.22—0.23 x ANT VI, aout 0.31 <x PT and 0.82—

0.86 x BASE. Abdomen with small marginal tubercles

on ABD IV. SIPH 1.00—1.22 x cauda, 0.09-0.11 x BL,

and 0.57-0.64 x ANT III. Setae on ABD I-V 0.010-

0.013 mm long, 0.50-0.76 x BD III. Setae on ABD VI-—

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A new Aphis species from Europe 197

VIII 0.015—0.030 mm long, 0.75—1.15 x BD HI. Cauda

with 46 setae.

Material examined. CZECH REPUBLIC: Cesky Krum-

lov-Vy&né, 27 May 1987, on Knautia arvensis, J. Holman

leg., 2 al, 19705, IECA:; Karl8Stejn, 10 October 1971, on

K. arvensis, J. Holman leg., 4 2 , 5 ¢ , 13289, IECA;

FRANCE: Nevache (1600m), dépt. Hautes-Alpes,

25 June 1969, on K. arvensis, F. Leclant leg., 2 apt, L3120,

IECA; HUNGARY: Nagykovacsi: Nagyszénas, 17 June

1964, on Scabiosa sp., H. Szelegiewicz leg., 6 apt, 2225,

R344, ZMPA; on S. ochroleuca, H. Szelegiewicz leg.,

6 apt, 2782, R434, ZMPA; POLAND: Lugwald near

Olsztyn, 27 June 1964, on K. arvensis, S. Huculak leg.,

3 apt (per one on each slide), 189, R341, ZMPA; Kaletnik

near Suwatki, 09 July 1955, on K. arvensis, H. Szele-

giewicz leg., 4 apt, 32 R348, 1 al, 32 R439, ZMPA; SLO-

VAKIA: Vrbovce, 20 May 1967, on K. arvensis, J. Hol-

man leg., 4 apt, 10879 (apt. 33-36), IECA; SWEDEN:

Vaxjo, S. Areda, 06 June 1981, R. Danielsson leg., 2 apt,

1 al, 5132: 24, IECA.

Host plants. Dipsacus sp. D. fullonum, Knautia arvensis,

K. integrifolia, K. dinarica, K. dipsacifolia, K. longifolia,

Scabiosa sp., S. argentea, S. atropurpurea, S. canescens,

S. columbaria, S. comosa, S. ochroleuca, S. sosnowskyi,

S. pratensis (Holman 2009)

Biology. This is a monoecious and holocyclic species

with sexuales in September and October. The aphids oc-

cur On upper parts of stems and inflorescences or on un-

dersides of leaves and at base of stems or on roots (Heie

1986).

Distribution. Aphis confusa is a common and widely

distributed species in Europe (except Balkans).

Aphis (Aphis) holmani sp. nov.

Aphis knautiae Holman in Holman & Pintera, 1981: 50

(nomen nudum)

Figs 1-7, Supplementary tables 1-4; Tables 1—2

Description

Apterous viviparous female (n=63).

Colour in life: unknown; pigmentation on slide: head

slightly sclerotized, light brown. Antennae yellow or

light yellow with ANT I and ANT I and light brown,

apical part of ANT V and BASE and PT pale brown. Legs

yellow with light brown distal parts of tibiae and tarsi.

Abdomen yellow with light brown to brown SIPH, cau-

da and anal plate (Fig. 1b). HW 0.36—0.45 x ANT. Head

setae 0.015—0.035 mm long, 0.83—1.25 x BD III. ANT

0.46—0.52 x BL in 5-segmented specimens and 0.60—0.62

x BL in 6-segmented specimens. In 6-segmented speci-

Bonn zoological Bulletin 68 (2): 189-203

mens ANT IV slightly shorter or as long as ANT V. ANT

VI with PT 1.75—2.20 x BASE. Other antennal ratios:

VII 0.86-1.33, V:II 0.45-0.62, IV: 0.54—0.57 (in

6-segmented specimens), PT: III 0.55—0.91, PT:IV 1.26—

1.69, PT: V 1.16—1.57. ANT bearing very short and point-

ed setae (Fig. 5d). ANT II setae shorter than the width

of the segment, 0.010—0.125 mm long, LS III 0.50—1.00

x BD HI. ANT I with 4-6, ANT II with 3-5, ANT III

with 5-11, ANT IV with 3-5, ANT V with 2-4, ANT VI

with 1—2 basal and 4 apical setae. Rostrum reaching from

hind coxae to ABD I. URS 0.36—0.54 x ANT III, 0.40—

0.43 x ANT VI, 0.59-0.66 x PT, 1.16-1.30 x BASE and

1.40-1.52 x HT II with 2-4 accessory setae. Mesoster-

nal furca separated or slightly fused, wide. I FEMO-

RA bearing long, fine, hair-like setae with pointed apices

(Fig. 5e), 0.015—0.050 mm long, IIT FEMORA LS 0.045—

0.050 mm long, 0.90—1.11 =< trochantero-femoral suture

length. Posterior seta on hind trochanter, 1.00-1.22 x

trochantero-femoral suture length (Fig. Sf). III TIBIAE

bearing long setae with pointed apices, 0.010—0.040 mm

long. HT I with 3—3-2 ventral setae, HT II 0.26—0.35 x

ANT III, 0.26—0.30 x ANT VI, 0.38-0.47 x PT and 0.83—

0.90 x BASE. Abdomen membranous, with well-devel-

oped marginal tubercles on ABD I and ABD VII and

sometimes small marginal tubercles on ABD IV. SIPH

tubular, very slightly tapering towards apex, 1.05—1.50 x

cauda, 0.09-0.14 x BL, and 0.59-0.91 x ANT III. Dor-

sal setae very short and pointed, 0.015—0.030 mm long,

0.83—-1.25 x BD III on ABD I-V and 0.020—0.055 mm

long, 1.11—1.76 x BD III on ABD VI-VIII. Anterior setae

on genital plate 0.045—0.050 mm long, 2.25—2.94 x BD

III. Cauda with 6—9 setae (Fig. 6c, d).

Alate viviparous female (n=5).

Colour in life: unknown; pigmentation on slide: head

and thorax sclerotized, brown. ANT light brown with

brown ANT I and ANT II and slightly paler basal half of

ANT III-V. Fore and middle legs light brown with slight-

ly darker distal parts of tibiae. Hind legs with brown fem-

ora with paler proximal parts and light brown tibiae with

darker distal part of tibiae and tarsi. Wings with light

brown pterostigma and veins. Abdomen pale with light

brown sclerotization, brown SIPH and cauda (Fig. 2b).

HW about 0.32 x ANT. Head setae 0.017—0.027 mm

long, 0.11-0.14 x BD HI. ANT 0.65-0.76 x BL. ANT

IIT with 4—6 secondary rhinaria. ANT IV as long as or

slightly shorter, as long as than ANT V with 0-1 sec-

ondary rhinaria. ANT VI with PT 2.40—2.47 x BASE.

Other antennal ratios: VI:III 1.40—1.61, V:HI 0.57—0.66,

IV: 0.51-0.61, PT:I 1.00-1.14, PT:IV 1.73—1.84,

PT:V 1.71—1.73. ANT bearing very short and blunt se-

tae. ANT HI setae shorter than the width of the segment,

0.012—0.015 mm long, LS HI 0.80—1.00 x BD III. ANT I

with 4-5, ANT II with 3-5, ANT III with 6-10, ANT IV

with 3-6, ANT V with 4-6, ANT VI with 2 basal setae.

Rostrum reaching from hind coxae to ABD I. URS 0.51-—

©ZFMK

198 Mariusz Kanturski & Ales Bezdék

Fig. 6. End of abdomen (left column) and ABD VIII setae (right column) of Knautia feeding Aphis species. (a—b) A. confusa, (c— d)

A. holmani sp. nov.; (e-f) A. longini; (g-h) A. thomasi.

Bonn zoological Bulletin 68 (2): 189-203 ©ZFMK

A new Aphis species from Europe 199

0.59 x ANT III, about 0.36 x ANT VI, 0.51-0.52 x PT,

1.25—1.28 x BASE and 1.50—1.56 x HT I. WI FEMORA

bearing long, fine and pointed setae, 0.012—0.032 mm

long, III FEMORA LS 0.80-0.86 = trochantero-femoral

suture length. Posterior seta on hind trochanter 0.80—1.00

x trochantero-femoral suture length. I] TIBIAE bearing

long, fine and pointed setae, 0.012—0.032 mm long. HT

II 0.34—0.38 x ANT III, 0.23-0.26 x ANT VI, 0.33-0.34

x PT and 0.80-1.00 x BASE. Abdomen with sclerite on

ABD I, small spinal sclerites on ABD ABD H-IV. ABD

V with solid or broken presiphuncular sclerites. ABD VI

with large spinal plate. ABD VII with wide spino-pleural

plate. ABD IV and VI with small MTu (Fig. 7b). SIPH

1.14—1.636 x cauda, 0.09-0.10 x BL, and about 0.57 x

ANT UI. Setae on ABD I-V 0.020-0.025 mm long, about

1.33 x BD III. Setae on ABD VI-VIII 0.025—0.040 mm

long, about 1.66 <x BD III. Genital plate anterior setae

0.030—0.040 mm long, 2.00—2.66 x BD HI. Cauda with

8 setae.

Oviparous female (n=47).

Colour in life: unknown; pigmentation on slide: head

sclerotized, brown. Antennae pale with ANT I, ANT I,

apical part of ANT V and ANT VI brown. Legs yellow

with slightly darker very apical parts and tarsi. Abdomen

yellow with brown SIPH, cauda and anal plate (Fig. 3b).

HW 0.44-0.46 x ANT. Head setae 0.017—0.040 mm

long, 0.85—1.47 x BD III. ANT 0.51-0.56 x BL. ANT IV

shorter than ANT V. ANT VI with PT 2.68—3.06 x BASE.

Other antennal ratios: VI:III 2.00—2.18, V:HI 0.87—0.92,

IV:IIl 0.70-0.85, PT:HI 1.48-1.51, PT:IV 1.86—2.09,

PT:V 1.64—1.75. ANT bearing very short and pointed se-

tae. ANT HI setae shorter than the width of the segment,

0.011-0.015 mm long, LS III 0.60—-0.88 x BD III. ANT

I with 4-5, ANT II with 4, ANT III with 3-4, ANT IV

with 2-3, ANT V with 2-3, ANT VI with 2-3 basal setae.

Rostrum reaching hind coxae. URS 0.70-0.81 x ANT

IH, 0.35-0.37 x ANT VI, 0.35-0.37 x PT, 1.37—1.43 x

BASE and 1.22—1.43 x HT II. Mesosternal furca fused

and wide. II] FEMORA with 25-42 pseudosensoria,

bearing long, fine and pointed setae, 0.015—0.050 mm

long, III FEMORA LS 0.84—1.00 = trochantero-femoral

suture length. Posterior seta on hind trochanter 0.74—1.11

x trochantero-femoral suture length. I] TIBIAE bearing

long, fine and pointed setae, 0.017—0.045 mm long. HT

II 0.50—0.62 x ANT III, 0.24—0.29 x ANT VI, 0.32-0.39

x PT and 1.00-1.12 x BASE. SIPH 1.07—1.12 x cauda,

0.08—0.09 x BL, and 0.87—0.96 x ANT III. Setae on ABD

I-V 0.022—0.027 mm long, 1.10—1.29 x BD III. Setae on

ABD VI-VIII 0.032—0.075 mm long, 1.88—2.00 x BD

Il. Genital plate anterior setae 0.050—0.055 mm long,

2.50—3.23 x BD III. Cauda with 6-10 setae.

Male (n=16).

Colour in life: unknown: pigmentation on slide: head

slightly sclerotized, brown. Antennae light rown to

Bonn zoological Bulletin 68 (2): 189-203

brown with yellow ANT III, basal part of ANT IV and

ANT V. Legs yellow with light brown very distal parts of

tibiae and tarsi. Abdomen pale with light brown to brown

SIPH, cauda and anal plate (Fig. 4b). HW 0.31—0.36 x

ANT. Head setae 0.017—0.025 mm long, 1.11—1.33 x BD

Ill. ANT 0.87—1.06 x BL. ANT III with 10—16 secondary

rhinaria. ANT IV as long as or slightly longer than ANT

V with 12-18 secondary rhinaria. ANT V with 11-13

secondary rhinaria. ANT VI with PT 2.94—3.28 x BASE.

Other antennal ratios: VI:III 1.76—1.86, V:HI 0.82-0.83,

IV: 0.82—0.88, PT:IIl 1.35-1.38, PT:IV 1.56—1.64,

PT:V 1.64—1.66. ANT bearing very short and pointed

setae. ANT III setae shorter than the width of the seg-

ment, 0.011—0.012 mm long, LS III about 0.83 = BD III.

ANT I with 4-5, ANT II with 4, ANT IIT with 4-5, ANT

IV with 3, ANT V with 2-3, ANT VI with 2 basal setae.

Rostrum reaching from hind coxae to ABD I. URS about

0.55 x ANT III, 0.29-0.31 x ANT VI, 0.40-0.41 x PT,

1.17—1.35 x BASE and 1.33-1.35 x HT I. WI] FEMORA

bearing long, fine and pointed setae, 0.017—0.040 mm

long, III FEMORA LS 1.06—1.08 = trochantero-femoral

suture length. Posterior seta on hind trochanter 1.00—1.06

x trochantero-femoral suture length. I TIBIAE bearing

long, fine and pointed setae, 0.012—0.032 mm long. HT

II about 0.41 x ANT III, 0.22—0.23 x ANT VI, about

0.30 x PT and 0.88—1.00 x BASE. Abdomen with small

marginal tubercles on ABD VI. SIPH 0.88—0.90 x cauda,

0.08—0.10 x BL, and 0.47—0.50 x ANT III. Setae on ABD

I-V 0.022—0.027 mm long, 1.46—1.80 x BD III. Setae on

ABD VI-VIII 0.030—0.052 mm long, 2.00—2.46 x BD

IH. Cauda with 5-6 setae.

Diagnosis

From so far known Aphis species feeding on Knautia,

apterous viviparous females of A. holmani sp. nov. are

most similar to A. confusa by hind femora and hind tro-

chanter setae lengths (which are longer than 0.50 ~ tro-

chantero-femoral suture length) and genital plate anteri-

or setae lengths (which are longer than 1.00 x BD III).

Both species differ from A. thomasi in those characters

(0.20—0.50 x trochantero-femoral suture setae length and

0.20-0.50 x BD III respectively). The new species differ

from A. confusa by:

¢ Pointed setae on femora, head and antennae (setae

with blunt apices in A. confusa)

¢ longer anterior setae on the genital plate, 2.25—2.94 x

BD III (1.00—2.00 in A. confusa)

¢ higher ratio of URS/HT I, 1.40-1.52 (1.00—1.22 in

A. confusa)

¢ lower ratio of PT/BASE, 1.75—2.20 (2.45-3.00 in

A. confusa)

¢ lower ratio of SIPH/cauda, 1.05—1.50 (1.60-1.88 in

A. confusa)

©ZFMK

200 Mariusz Kanturski & Ales Bezdék

0.20 mm

Fig. 7. Key morphological differences between known alate viviparous females of Knautia feeding Aphis species. (a) abdomen

of A. confusa without presiphuncular sclerites; (b) abdomen of A. holmani sp. nov. with presiphuncular sclerites; (c) abdomen of

A. thomasi without presiphuncular and spino-pleural sclerites.

Detailed morphological differences between particular

morphs of both species are given in Tables | and 2.

Etymology. The authors have the pleasure to give the

name to honour the late Jaroslav Holman (1931-2014)

— an outstanding European aphidologist and a long-time

employee of the Biology Centre CAS in Ceské Budéjov-

ice, Czech Republic.

Biology and distribution. The new aphid species is

associated with Knautia drymeia Heuff., from which

it was collected in Central (the Czech Republic) and

South-Eastern Europe (Bulgaria and Romania). It is a

holocyclic aphid with the sexual phase in September.

Material examined. HOLOTYPE: CZECH REPUB-

LIC, Jihomoravsky kraj, Pusty Zleb, Moravsky kras, 20

June 1967, on Knautia drymeia, J. Holman leg., 1 apt,

10838 B (apt. 8), IECA. PARATYPES: CZECH REPUB-

LIC, the same data as in the holotype, 11 slides (1 apt on

each slide), 10838 B (apt. 1-7, 9-12); Macocha, 19 May

1967, on K. drymeia, J. Holman leg., 1 apt, 10877 (apt.

1), 16 apt (two on each slide), 10877 (apt. 2-17), IECA,

slide no apt.2 DZUS; Pusty zleb, 22 September 1970, on

K. drymeia, J. Holman leg., 40 9 (four on each slide),

12885 (Q 21-48, 9 61-68) 6 9, 10877 (73-78); 12 2

(four on each slide), 12885 (4 1-12), 1 9,4 0 (4 13-

16, 2 49), IECA; BULGARIA, Mt. VitoSa nr. Simeno-

vo, 22 May 1990, on K. drymeia, J. Holman leg., 2 apt,

21725 A (apt. 1-2), 4 apt, 21725 (apt. 3-6), 16 apt (four

on each slide), 21722 B (apt. 1-16), 2 al, 21722 B (al.

1-2), 3 al, 21722 B (al. 3-5), IECA; ROMANIA, Baile

Herculane jud. Caras-Sev, 20 July 1976, on K. drymeia,

J. Holman leg., 12 apt (six per each slide), 16309 (apt.

1-12), IECA.

Bonn zoological Bulletin 68 (2): 189-203

Aphis (Aphis) longini Huculak, 1968

Huculak, 1968: 333

Figs 1, 5, 6; Supplementary table 1

Apterous viviparous female (n=5).

Colour in life: matt dirty green to olive brown (Huc-

ulak, 1968); pigmentation on slide; head sclerotized,

brown. Antennae brown with yellow ANT III and basal

part of ANT IV. Femora of legs brown with lighter basal

parts, tibiae light brown to yellow with brown to dark

brown distal and apical parts and tarsi. Abdomen yellow

to light brown SIPH brown with lighter basal part, cau-

da and anal plate brown to dark brown (Fig. 1c). HW

0.34-0.35 x ANT. Head setae 0.015—0.045 mm long,

0.60-1.00 x BD HII. ANT 0.65—0.67 x BL. ANT IV as

long as or longer than ANT V. ANT VI with PT 3.58—

4.27 x BASE. Other antennal ratios: VI-III 2.07—2.11,

VIII 0.61-0.67, IV:HI 0.67—0.80, PT:III 1.65—1.67,

PT:IV 2.04—2.47, PT:V 2.47—2.68. ANT bearing long,

fine and pointed. ANT III setae as long as and longer

than the width of the segment (Fig. 5g), 0.030—0.040 mm

long, LS III 1.20—2.00 x BD HI. ANT I with 4-5, ANT

II with 5, ANT II with 1417, ANT IV with 11-12, ANT

V with 7, ANT VI with 2 basal setae. Rostrum reaching

from hind coxae to ABD I. URS 0.57-0.59 x ANT III,

0.27-0.28 x ANT VI, 0.34—0.36 x PT, 1.29-1.45 x BASE

and 1.14~-1.19 x HT II. Mesosternal furca robust, fused

and wide. II] FEMORA bearing long, fine and pointed

setae (Fig 5h), 0.025—0.055 mm long, II] FEMORA LS

1.00—1.10 x trochantero-femoral suture length. Posterior

seta on hind trochanter about 0.90 x trochantero-femoral

suture length (Fig. 51). III TIBIAE bearing long, fine and

pointed setae, 0.050-0.055 mm long. HT II about 0.50

x ANT III, 0.23-0.24 x ANT VI, 0.29-0.30 x PT and

1.08—1.27 x BASE. Abdomen with small marginal tuber-

cles on ABD IV. SIPH about 1.33 x cauda, 0.09-0.10 x

BL, and 0.76—0.78 x ANT HI. Setae on ABD I-V 0.010—

©ZFMK

A new Aphis species from Europe 201

0.015 mm long, 0.40-0.62 x BD III. Setae on ABD VI-—

VIII 0.015—0.032 mm long, 0.70-0.75 x BD II. Genital

plate anterior setae 0.060—0.070 mm long, about 3.50 x

BD III. Cauda with 16-17 setae (Fig. 6e, f).

Remarks: Despite Huculak (1968) gave short descrip-

tions of oviparous females and males in the original

description, no material has been found in the ZMPA

collection. The alate viviparous females are unknown.

Measurements of the specimens examined in this study

vary from those given by Huculak in the original de-

scription (1968) and in Blackman & Eastop (2019). The

differences are maybe due to the fact that Huculak mea-

sured more specimens which were not available for us.

Material examined. Holotype: POLAND: Trzebinia,

26 August 1967, on Knautia arvensis, L. Olesinski leg.,

1 apt, APH-1469, ZMPA; Paratype: the same data as ho-

lotype, 1 apt, APH-1479, ZMPA, 1 apt, APH-1477; other

material: Trzebinia, 01 October 1967, on K. arvensis, L.

Olesinski leg., 1 apt. 2651, R621, ZMPA;

Host plants. The species is known only from K. arvensis

(Holman 2009).

Biology. It is a monoecious and holocyclic species with

sexual generation in October. The aphids form large,

ant-attended colonies on the bases of stems on the plant

(Huculak 1968).

Distribution. Poland, Russia (Holman, 2009). Olesins-

ki & Szelegiewicz (1974) provided that this species was

also found in “Czechoslovakia” but without existing ma-

terial it was impossible to determine if it was the Czech

Republic or Slovakia.

Aphis (Aphis) thomasi (Borner, 1950)

Doralina thomasi Borner, 1950: 7

Figs 1-7; Supplementary tables 1-4

Apterous viviparous female (n=12).

Colour in life: straw yellow (Blackman & Eastop,

2019); pigmentation on slide: head sclerotized, light

brown. Antennae light brown with pale basal half of

ANT III and base of ANT IV. Tibiae of legs light brown

with slightly paler basal distal parts, tibiae light brown

to pale with light brown apical parts, tarsi light brown.

Abdomen yellow with light brown SIPH, cauda and

anal plate (Fig. 1d). HW 0.47-0.58 x ANT. Head setae

0.005—0.010 mm long, 0.25—0.29 x BD HI. ANT 0.49—

0.54 x BL. ANT IV as long as or shorter than ANT V.

ANT VI with PT 2.00-—3.20 x BASE. Other antennal ra-

tios: VIII 1.42—2.10, V:HI 0.50-0.69, IV: 0.46—0.60,

PT:III 1.00-1.60, PT:IV 1.90—2.66, PT:V_ 1.66—2.26.

ANT bearing very short and blunt setae (Fig. 5)). ANT

III setae shorter than the width of the segment, 0.003—

Bonn zoological Bulletin 68 (2): 189-203

0.007 mm long, LS HI 0.25—0.65 x BD III. ANT I with

3-6, ANT II with 3—5, ANT HI with 4-6, ANT IV with

2-3, ANT V with 1-4, ANT VI with 1-2 basal setae.

Rostrum reaching to ABD I. URS 0.65-0.80 x ANT

Il, 0.38-0.46 x ANT VI, 0.50-0.70 x PT, 1.40-1.60 x

BASE and 1.31—1.66 x HT II. Mesosternal furca slightly

fused, wide. III FEMORA bearing very short, rigid and

blunt setae (Fig. 5k), 0.005—0.007 mm long, HI FEM-

ORA LS 0.12-0.25 x trochantero-femoral suture length.

Posterior seta on hind trochanter 0.18—0.25 x trochan-

tero-femoral suture length (Fig. 51). II] TIBIAE bearing

short rigid and blunt setae, 0.005—0.025 mm long. HT II

0.43-0.60 x ANT II, 0.25-0.35 x ANT VI, 0.37-0.50

x PT and 0.92-—1.20 x BASE. SIPH 1.09-1.60 x cauda,

0.11-0.16 x BL, and 0.80—1.53 x ANT III. Setae on ABD

I-V 0.006—0.010 mm long, 0.30—0.44 x BD III. Setae on

ABD VI-VIII 0.007—0.025 mm long, 0.37—-0.73 x BD

Il. Genital plate anterior setae 0.010—0.035 mm long,

0.50-1.47 x BD II. Cauda with 6-8 setae.

Alate viviparous female (n=2).

Colour in life: unknown; pigmentation on slide: head

sclerotized, brown. Antennae yellow or light brown with

paler basal parts of ANT II-VI BASE. Femora of legs

uniformly light brown with pale distal parts. Tibiae pale

with light brown distal parts of tibiae and light brown tar-

si. Abdomen pale with light brown SIPH, cauda and anal

plate (Fig. 2c). HW 0.35—0.36 x ANT. Head setae 0.008—

0.010 mm long, about 0.66 x BD HI. ANT 0.62-0.65 x

BL. ANT UI with 8-10 secondary rhinaria. ANT IV as

long as or shorter than ANT V, with 2-4 secondary rhi-

naria. ANT V sometimes with one secondary rhinarium.

ANT VI with PT 2.30—2.53 x BASE. Other antennal ra-

tios: VI: 1.43—1.52, V:II 0.59-0.65, IV: 0.56—0.59,

PT:II 1.00-1.09, PT:IV 1.76-1.84, PT:V_ 1.53-1.84.

ANT bearing very short and blunt setae. ANT HI setae

shorter than the width of the segment, 0.005—0.010 mm

long, LS III 0.60-0.66 x BD III. ANT I with 4-5, ANT

II with 4, ANT II with 5-6, ANT IV with 3-4, ANT V

with 3—4, ANT VI with 1-3 basal setae. Rostrum reach-

ing hind coxae. URS 0.45-0.50 x ANT II, 0.31-0.32

x ANT VI, about 0.45 x PT, 1.05-1.15 x BASE and

1.22-1.23 x HT II. II] FEMORA bearing short, rigid

and blunt setae, 0.007—0.012 mm long, IIT FEMORA LS

0.31-0.35 x trochantero-femoral suture length. Posterior

seta on hind trochanter 0.31—0.39 x trochantero-femoral

suture length. HI TIBIAE bearing short rigid and blunt

setae, 0.010—0.022 mm long. HT II 0.36—0.40 x ANT III,

0.25—0.26 x ANT VI, 0.36-0.37 x PT and 0.85—0.94 x

BASE. Abdomen with sclerite on ABD I, large marginal

plates on ABD H-IV and small ones on ABD V, which

is without presiphuncular sclerites. ABD VI with large

postsiphuncular sclerites and very small spinal sclerite.

ABD VII with wide spino-pleural plate. ABD I and VII

with visible marginal tubercles (Fig. 7c). SIPH 1.09-1.61

x cauda, 0.08—0.11 x BL, and 0.54—0.73 x ANT III. Setae

©ZFMK

202 Mariusz Kanturski & AleS Bezdék

on ABD I-V 0.007—0.012 mm long, 0.50-—0.66 x BD III.

Setae on ABD VI-VIII 0.010—0.017 mm long, about 0.66

x BD UI. Genital plate anterior setae 0.012—0.015 mm

long, 0.83—1.00 x BD III. Cauda with 5-8 setae (Fig. 7c).

Oviparous female. Description (n=9).

Colour in life: unknown; pigmentation on slide: head

sclerotized, light brown. Antennae yellow or light yellow

with ANT I, ANT II, ANT V and ANT VI light brown.

Femora yellow or light brown with paler proximal parts.

Fore and middle tibiae yellow or light brown with dark-

er distal parts and tarsi. Hind tibiae light uniformly light

brown or with slightly darker apical half. Abdomen yel-

low with light brown SIPH, cauda and anal plate (Fig. 3c).

HW 0.51-0.56 x ANT. Head setae 0.005—0.011 mm long,

0.28-0.37 x BD III. ANT 0.44—0.48 x BL. ANT IV as

long as or slightly shorter than ANT V. ANT VI with PT

2.13—2.56 x BASE. Other antennal ratios: VI:II 2.35—

2.59, V:IIT 0.75—0.81, IV:II 0.60—-0.81, PT:III 1.60—1.86,

PT:IV 2.27-2.66, PT:V 2.00—2.27. ANT bearing very

short and blunt setae. ANT III setae shorter than the width

of the segment, 0.005—0.075 mm long, LS II 0.28-0.58

x BD III. ANT I with 4-6, ANT IT with 3-5, ANT HI

with 3-5, ANT IV with 3, ANT V with 3-4, ANT VI

with 2-3 basal setae. Rostrum reaching hind coxae. URS

0.91-1.10 x ANT III, 0.36—-0.46 x ANT VI, 0.51-0.68 x

PT, 1.29-1.46 x BASE and 1.31—1.46 x HT II. Mesoster-

nal poorly visible almost separated. IIT FEMORA bear-

ing very short, rigid and blunt setae, 0.005—0.010 mm

long, III FEMORA LS 0.15—0.23 x trochantero-femoral

suture length. Posterior seta on hind trochanter about

0.17 x trochantero-femoral suture length. III TIBIAE

with 27-53 pseudosensoria, bearing short to medium in

length, rigid and blunt setae, 0.007—0.022 mm long. HT

II 0.66—0.75 x ANT III, 0.28—0.31 <x ANT VI, 0.39-0.46

x PT and 0.94-1.00 x BASE. SIPH 1.13-1.33 x cauda,

0.10—0.11 x BL, and 1.25—1.50 x ANT III. Setae on ABD

I-V 0.005-0.008 mm long, about 0.25—-0.42 x BD II.

Setae on ABD VI-VIII 0.010-0.017 mm long, 0.50—0.80

x BD III. Genital plate anterior setae 0.017—0.020 mm

long, 0.87—1.14 x BD III. Cauda with 6-10 setae.

Male. Description (n=1).

Colour in life: unknown; pigmentation on slide: head

sclerotized, light brown. Antennae light brown with paler

basal part of ANT III. Legs yellow with lighter central

parts of tibiae. Abdomen pale, SIPH light brown with

paler basal parts, cauda and anal genitalia light brown

(Fig. 4c). HW 0.49-0.52 x ANT. Head setae 0.008—

0.010 mm long, about 0.40 x BD III. ANT 5—segment-

ed, 0.57—0.60 x BL. ANT III with 10, ANT IV with 1-3

secondary rhinaria. ANT V with PT 2.28—3.00 x BASE.

Other antennal ratios: V:III 1.45—1.48, [V:HI 0.48—0.51,

PT:IIT 1.03-—1.09, PT:IV 2.00-2.25. ANT bearing very

short and blunt setae. ANT III setae shorter than the

width of the segment, 0.005—0.006 mm long, LS III

Bonn zoological Bulletin 68 (2): 189-203

0.30-0.50 x BD II. ANT I with 5, ANT II with 4, ANT

IH with 5—6, ANT IV with 3-4, ANT V with 1-2 basal

setae. Rostrum reaching ABD IIT. URS 0.60—0.64 x ANT

IH, 0.41-0.43 x ANT VI, 0.55-0.62 x PT, 1.42-1.66 x

BASE and about 1.42 x HT I. Mesosternal furca poorly

visible, separated. II] FEMORA bearing very short, rigid

and blunt setae, 0.005—0.007 mm long, IIIT FEMORA LS

about 0.16 x trochantero-femoral suture length. Posterior

seta on hind trochanter about 0.22 x trochantero-femoral

suture length. III TIBIAE bearing short, rigid and blunt

setae, 0.005—0.017 mm long. HT IH 0.42—0.45 x ANT III,

0.29-0.30 x ANT V, 0.38-0.43 x PT and 1.00-1.16 x

BASE. Abdomen with small marginal tubercles on ABD

IV. SIPH 1.00—1.10 x cauda, 0.10—0.11 x BL, and 0.64—

0.66 x ANT III. Setae on ABD I-V 0.007-0.010 mm

long, 0,37—0.50 x BD III. Setae on ABD VI-VII 0.010—

0.015 mm long, 0.50-0.75 x BD III. Cauda with 6 setae.

Material examined. BULGARIA: Liljanovo (10 km

E of Sandenski) reg. Blagoevgrad, 28 May 1990, on

Scabiosa sp., J. Holman leg., 4 apt, 21840 A (apt. 1-4),

IECA; CZECH REPUBLIC: Luka p. Mednikem, 30 Sep-

tember 1964, on S. canescens, J. Holman leg., 7 2, 8851

(2 9-15), IECA; POLAND: Augustow, 05 July 1967, on

S. ochroleuca, 8. Huculak leg., 1 apt, 1248, R838; ZMPA;

Warszawa-Bielany, 30 September 1965, on S. ochloreu-

ca, H. Szelegiewicz leg., 5 apt, 2991, R839, ZMPA; 2 apt,

29,14, 2991, R839, 363, ZMPA; UKRAINE: Catyrdag,

Krimea, 21 July 1960, on S. gramuntia (= S. trandria), J.

Holman leg., 2 al, 3898 (al. 1-2), IECA.

Host plants. Knautia arvensis, Scabiosa sp., S. argentea,

S. atropurpurea, S. canescens, S. columbaria, S. comosa

(= S. lachnophylla), S. ochroleuca, S. triandra (Holman,

2009), Pycnocomon rutifolium (Blackman & Eastop

2019).

Biology. Aphis thomasi is a monoecious and holocyclic

species with sexual phase in the end of September. The

aphids live on basal parts on the host plants.

Distribution. Bulgaria, Czech Republic, Germany, Po-

land, Romania, Slovakia, Spain, Switzerland, Ukraine,

probably Hungary (de Jong et al. 2014).

Key to Knautia feeding Aphis based on Blackman &

Eastop (2019)

1 ANT tubercles undeveloped or weakly developed.

ABD TERG 1| and 7 with marginal tubercles (MTu)

UE 5 cle cre NL RR ane aah tree EP IR, 2

— ANT tubercles well developed. ABD TERG 1| and 7

AWVACHOUL IVIERU eet eee Peele me codeine tates anes 4

2. Cauda with 14-24 hairs. Longest hairs on hind femur

as long as or longer than diameter of trochantero-

femoral suture. Well-developed MTu present on

©ZFMK

A new Aphis species from Europe 203

ABD TERG 2-4 as well as 1 and 7. ANT PT/BASE

ae SS, Feet Dis ee AS tees is IRR a Aphis longini

— Cauda with 4-9 hairs. Longest hairs on hind femur

0.15—0.7 of diameter of trochantero-femoral suture.

Small MTu present or absent from ABD TERG

2—4(-5). ANT PT/BASE 1.5-3.3 .0.0000ccccceeee 3

3. Posterior hair on hind trochanter 0.2—0.5 = diameter

of trochantero-femoral suture. Hairs on anterior half

of subgenital plate 0.2-0.5 x ANT BD IIL ................

5 EAM RR OEP hye de LoTR a Ph, oh te NSE: Aphis thomasi

— Posterior hair on hind trochanter 0.6—1.1 x diameter

of trochantero-femoral suture. Hairs on anterior half

of subgenital plate 1.0-3.0 x ANT BD HI ............. 4

4. Hairs on hind femora and antennae pointed. Hairs on

anterior half of subgenital plate 2.25—2.94 x Ant BD

IH, RIV+V 1.40-1.52 x HT ID... eee.

gent oe pe te Fea Aphis holmani sp. nov.

— Hairs on hind femora and antennae with blunt apices.

Hairs on anterior half of subgenital plate 1.00—2.00 x

Ant BD HI, RIV+V 0.90-1.10 x HT TD.

Pee Ls ee en ee, Pe nn Aphis confusa

5. Head and SIPH black. ANT HI with 7—25 rhinaria.

SIPH with polygonal reticulation on distal 0.1—0.2 of

length (8 or more rows of closed cells)................. 5

— Head and SIPH pale. ANT III with 0-3 rhinaria.

SIPH without polygonal reticulation or with only

1AATOWS?OF CLOSE: COM SF es sintatie lncctsentiese derstand hen 6

Acknowledgements. We would like to thank the editor and two

anonymous referees for valuable comments and suggestions

which greatly improved the first version of the manuscript.

Mariusz Kanturski gratefully acknowledges the Scholarship

for Outstanding Young Scientists from the Ministry of Science

and Higher Education of Poland (1165/E-340/STYP/12/17).

Ale’ Bezdék was partially supported by the research programm

Strategy AV21 of the Czech Academy of Sciences — Diversity

of Life and Health of Ecosystems”.

REFERENCES

Blackman RL, Eastop VF (2006) Aphids on the World’s Her-

baceous Plants and Shrubs. John Wiley & Sons, Ltd., West

Sussex, 1439 pp.

Blackman RL, Eastop VF (2019) Aphids on the World’s Plants.

An online identification and information guide.

http://www.aphidsonworldsplants.info (last access on June

15, 2019)

Borner C (1950) Neue europaische Blattlausarten. Naumburg

(privately published), 19 pp.

Ehrendorfer F (1962) Beitrage zur Phylogenie der Gattung

Knautia (Dipsacaceae), I. Cytologische Grundlagen und all-

gemeine Hinweise. Osterreichische Botanische Zeitschrift

109: 276-343

Ehrendorfer F (1976) Knautia L. Pp. 60-67 in: Tutin T, Hey-

wood VH, Burges NA, Valentine DH (eds) Flora Europaea,

vol. 4. Cambridge University Press, Cambridge

Favret C (2019) Aphid Species File. Version 5.0/5.0.

http://www.Aphid.SpeciesFile.org (last access on June 15,

2019)

Bonn zoological Bulletin 68 (2): 189-203

Fischer MA, Oswald K, Adler W (2008) Exkursionsflora fiir

Osterreich, Liechtenstein und Swdtirol, 3"! ed. Biologiezen-

trum der Ober6sterreichischen Landesmuseen, Linz, Austria,

1391 pp.

Heie OE (1986) The Aphidoidea (Hemiptera) of Fennoscandia

and Denmark III. Pterocommatinae and Aphidinae, Aphidin1.

Fauna Entomologica Scandinavica 17: 1-314

Holman J, Pintera A (1981) Ubersicht der Blattlause (Homop-

tera, Aphidoidea) der Rumanischen Sozialistischen Repub-

lik. Studie CSAV (Prague) 15: 1-125

Holman J (2009) Host Plant Catalog of Aphids, Palaearctic Re-

gion. Springer Science + Business Media B.V., 1216 pp

Huculak S (1968) Eine neue Art der Gattung Aphis L. (Homop-

tera, Aphididae) am Knautia arvesis (L.) Coult. aus Polen.

Annales Zoologici XX VI: 333-342

Ilharco FA, van Harten A (1987) Systematics. In: Minks AK

& Harrewijn P (eds) Aphids: their biology, natural enemies

and control. Elsevier Science Publishers, Amsterdam: 51—77

ICZN [International Commission on Zoological Nomenclature]

(1999) International Code of Zoological Nomenclature. 4th

Edition. The International Trust for Zoological Nomencla-

ture, London, 306 pp

Jong Y de, Verbeek M, Michelsen V, Bjorn P, Los W, Steeman

F, Bailly N, Basire C, Chylarecki P, Stloukal E, Hagedorn

G, Wetzel F, Glockler F, Kroupa A, Korb G, Hoffmann A,

Hauser C, Kohlbecker A, Miller A, Gtintsch A, Stoev P,

Penev L (2014) Fauna Europaea — all European animal

species on the web. Biodiversity Data Journal 2: e4034.

https://doi.org/10.3897/BDJ.2.e4034

Olesinski L, Szelegiewicz H (1974) Mszyce (Homoptera, Aphi-

dodea) okolic Chrzanowa. Fragmenta Faunistica XIX (12):

319-347

ReSetnik I, Frajman B, Bogdanovic S, Ehrendorfer F,

Schonswetter P (2014) Disentangling relationships among

the diploid members of the intricate genus Knautia (Capri-

foliaceae, Dipsacoideae). Molecular Phylogenetics and Evo-

lution 74: 97-110

ReSetnik I, Frajman B, Schoénswetter P (2016) Heteroploid

Knautia drymeia includes K. gussonei and cannot be separat-

ed into diagnosable subspecies. American Journal of Botany

103(7): 1300-1313

Walker F (1849) Descriptions of new British Aphides. The Zo-

ologist 7 (appendix): xliit-lvil

APPENDIX I

(electronic supplement, available at www.bonnzoologicalbulletin.de)

Suppl. table 1. Measurements (in mm) of apterous viviparous

females of Knautia feeding Aphis species; “measurements of

ANT III for specimens with 5-segmented antennae. ““measure-

ments of ANT III for specimens with 6-segmented antennae.

Suppl. table 2. Measurements (in mm) of known alate vivipa-

rous females of Knautia feeding Aphis species.

Suppl. table 3. Measurements (in mm) of available for exam-

ination oviparous females of Knautia feeding Aphis species.

Suppl. table 4. Measurements (in mm) of available for ex-

amination males of Knautia feeding Aphis species (male of A.

thomasi with 5-segmented antennae).

©ZFMK

BHL

Blank Page Digitally Inserted

APPENDIX 1

Suppl. table 1. Measurements (in mm) of apterous viviparous females of Knautia feeding Aphis species.

(APPENDIX 1)

A new Aphis species from Europe

“measurements of ANT III for specimens with 5-segmented antennae; “measurements of ANT III for specimens with 6-segmented

antennae.

Character Aphis confusa Aphis holmani sp. nov. Aphis longini Aphis thomasi

BL 1.75—2.07 1.50—2.00 2.00—2.10 1.20-1.55

MAX W 0.87—1.32 0.85—1.25 1.25—1.32 0.71-0.92

HW 0.36-0.45 0.38-0.44 0.48-0.49 0.35-0.38

ANT 0.94—-1.21 0.85—1.20 1.35-1.41 0.59-0.78

0.29-0.38* 0.13—-0.15*

ANT II 0.22-0.32 0.17

0.24—0.33** 0.13—-0.19**

ANT IV 0.14-0.19 0.13-0.19 0.19-0.21 0.06-0.10

ANT V 0.14—0.18 0.14-0.19 0.16-0.19 0.09-0.11

ANT VI 0.32-0.39 0.30-0.37 0.55—0.58 0.22-0.31

BASE 0.08-0.11 0.10-0.13 0.11-0.12 0.070—0.080

PT 0.240 .28 0.20-0.24 0.43-0.47 0.15—-0.24

URS 0.10-0.12 0.13-0.16 0.15—-0.16 0.105—0.125

Hi FEMUR 0.37—-0.50 0.35—-0.49 0.48-0.50 0.23—-0.34

I FEMUR LS 0.030-0.035 0.045-0.050 0.050-0.055 0.010-0.011

PHT 0.035-0.050 0.050-0.055 0.050 0.010-0.012

HI TIBIA 0.69-0.93 0.64—-0.94 0.92-0.95 0.44—0.60

HT Il 0.09-0.12 0.085-0.100 0.13-0.14 0.075—0.095

SIPH 0.24—0.38 0.18-0.27 0.20-0.22 0.20-0.24

CAUDA 0.15-0.21 0.15-0.18 0.18-0.18 0.13-0.15

GPL 0.10-0.13 0.11-0.12 0.14-0.16 0.090-0.100

GP W 0.20-0.26 0.22-0.43 0.27-0.29 0.19-0.20

Bonn zoological Bulletin 68 (2): 189-203

©ZFMK

a Mariusz Kanturski & AleS Bezdék

Suppl. table 2. Measurements (in mm) of known alate viviparous females of Knautia feeding Aphis species.

Character Aphis confusa Aphis holmani sp. nov. Aphis thomasi

BL 1.55—1.70 1.20—-1.57 1.42-1.50

MAX W 0.72—0.80 0.50—-0.66 0.50—0.60

HW 0.36—-0.37 0.30—0.34 0.33-0.35

ANT 1.015-1.055 0.920-1.035 0.92-0.94

ANT I 0.22—0.26 0.21-0.26 0.22-0.23

ANT IV 0.15—0.18 0.13-0.15 0.13

ANT V 0.15—0.17 0.14-0.15 0.13-0.15

ANT VI 0.34—0.39 0.34—0.36 0.33

BASE 0.10-0.11 0.100—0.105 0.095-0.100

PT 0.24—0.29 0.24—0.26 0.23—-0.24

URS 0.10-0.11 0.125-0.135 0.105-0.110

If FEMUR 0.38—-0.42 0.34—0.40 0.31-0.32

If FEMUR LS 0.022—0.025 0.032—0.035 0.010-0.012

PHT 0.015—0.030 0.032—0.037 0.011-0.012

I TIBIA 0.75—0.81 0.72—0.80 0.61-0.65

HT II 0.085-0.100 0.080-0.090 0.085-0.090

SIPH 0.21-0.23 0.12-0.15 0.12-0.17

CAUDA 0.13—-0.15 0.10-0.11 0.105-0.110

GPL 0.09-0.10 0.08-0.10 0.08—-0.10

GPW 0.18—0.20 0.18—-0.22 0.18—-0.20

Bonn zoological Bulletin 68 (2): 189-203

©ZFMK

A new Aphis species from Europe

(APPENDIX 1)

Suppl. table 3. Measurements (in mm) of available for examination oviparous females of Knautia feeding Aphis species.

Character

Aphis confusa

Aphis holmani sp. nov.

Aphis thomasi

1.45—1.52

0.70—0.79

1.45-1.60

0.80—-0.92

1.25-1.52

0.60—1.02

0.33—0.37

0.70—0.73

0.36—-0.38

0.77—-0.85

0.30—-0.38

0.57—-0.68

0.12—0.14

0.09-0.10

0.13—-0.16

0.11-0.13

0.10-0.12

0.06—0.09

0.11—0.12

0.12-0.14

0.29-0.32

0.08—0.09

0.23—0.28

0.20

0.08

0.21-0.24

0.075—0.085

0.16—-0.20

Hl FEMUR

0.09-0.10

0.26—0.30

0.11

0.29-0.32

0.10-0.11

0.20—-0.26

Iii FEMUR LS

PHT

0.025—0.035

0.027—0.045

0.04—0.05

0.037—0.050

0.007—0.010

II TIBIA

0.46—0.51

0.075—0.080

0.54—0.60

0.08—0.09

0.38—-0.46

0.075—0.08

0.16—-0.18

0.13—-0.14

0.13-0.14

0.12-0.13

0.12—0.16

0.11—0.13

GPW

0.10-0.11

0.23—0.25

0.10—-0.12

0.24—0.28

0.12-0.13

0.20-0.25

Bonn zoological Bulletin 68 (2): 189-203

©ZFMK

Mariusz Kanturski & AleS Bezdék

Suppl. table 4. Measurements (in mm) of available for examination males of Knautia feeding Aphis species (male of A. thomasi

with 5-segmented antennae).

Character Aphis confusa

Aphis holmani sp. nov.

Aphis thomasi

1.00-1.05

0.46—-0.57

0.80—1.07

0.43—0.55

0.97

0.31-0.33

0.76—0.84

0.27—-0.34

0.85—0.93

0.55-0.58

0.17

0.10-0.12

0.17—0.18

0.14-0.16

0.155—0.165

0.13-0.14

0.28-0.31

0.14—0.15

0.30—0.33

0.23—0.24

0.07—0.08

0.20-0.22

0.07—0.08

0.23—0.25

0.06—0.07

0.16-0.17

Hl FEMUR

0.085

0.23—0.25

0.09-0.10

0.26—0.27

0.10

I FEMUR LS

0.025—0.032

0.040-0.045

0.032—0.040

0.032—0.037

II TIBIA

0.45—0.50

0.065—0.070

0.52—0.56

0.070—0.075

0.07

HT II

SIPH

CAUDA

0.10-0.11

0.09-0.10

0.08—0.09

0.09-0.10

0.10-0.11

0.10

Bonn zoological Bulletin 68 (2): 189-203

©ZFMK

Bonn zoological Bulletin 68 (2): 205-207

2019 - Aquino D.A. et al.

https://do1.org/10.20363/BZB-2019.68.2.205

ISSN 2190-7307

http://www.zoologicalbulletin.de

Scientific note

urn:|sid:zoobank.org:pub:0D573D65-6DD1-4B3C-9ED9-D44F48638C54

First record of males of the invasive eucalyptus pest species

Leptocybe invasa Fisher & LaSalle, 2004

(Hymenoptera: Eulophidae: Tetrastichinae) from South America

Daniel Alejandro Aquino’, Carmen Marcela Hernandez” & Andrea Verénica Andorno®

‘Centro de Estudios Parasitologicos y de Vectores (CEPAVE, CONICET-UNLP), Boulevard 120 entre 60 y 64,

AR-1900 La Plata, Argentina

?3 Instituto Nacional de Tecnologia Agropecuaria (INTA). Instituto de Microbiologia y Zoologia Agricola, AR-Argentina

* Corresponding author: Email: daquino@fcnym.unlp.edu.ar

'urn:lsid:zoobank.org:author:670E065F-04CF-4247-8515-AC41D151017A

2urn:lsid:zoobank.org:author:778230C8-8EA2-4C21-892B-ACB488810D25

3urn:lsid:zoobank.org:author:47B448E3-6B06-4343-BB2F-8E87D4E17750

Abstract. The “blue gum chalcid”, Leptocybe invasa is a pest of eucalyptus worldwide. It has an Australian origin and has

expanded into Asia, Europe, Africa and America. L. invasa females were reported from South America in the last decade.

Thelytokous parthenogenesis is the most common reproductive mechanism of this pest. However, male adults have been

reported from Asia and Southeastern Europe. In this work, L. invasa males are reported for the first time from South Amer-

ica (Argentina) and information on morphological characters of males is provided. The importance of this discovery is

highlighted based on recent studies that suggested the existence of two cryptic Leptocybe species in invasive populations.

Key words. Blue gum chalcid, gall, forest, chalcidoidea.

The “blue gum chalcid”, Leptocybe invasa Fisher & La-

Salle, 2004 (Hymenoptera: Eulophidae: Tetrastichinae)

is a global pest in Eucalyptus plantations. The wasp lays

eggs in plant tissues causing the formation of galls on

the midribs, petioles and stems of young leaves. Severely

infested trees have a gnarled appearance, show stunted

growth, and in young trees, this pest can cause significant

injury (Mendel et al. 2004; Nyeko, 2005).

The rapid colonization after L. invasa spread into a

novel environment can be attributed to the absence of

natural enemies, the presence of large amounts of suit-

able host plants, the resistance to low temperatures of

adults and the occurrence of two reproductive modes

(Zheng et al. 2014b). This pest displays thelytokous re-

production but since the discovery of males, sexual re-

production has been considered a second reproductive

strategy. Nevertheless, factors influencing reproductive

strategies and the relation between reproductive strate-

gy and population expansion are unknown (Zheng et al.

2014a). In the current study, we describe male specimens

of L. invasa from Argentina and report the sex ratio in

eucalyptus plantations in the country.

In 2017 and 2018, a survey of L. invasa populations

and natural enemies was conducted in infested eucalyp-

tus plantations in an extensive productive area in Buenos

Aires and the Mesopotamia region (Entre Rios, Corri-

entes and Misiones). Almost fifty samples of 20-30 cm

Received: 13.03.2019

Accepted: 08.11.2019

long, L. invasa gall infested branches were collected. The

plant material was kept in glass containers with voile lid

and absorbent paper under controlled laboratory condi-

tions (T: 25 + 2°C, HR: 50-70% and natural lighting).

The insects that emerged from each sample were kept

in 70% ethanol and were identified by the first author.

The specimens were card- and slide-mounted following

Noyes (1990) and were compared with the descriptions

of the males in Doganlar (2005), Chen et al. (2009) and

Zheng et al. (2014b). Voucher specimens were deposited

at the Division Entomologia of Museo de La Plata. Sex

ratios were calculated as X9/X(9+). Measurements

were taken with a Leitz Wetzlar SM/LUX; specimens

were photographed using a Leica DFC295 digital cam-

era attached to a stereomicroscope Biotraza. Series of

partially focused digital images were stacked using the

Helicon Focus software (Version 6.8.0 Pro) by ©Helicon

Soft Ltd., 2000. Photoshop CC2018 was used to produce

final images with enhanced quality.

Material examined. Argentina, Entre Rios, Concordia

en 8.111.2018 and 5.x.2018 (sex ratio 3m:104f (0,972)

and 17m:1290f (0.987)); Argentina, Corrientes, Alvear

28.1x.2018 (sex ratio 1m:24f (0,960)).

According to Doganlar (2005), the male is similar to the

female described by Fisher & LaSalle (2004), except:

Corresponding editor: R. Peters

Published: 11.12.2019

206 Daniel Alejandro Aquino et al.

0,2 mm

Figs. 1-4. Leptocybe invasa male from Argentina. 1. lateral

habitus; 2. antenna; 3. genitalia; 4. forewing.

head and mesosoma brown with distinct blue to green

metallic shine (Fig. 1); ocellar triangle and mouth mar-

gin light brown; eyes and ocellus red; malar sulcus dark

brown; metasoma brown; legs almost white, except mid

and hind coxae same color of the body and hind femur

light brown. Antenna with scape and pedicel light brown

darkened dorsally and apically; funicle and clava light

brown to almost white. Wings hyaline, veins light brown

(Fig. 4). Genitalia (Fig. 3) with digitus having one claw.

The specimens collected in this study are similar to those

described by Doganlar (2005) except for the following

measurements and ratios: Head 1.28 times as broad as

high. Antenna (Fig. 2) with scape 3.13 times as long as

broad, ventral plaque 0.18—0.24x length of scape; pedicel

1.9-2.8x as long as broad, longer than anelli plus Fl; Fl

shorter than F2 (in figure 1b in Doganlar (2005) it is ob-

servable that F1 is shorter than F2 although he mentions

Bonn zoological Bulletin 68 (2): 205—207

that “F'/ is 1.46x longer than F2”), about 0.75x length of

F2, as long as broad to 0.7 times as long as broad, F2 to

F4 subequal in length, each slightly longer than broad:

clava about 2.75 times as long as broad, 1.e., slightly lon-

ger than that described by Doganlar (2005), with C1—C3

decreasing in length, except in one specimen in which Cl

is shorter than C2 and C3.

Leptocybe invasa males were reported from Turkey,

China, India, Tatwan and Thailand, and populations of

this pest around the world are known to have different

sex ratios over their geographic distribution (Nugnes

et al. 2015). Males are rare in Turkey (sex ratio: 0,992, n=

125) (Doganlar 2005) and India (sex ratio: 0,992, n=141)

(Akhtar et al. 2012). In China a low proportion of males

was reported from Fujian, Guangdong, Hainan, Guangxi,

Jiangxi and Sichuan (sex ratio > 0,95), but a high pro-

portion of males was reported from other regions (males

proportion ranging from 18—48%) and in Thailand (sex

ratio: 0,663) (Liang et al. 2010; Santongprow et al. 2011;

Zheng et al. 2018).

In this work, in only two out of twenty sampling sites

males were found (in Entre Rios and in Corrientes) and

a clearly female-biased sex ratio was found, with values

ZOrOG.

While L. invasa females were recorded from several

countries in America since 2007 (Brazil (2007), Argen-

tina (2009), Paraguay (2012), Uruguay (2013), Chile

(2014), Mexico (2014) and USA (2008) (Costa et al.

2008; Wiley and Skelley, 2008; Aquino et al. 2011; SAG

2014; Benitez et al. 2014; Vanegas et al. 2015; Jorge

et al. 2016)), this is the first report of L. invasa males

from America. Recent molecular analyses suggested that

L. invasa is, in fact, a complex of at least two cryptic

species involved in the rapid and efficient spread of the

wasp (Nugnes et al. 2015; Dittrich-Schroéder et al. 2018).

Nugnes et al. (2015) proposed that there is a link between

the presence of males in the invasive range and lineage

identity. Further molecular analyses are necessary to un-

derstand the invasion pathways and to characterize the

populations of Leptocybe species in Argentina and other

countries in America.

Acknowledgements. The authors thank Sergio Ramos, Cintia

Meneses and Edgar Eskiviski for providing samples. We grate-

fully acknowledge to CONICET and the financial support from

PICT-2016-0834 (FONCYT, MINCYT, Argentina).

REFERENCES

Akhtar MS, Patankar NV, Gaur A (2012) Observations on the

biology and male of eucalyptus gall wasp Leptocybe invasa

Fisher & LaSalle (Hymenoptera: Eulophidae). Indian Journal

of Entomology 74: 173-175

Aquino DA, Botto EN, Loiacono MS, Pathauer P (2011) Avispa

de la agalla del Eucalipto, Leptocybe invasa Fisher & LaSal-

le (Hymenoptera: Eulophidae: Tetrastichinae) en Argentina.

Revista de Investigaciones Agropecuarias 37 (2): 159-164

©ZFMK

First record of males of the invasive eucalyptus pest species Leptocybe invasa from South America 207

Benitez EA, Costa, VA, de Moraes GJ, Godziewsky D (2014)

First record of Leptocybe invasa Fisher & LaSalle (Hyme-

noptera: Eulophidae) and Rhombacus eucalypti Ghosh &

Chakrabarti (Acari: Eriophyidae) from Paraguay. Boletin

del Museo Nacional de Historia Natural de Paraguay 18:

129-132

Chen HY, Yao JM, Xu ZF (2009) First description of the male

of Leptocybe invasa Fisher & LaSalle (Hymenoptera: Eulo-

phidae) from China. Journal of Environmental Entomology

31: 285—287 (in Chinese with English abstract)

Costa VA, Filho EB, Wilcken CF, Stape JL, LaSalle J, Teixeira

L (2008) Eucaliptus gall wasp, Leptocybe invasa Fisher &

LaSalle (Hymenoptera: Eulophidae) in Brazil: New for-

est pest reaches the New World. Revista de Agricultura 83:

136-139

Dittrich-Schréder G, Hoareau TB, Hurley BP, Wingfield MJ,

Lawson S, Nahrung HF, Slippers B (2018) Population genet-

ic analyses of complex global insect invasions in managed

landscapes: a Leptocybe invasa (Hymenoptera) case study.

Biological Invasions 20: 2395

Doganlar O (2005) Occurrence of Leptocybe invasa Fisher &

LaSalle, 2004 (Hymenoptera: Chalcidoidea: Eulophidae) on

Eucalyptus camaldulensis in Turkey, with description of the

male sex. Zoology in the Middle East 35: 112-114

Jorge C, Martinez G, Gomez D, Bollazzi M (2016) First record

of the eucalypt gall-wasp Leptocybe invasa (Hymenoptera:

Eulophidae) from Uruguay. BOSQUE 37(3): 631-636

Kumari KN, Kulkarni H, Vastrad AS, Goud KB (2010) Biology

of eucalyptus gall wasp, Leptocybe invasa Fisher & LaSalle

(Hymenoptera: Eulophidae). Karnataka Journal of Agricul-

tural Sciences 23: 211-212

Liang YP, Li JY, Wen XJ, Li YX (2010) Observation on the

male ratio of Leptocybe invasa adult. Forest Pest and Disease

29: 21-22 (in Chinese with English abstract).

Mendel Z, Protasov A, Fisher N, LaSalle J (2004) Taxonomy

and biology of Leptocybe invasa gen. & sp. n. (Hymenop-

tera: Eulophidae), an invasive gall inducer on eucalyptus.

Australian Journal of Entomology 43: 101-113

Nugnes F, Gebiola M, Monti MM, Gualtieri L, Giorgini M,

Wang JG, Bernardo U (2015) Genetic diversity of the in-

vasive gall wasp Leptocybe invasa (Hymenoptera: Eu-

lophidae) and of its Rickettsia endosymbiont, and asso-

Bonn zoological Bulletin 68 (2): 205-207

ciated sex ratio differences. PLoS One 10(5): e0124660.

https://doi.org/10.1371/journal.pone.0124660

SAG (Servicio Agricola y Ganadero, CL) (2014) Leptocy-

be invasa Fischer & LaSalle (Hymenoptera: Eulophidae)

Microavispa formadora de agallas en eucalipto. Servicio

Agricola y Ganadero. https://www.sag.gob.cl/ambitos-de-ac-

cion/leptocybe-invasa-fischer-lasalle-hymenoptera-eulophi-

dae-microavispa-formadora-de

Sangtongpraow B, Charernsom K, Siripatanadilok S (2011)

Longevity, fecundity and development time of Eucalyptus

gall wasp, Leptocybe invasa Fisher and LaSalle (Hymenop-

tera: Eulophidae) in Kanchanaburi province, Thailand. Thai

Journal of Agricultural Science 44: 155-163

Tung GS, LaSalle J (2010) Pest alert — a newly discovered in-

vasion of gall forming wasps, Leptocybe invasa Fisher & La-

Salle, on eucalyptus trees in Tatwan. Formosan Entomology

30: 241-244

Vanegas-Rico JM, Lomeli-Flores JR, Rodriguez-Leyva E,

Jiménez-Quiroz E, Pujade-Villar J (2015) First record of

eucalyptus gall wasp Leptocybe invasa (Hymenoptera: Eu-

lophidae) in Mexico. Revista Mexicana de Biodiversidad

86(4): 1095-1098

Wiley J, Skelley PA (2008) A eucalyptus pest, Leptocybe in-

vasa Fisher & LaSalle (Hymenoptera: Eulophidae), genus

and species new to Florida and North America. Pest Alert.

Florida, USA: Florida Department of Agriculture and Con-

sumer Services, Division of Plant Industry. http://www.

freshfromflorida.com/content/download/68487/1614796/

Pest_Alert_- Leptocybe_invasa Blue Gum_Chalcid.pdf

Zheng X, Huang Z, Li J, Yang Z, Lu W (2018) Reproductive

Biology of Leptocybe invasa Fisher & LaSalle (Hymenop-

tera: Eulophidae). Neotropical Entomology 47(1): 19-25

Zheng XL, Li J, Yang ZD, Xian ZH,Wei JG, Lei CL, Wang

XP, Lu W (2014a) A review of invasive biology, prevalence

and management of Leptocybe invasa Fisher & LaSalle (Hy-

menoptera: Eulophidae: Tetrastichinae). African Entomology

22: 68-79

Zheng XL, Yang ZD, LiJ, Xian ZH, Yang J, Liu JY, Su S, Wang

XL,. Lu W (2014b) Rapid identification of both sexes of Lep-

tocybe invasa Fisher & LaSalle (Hymenoptera: Eulophidae:

Tetrastichinae): a morphological perspective. African Ento-

mology 22(3): 643-650

©ZFMK

BHL

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Bonn zoological Bulletin 68 (2): 209-273

2019 - Stuke J.-H. & Clements D.K.

https://do1.org/10.20363/BZB-2019.68.2.209

ISSN 2190-7307

http://www.zoologicalbulletin.de

Reasearch article

urn:lIsid:zoobank.org:pub:70A02482-0040-4136-ABB0-9C840A4D771A

New conopid records from the Afrotropical Region (Diptera) —

Part 3: Physocephalini

Jens-Hermann Stuke!* & David K. Clements?

'Roter Weg 22, D-26789 Leer, Germany

?7 Vista Rise, Llandaff, Cardiff, CF5 2SD, U.K.

* Corresponding author: Email: jstuke@zfn.uni-bremen.de

'urn:lsid:zoobank.org:author:27 ED8CBD-9201-4983-8FE3-698FB499A0E0

2urn:Isid:zoobank.org:author: 1 8B2580D-BA69-4BAC-95BE-A519F880D9B3

Abstract. The tribe Physocephalini in the Afrotropical Region is now taken to include only the genus Physocephala:

both Pseudophysocephala Krober, 1940 and Dacops Speiser, 1923 are herewith treated as junior subjective synonyms of

Physocephala Schiner, 1861. Some 45 Physocephala species are recorded in the region. Three species are new to science,

comprising P. guillarmodi spec. nov. (Burundi, Lesotho, South Africa), P. ssymanki spec. nov. (Namibia) and P. goergeni

spec. nov. (Togo). Eighteen new synonyms are introduced: P. antiqua (Wiedemann, 1830) = P. maculipes (Bigot, 1887)

syn. nov. = P. madagascariensis Krober, 1915 syn. nov. = P. gracilia Krober, 1915 syn. nov. = P. minutissima Krober, 1933

syn. nov.; P. /arvata (Speiser, 1911) =P. similis Krober 1915 syn. nov.; P. abyssinica Krober, 1915 = P. fumivena Camras,

2001 syn. nov. = P. longitheca Camras, 2001 syn. nov. = P. atronata Camras, 2001 syn. nov.; P. digitata (Speiser, 1909)

= P. simplex Krober, 1915 syn. nov. = P. ugandae Krober, 1915 syn. nov. = P. bequaertorum Camras, 1962 syn. nov. =

P. lineifrons Camras, 1962 syn. nov. = P. ethiopica Camras, 1962 syn. nov.; P. microvena Brunetti, 1925 = P. nigritarsis

(Krober, 1939) syn. nov.; P. vitripennis Curran, 1928 = P. intermedia Krober, 1936 syn. nov. = P. bouvieri (Séguy, 1936)

syn. nov. = P. meii Camras, 2001 syn. nov.; P. nigrita (Camras, 1962) = P. brevivertex (Camras, 2001) syn. nov. Phys-

ocephala nigerrima Krober, 1915 is treated as an unrecognised taxon (nomen dubium). Lectotypes are designated for

Physocephala pubescens Brunetti 1925 and P. curta Krober, 1936. Physocephala kroeberi (nom. nov.) is introduced for

Pseudophysocephala annulips Krober, 1939, the latter being a junior secondary homonym of Conops annulipes Wiede-

mann in Meigen 1824. Physocephala acroschista (Speiser, 1911) is treated as valid species (status rev.). Diagnostic keys

are presented for the Afrotropical Physocephala together with new faunistic records for 33 species.

Key words. Diptera, Conopidae, Physocephala, Pseudophysocephala, Dacops, new species, primary types, new syn-

onyms, type species, lectotype designations, nomen dubium, nomen nov.m, status review, faunistic records, identification

key.

INTRODUCTION

This is the third and final part of a work presenting new

faunistic records of the Diptera family Conopidae from

the Afrotropical Region. The first part dealt with the sub-

families Myopinae and Stylogastrinae (Stuke 2015a), and

the second part with all of the Conopinae except for the

Physocephalini (Stuke in press). The present paper deals

with the latter. As in the two previous papers, the original

aim of the project was primarily to present new faunistic

information, but this has necessitated extensive revision-

ary work in order to clarify the diagnosis of several of the

species. As a result, new keys for all Afrotropical Phys-

ocephala species have had to be prepared, comprising a

key to the nine species-groups currently recognised in the

region as well as keys to the individual species in each

group. The three publications together give a detailed

overview of all of the valid Conopidae recognised in the

Received: 09.08.2019

Accepted: 27.11.2019

region at the present time, as well as summarising the

known distribution.

MATERIALS AND METHODS

As in the previous two parts of this series, the morpho-

logical terminology used in the species descriptions 1s

mainly adopted from Cumming & Wood (2009). The

terminology used to describe the postabdomen is addi-

tionally illustrated in Figs 5, 8 and 22—23. Since Kotrba

(2000) concluded that sternite 9 is absent in the Cyclor-

rhapha, the structure referred to as ‘syntergite 8+9’ in the

previous papers is henceforward termed ‘tergite 8’ and

the structure referred to as ‘sternite 9’ in the previous pa-

pers is henceforward termed ‘sternite 8’. The structures

referred to as ‘sternite 8’ in the previous papers is not

identified yet and henceforward termed ‘ventral scleroti-

sation’ (Londsdale, personal communication). The term

Corresponding editor: X. Mengual

Published: 11.12.2019

210

‘hair’ is discarded in favour of ‘setula’ since it is often

impossible to distinguish between the two. Numbers of

setae refer to one side of the body only. Any form of ‘mi-

crotomentosity’ is referred to by the more common term

‘dusting’.

The historically important collections held by MRAC,

ZMHB and NHML were revisited and the material con-

tained therein almost completely re-identified by JHS. In

the NHML, however, there were many specimens held

under ‘Physocephala bimarginipennis’ and ‘Physoceph-

ala maculigera’ which could not all be re-identified in

detail due to time constraints.

Where new synonyms are introduced, the affected spe-

cies names are given with the original combination in

square brackets.

Faunistic records previously published elsewhere by

JHS are not repeated here, other than where these require

correction. Faunistic data is relayed from the specimen

labels with as few changes as possible. A few locations

have been altered to more commonly used names, and in

a few cases provinces or an interpretation of the location

have been added, all in square brackets. Locations which

could not be found with an internet search, or abbrevia-

tions or characters which could not be resolved, are given

in quotation marks. Coordinates are only included where

these were present on the labels.

For primary type material, the labels are rendered as

citations. The labels are listed and numbered in the order

found, commencing with the uppermost. Line-breaks are

indicated by a slash-mark [“/”] and where there are actu-

al slash-marks on the labels themselves these are includ-

ed without spaces before and after. Where text on labels

could not be deciphered with certainty this is indicated

by “[?]”. Persons mentioned on the labels of primary type

material are given in small caps. Determination labels are

also cited to assist in the interpretation of identifications

given by previous researchers.

Abbreviations used for collections referred to in the

text

AMGS = Albany Museum, Grahamstown,

South Africa, Cape Province

BMSA = National Museum Bloemfontein

South Africa

CAS = California Academy of Sciences,

San Francisco, USA

CULSP= Czech University of Life Sciences Prague,

Czech Republic

FMNH = Field Museum of Natural History, Chicago,

USA

IITA = International Institute of Tropical Agriculture,

Abomey-Calavi, Benin

ISNB- = Institut Royal des Sciences Naturelles de

Belgique, Brussels, Belgium

Bonn zoological Bulletin 68 (2): 209-273

Jens-Hermann Stuke & David K. Clements

MNHN= Muséum National d’ Histoire Naturelle,

Paris, France

MRAC = Musee Royal de I’ Afrique Centrale,

Tervuren, Belgium

MZLU = Lund University, Lund, Sweden

NHML = The Natural History Museum of London

[formerly the British Museum

(Natural History) BMNH], London, UK

NHRS = Naturhistoriska Riksmuseet,

Stockholm, Sweden

NMKE = National Museum of Kenya, Nairobi, Kenya

PASS = priv. coll. Axel Ssymank (Germany, Bonn)

PCFK = priv. coll. Christian F. Kassebeer

(Germany, Damlos)

PHJF = priv. coll. Hans-Joachim Fligel

(Germany, Kntllwald)

PJHS = priv. coll. Jens-H. Stuke (Germany, Leer)

PMHA = priv. coll. Martin Hauser (USA, Sacramento)

PMME = priv. coll. Maurizio Mei (Italy, Rome)

RMNH = Nationaal Natuurhistorische Museum

(“Naturalis”), Leiden, Netherlands

SMNS = Staatliches Museum fiir Naturkunde,

Stuttgart, Germany

SMTD = Staatliches Museum fiir Tierkunde,

Dresden, Germany

SMWN= National Museum of Namibia,

Windhoek, Namibia

TAUI = Tel Aviv University, Tel Aviv, Israel

UCDC = R.M. Bohart Museum of Entomology,

University of California, USA

USNM = Smithsonian Institution National Museum

of Natural History [formerly the United States

National Museum], Washington DC, USA

ZFMK = Zoologische Forschungmuseum Alexander

Koenig, Bonn, Germany

Museum fiir Naturkunde der

Humboldt-Universitat, Berlin, Germany

ZSM_ = Zoologische Staatssammlung,

Munchen, Gemany

Duplicates of newly designated type material are retained

in the collection of JHS for further research.

The nomenclature used in this work is based on that

given in the world checklist of Stuke (2017a), which list-

ed all synonyms, type specimen depositories and repre-

sentative faunistic records known at the time, together

with relevant sources and literature references etc. Here-

with we therefore include only new, additional or amend-

ed information on these matters where this is relevant to

the Afrotropical Region.

©ZFMK

New conopid records from the Afrotropical Region (Diptera) — Part 3: Physocephalini 211

RESULTS

CONOPINAE Macquart, 1834

The genera of Afrotropical Conopinae can be identified

using the key of Stuke (in press).

Physocephalini Smith & Peterson, 1987

To date three Afrotropical genera have been placed in

the tribe Physocephalini, two of which (Dacops, Pseu-

dophysocephala) are endemic or almost endemic to the

region (Stuke 2016). The cladistic analysis of Gibson &

Skevington (2013) concluded that Physocephalini is

monophyletic, a position which has not been rejected

since the tribe was introduced by Camras (1965) and sub-

sequently defined by Smith & Peterson (1987).

Krober (1939) differentiated the genus Pseudophys-

ocephala only within a key, based on the characters

“Kurze, gedrungene Arten mit grossem, flachem Kopf

und mit lackartig glanzendem Einschnitt am Hinterrand

des Auges” [short, stocky species with large, flat head

and with shining depression at hind margin of eye]. The

genus Pseudophysocephala was subsequently only re-

ferred to in publications by Camras (1962b, 2001), where

he distinguished the genus using almost the same charac-

ter set: “Head short (flat). Front, face, and cheek relative-

ly narrow. Front usually not higher than eye. Indentation

and triangle of eye large. Terminal female abdominal

segments moderately to very aberrant.” Camras (1962b)

stated that “By using the head characters, Pseudophyso-

cephala has been maintained, although the intergradation

of any one character is complete”. All of the diagnostic

characters used by both Camras and Kroéber are variable

within both Physocephala and Pseudophysocephala,

however, and although it is possible to recognise some

distinct species-groups with obvious characters such as

an aberrant theca or characteristic setae on tarsi, legs or

pleura, none of the diagnostic characters mentioned by

these two authors, nor any combination of these charac-

ters, is actually suitable to differentiate two genera. The

type species of the genus Pseudophysocephala, Conops

platycephalus Loew, 1853, is particularly intermediate

and indeed very hard to identify at all. This difficulty has

resulted in many problems in species identification given

that the separation of Physocephala and Pseudophyso-

cephala is the starting point in the recent keys of Camras

(1962b, 2001). Pseudophysocephala and Physocephala

fall together in the phylogenetic tree of Gibson & Skev-

ington (2013). In their analysis, however, only two Pseu-

dophysocephala species were included, both of which are

treated in this paper as synonyms and neither of which

reflects the huge morphological variability present within

the genus. Only one character was mentioned by these

authors to distinguish Pseudophysocephala and Physo-

Bonn zoological Bulletin 68 (2): 209-273

cephala: [95-1] “Narrow female abdominal segments

5—7 present”. This character does not in fact distinguish

the only species they used in their analysis from many

Physocephala species, however, and is absent in several

other Pseudophysocephala species. As a result, we here-

with treat Pseudophysocephala Krober, 1940 as a junior

synonym of Physocephala Schiner, 1861 (syn. nov.).

Both of the known Dacops species are characterised by

a single synapomorphy — @ postabdomen with a unique

ventral spoon-shaped structure (Fig. 3) — and they are

therefore without doubt sister species. No synapomor-

phic character has ever been found to show Physocepha-

la to be monophyletic when excluding Dacops, however.

Gibson & Skevington (2013) argued that Physocephala

(including Pseudophysocephala) was monophyletic by

virtue of one apomorphic character: [51-1] “metafemur

distinctly broadened basally”. This character is suitable

for diagnosing the tribe Physocephalini, but the slight-

ly broadened hind femur of Dacops also falls within

the variability found in this tribe. It may be that Phys-

ocephala without Dacops is paraphyletic, but no char-

acter has been found which consistently separates the

two genera, and no taxonomic revision or phylogenetic

study has so far considered enough species to cover the

full range of variation. We therefore synonymise Dacops

Speiser, 1923 with Physocephala Schiner, 1861 herewith

(Syn. nov.).

With the introduction of these new synonyms only the

genus Physocephala remains in the Afrotropical Physo-

cephalini. There are clearly distinct species-groups with-

in this genus, as previously described by Camras (2001)

and as indicated in Key 1, and it is possible that in future

at least some of these will be found to be monophylet-

ic, but probably not all. A full and detailed phylogenetic

analysis which includes most of the Afrotropical spe-

cies, and which follows the important comments made

by Borkent (2018: 113), will be necessary in the future

in order to fully clarify the position and, where neces-

sary, allow the rational division of Physocephala into

well-supported genera and/or subgenera.

Physocephala Schiner, 1861

= Pseudodacus Krober 1915, homonym of Pseudodacus

Hendel, 1914 [Tephritidae]

= Dacops Speiser, 1923 (syn. nov.)

= Archiphysocephala Krober, 1939 [Camras 1957,

Smith & Cunningham-van Someren 1970]

= Pseudophysocephala Krober, 1940 (syn. nov.)

Although Physocephala species are among some of the

most conspicuous and beautiful Conopidae the identifi-

cation of species is very difficult due to high infraspecific

variation and the lack of stable characters to divide the

genus in groups. Key | presents a new attempt to distin-

guish species-groups but we are very aware that this re-

©ZFMK

212 Jens-Hermann Stuke & David K. Clements

mains problematic. In many cases it will not be possible

to identify single specimens of difficult species without

comparative material. Much patience is necessary, and

no little frustration encountered, when identifying Afro-

tropical Physocephala!

It should be noted that the species-groups given be-

low are not intended to represent natural monophyletic

groupings but are merely an attempt to provide a starting

point for the identification of species within this large ge-

nus. Where our species-groups are identical to those pre-

viously identified by Camras (2001) we have retained the

same species-group name, but where they are constituted

or interpreted differently we have used a different name

in order to minimise confusion in the future.

Key 1 — Identification of Afrotropical Physocephala

species-groups

1. Mediotergite with distinct setae (e.g. Fig. 148); tibiae

may have black setulae arranged to form a dorsal

line (e.g. Fig. 140); vertex occupying almost half

length of frons (except in brevivertex); semi-circular,

setose and may have distinct ocellar scar (e.g. Figs

129, 131); apical aristomere may be extremely long

in some species, twice as long as first aristomere

including ventral projection (e.g. Figs 132, 149):

wing completely covered with microtrichia; cell

r,,, basal cell and basal medial cell completely dark

brown to black (e.g. Fig. 142); facial carina medially

black to brown.............. P. pubescens species-group

— Mediotergite at most with some barely visible short

setae; tibiae lack black setulae arranged in dorsal

lines; vertex clearly shorter than half length of

frons; other characters variable (e.g. Fig. 15); apical

aristomere never extremely long, much less than

twice length of first aristomere including ventral

projection; wing in some species with areas lacking

microtrichia; wing less darkened in some species;

facial carina completely yellow in some species .... 2

2. Knob of haltere at least partly velvety black (e.g.

Fig. 67); arista three-segmented (e.g. Fig. 39); pleura

hacks=vertical" (us tne Stripe lec)... erred. clues «i ceeps 3

— Knob of haltere at most with some obscure

darkening but never velvety black; arista with two or

three distinct aristomeres; pleura may have vertical

GUSTNGSUMIPE CI FIG ONeill dolores ti scuel 4

3. Frontoclypeal tubercle larger than adjoining

lateral facial groove, dorsally rounded and lacking

keel (Fig. 37); hind margin of eye lacks a shining

triangular indentation; scape shorter, about 2.5

times as long as high, apical aristomere as shown

in Fig. 39; vertex short, and with longitudinal

grooves (Fig. 38); wing as in Fig. 41: basal cell and

basal medial cell completely brown and covered

Bonn zoological Bulletin 68 (2): 209-273

with microtrichia; vena spuria starts in middle of

crossvein rm and reaches the hind margin of cell r,, .,

therefore separating off a narrow triangular area; 3

with distinctly pointed tip to abdomen in side view

(Fig. 40); larger species, abdomen not so obviously

narrow and elongated (Fig. 40); wing length 15-

ZO TYTHs 27e: 2 00 208. P. bimarginipennis species-group

Frontoclypeal tubercle smaller than adjoining lateral

facial groove, dorsally with sharp keel (e.g. Fig. 66);

hind margin of eye has distinct shining triangular

indentation (e.g. Fig. 80); scape long, about four

times as long as broad (e.g. Fig. 66), apical aristomere

elongated; vertex not obviously short and at most

with a few longitudinal grooves; wing as e.g. Fig.

70: basal cell and basal medial cell partly hyaline and

partly without microtrichia; vena spuria starta close

to hind margin of cell r,,., therefore not separating

off any distinct area; 6 abdomen with rounded tip

in side view (1.e. lacking distinct point; e.g. Fig.

67); smaller species with very narrow and elongated

abdomen reaching well beyond wing tips (e.g. Fig.

OP) pawingslenothy =) OAM tere comes, some oe one

lets Aenean AY aad Jes te P. halterata species-group

Vena spuria in cell r,,, usually well developed,

starting in anterior half of radial-medial crossvein

and therefore distinctly separated from media at least

in basal % of cell r,,. (e.g. Fig. 118). In occasional

specimens where the radial-medial crossvein 1s

reduced, this character may not be distinct; cell

r,,, hyaline between media and vena spuria (e.g.

Fig. 118, 122) and sometimes lacking microtrichia

there; dark species lacking distinct colour pattern

(e.g. Fig. 108, 122); scutum and scutellum black to

brown, tergites 1-3 dark brown to reddish-brown,

tergites 3-6 black to dark brown (except in P. rufa,

which is principally reddish-brown); abdomen lacks

obvious dense dusting, only tergite 3 may have

posterolateral silver-grey dusted spots (e.g. Fig. 108,

122); anterior part of abdomen obviously narrow and

elongated (e.g. Fig. 122); hind margin of eye with

distinct shining triangular indentation, and occiput

obviously bulging forward at this point; scape

elongated, at least as long as protruding part of face;

gena very narrow, not broader than maximum width

of proboscis; frons with darker black or brown mark,

or indistinct light brown marking; mediotergite

usually has some barely visible short setae; 2 fore

tarsi sometimes obviously broad and/or with unusual

structures such as long setae (e.g. Fig. 114) or long

and narrow or spine-like pulvilli (e.g. Fig. 120);

hind tarsi sometimes extremely short (e.g. Fig.

111); & theca reduced, aberrant and not protruding

far ventrally (e.g. Figs 110, 115, 124); @ sternite 5

sometimes v-shaped or u-shaped posteriorly ............

OO 7 Hae, XY MPO CF P. pilitarsis species-group

©ZFMK

New conopid records from the Afrotropical Region (Diptera) — Part 3: Physocephalini 213

Characters never in the above combination; vena

spuria in cell r,,. sometimes indistinct or missing

(e.g. Fig. 100); cell r,,, sometimes completely brown

between media and vena spuria, if vena spuria 1s

developed at all; several species are paler and have a

distinct colour pattern, e.g. orange scutum with black

markings; abdomen usually with obvious dusting,

tergites 3-4 with densely dusted hind margin and

at least tergites 5-6 with obvious golden dusting;

nterior part of abdomen sometimes wider and shorter

(e.g. Fig. 57); if hind margin of eye has a distinct

shining triangular indentation, then occiput usually

less obviously bulging forward at this point;scape

sometimes shorter than protruding part of face; gena

sometimes wider than maximum width of proboscis;

frons may lack dark marking; mediotergite lacks any

short setae in several species; @ tarsi lacking unusual

characters; 9 theca in almost all species normally

developed and obviously protruding ventrally; ¢@

sternite 5 straight-edged or only slightly concave

PO SLETIOT Ig a ee hcg na ee sionals eandee to sete 5

2 postabdomen ventrally with unique protruding

spoon-shaped structure (e.g. Fig. 3), theca absent; ¢

abdomen slightly pointed in side view (e.g. Fig. 2);

epandrium long, small cerci reaching only about 1/4

of the length of epandrium (e.g. Fig. 4); hypandrium

sheath narrow and elongated; distiphallus elongated

and aedeagus may therefore be obviously extruded

(e.g. Fig. 5); cell r,,, only dark in apical half (e.g.

Fig. 1) or cell r,,, completely dark (e.g. Fig. 6)........

Ly hopsotda tabi hitatd's aatttncal oben P. abdominalis species-group

2 postabdomen ventrally lacks protruding spoon-

shaped structure, but has theca; 3 abdomen not

pointed in side view but rounded (e.g. Fig. 66, 71);

epandrium smaller and therefore cerci reaching more

than 1/4 of the length of epandrium; hypandrium

sheath different (as far as is known) and distiphallus

less elongated; wing pattern may be different, or

SAINIC S AION Gah erry A aes Hts ee id YS 8 SUN med 6

Arista extremely short, appearing as if broken (e.g.

Figs 16, 32); aristomeres all shorter than height of

basal aristomere; in most species only two visible

aristomeres; scape in most species shorter, about

twice as long as high (e.g. Fig. 13); shining triangular

area at hind margin of eye barely developed or

absent; vertex anteriorly with more or less distinct

longitudinal groove (e.g. Fig. 15); radial-medial

crossvein usually not obviously short and never

completely reduced (e.g. Figs 12, 20, 21); basal cell

may be partially bare of microtrichia (e.g. Figs 9, 10);

anepimeron may have setulae; 3 sternite 8 slightly to

distinctly bulging over protandrium; typically with

very fine black setulae (distinctly smaller than the

scattered setulae on protandrium) arranged in distinct

Bonn zoological Bulletin 68 (2): 209-273

line at border between protandrium and _ sternite

8; 3 epandrium as e.g. Figs 7-8, 22-23: blackish

teeth on posterior margin of epandrium with more

or less broad blackened bases; epandrium with long

setulae anteromedially and dorsally; in lateral view

large apically and almost semi-circular; postgonite

evagination on inner side with broad-based spines

giving this structure a fish-scaled appearance (may

be hard to see in strongly macerated specimens,

character not proven in all species); 2 postabdomen

lacking obvious reductions or aberrant theca (e.g.

Fig. 14, 19), tergite 6 always large and elongated

CSET ON cra ise: Pee eS es iw a ee ees 5 |

Arista usually longer and v-shaped (e.g. Figs 48, 65),

both aristomeres distinctly longer than high; scape

sometimes more than twice as long as high (e.g.

Figs 47, 66); distinct shining triangular area at hind

margin of eye (e.g. Figs 72, 80); vertex lacks anterior

longitudinal groove in some species (e.g. Figs 84,

85, 87); radial-medial crossvein sometimes very

short or absent (e.g. Fig. 100); basal cell completely

covered with microtrichia; anepimeron lacks

setulae; 3 shining sternite 8 not usually bulging

over protandrium; border between protandrium and

sternite 8 usually lacks line of fine black setulae; ¢

epandrium sometimes different to above (but is

unknown in some species); 2 postabdomen may

have obvious reductions or an aberrant theca, tergite

6 sometimes very small (e.g. Fig. 55)... 8

Hind coxa not dusted, or at least not more densely

dusted than middle and fore coxae; pleura usually

lacks dense dusting; if dusting stripe is present

it starts at a point clearly separated from middle

coxa and becomes narrower before reaching

notopleuron; typically, mediotergite dorsally lacks

dense dusting connecting with dense dusted spot

on katatergite; costal cell and subcostal cell always

hyaline, obviously paler than base of cell r,,,. cell r,,,

typically hyaline distally, with isolated spot around

METIS te NMG characte: P. vittata species-group

Hind coxa more densely dusted than middle and

fore coxae (e.g. Fig. 14); pleura usually with dusting

stripe running vertically from middle coxa and often

reaching notopleuron without narrowing; typically,

mediotergite dorsally with dense dusting connecting

with dense dusted spot on katatergite; wing with

different colour pattern..... RP antiqua species-group

Ventral projection of basal aristomere towers over

apical aristomere (e.g. Figs 48, 50, 58); radial-medial

crossvein always distinct; vena spuria in cell r,,.

distinct, starting at radius R,,. and usually fused with

or closely approximated to media; wing membrane

between vena spuria and media hyaline (e.g. Fig. 49);

scutum usually with two sublateral dusting stripes

©ZFMK

214

starting at inner side of postpronotum and fused

before scutellum (e.g. Fig. 42); 2 theca normally

developed but usually depressed towards or against

the abdomen (..e. not projecting perpendicularly, e.g.

Fig. 54); 2 abdomen obviously short, tergites 5 and

6 much shorter than tergite 4 (e.g. Figs 55, 57)........

SEARS, oeaas oer TaN ore P. caenoneura species-group

— Apical aristomere usually longer than ventral

projection of basal aristomere (e.g. Fig. 79); radial-

medial crossvein sometimes reduced or absent

(e.g. Fig. 100); vena spuria in cell r,,, absent, or if

developed then membrane between vena spuria and

media not hyaline (e.g. Fig. 83); scutum may lack

sublateral dusting stripes; 9 theca sometimes very

reduced, or projects more perpendicularly from

abdomen (e.g. Fig. 77, 102); 2 abdomen may be

short or long ................. P. microvena species-group

Physocephala abdominalis species-group

Species of the Physocephala abdominalis group were

previously placed in the genus Dacops. Females are

easily recognised by the unique spoon-shaped structure

which projects ventrally at the apex of the abdomen, and

the lack of any theca (Figs 2, 3). The precise morpho-

logical derivation of the spoon-shaped structure 1s not

obvious to us, and does not appear to have been estab-

lished elsewhere, but given the scarcity of the available

material we have not carried out any dissections as the

structure is clearly visible without. Males are much more

difficult to diagnose because all of the characters used

in Key 1 are variable, and atypical forms occur. Males

of Physocephala kaplanae may occasionally have a re-

duced radial-medial crossvein and barely developed vena

spuria, making them easy to misidentify as a member of

the microvena group, although the characteristic male

postabdomen shape, as described in Key 1, should never-

theless allow these to be safely assigned. All members of

the abdominalis species-group are confined to the Afro-

tropical Region.

Key 2 — Identification of the Physocephala abdomina-

lis species-group

1. Basal cell completely hyaline and not covered with

microtrichia (Fig. 1); wing between costa and radius

R,,, hyaline (Fig. 1); black colouration on scutum not

reaching hind margin; overall appearance reddish-

brown, but may sometimes be brownish to blackish

P. abdominalis (Kroéber, 1915)

— Basal cell completely or almost completely dark, and

completely covered with microtrichia (Fig. 6); wing

between costa and radius R,,, dark (Fig. 6); black

colouration on scutum may reach to hind margin;

overall appearance black....... P. kaplanae (Camras,

2001)

Bonn zoological Bulletin 68 (2): 209-273

Jens-Hermann Stuke & David K. Clements

P. abdominalis (Kroéber, 1915)

(Figs 1-5)

Pseudodacus abdominalis Krober 1915

= Pseudodacus apicalis Krober, 1915

= Physocephala fascipennis Brunetti, 1925

= Conops patelliformis Séguy, 1933

Primary type material examined. |< syntype of Phys-

ocephala fascipennis Brunetti, 1925: (1) “Syn - / type”;

(2) “12.11.11 / Caia / Zambesi / Dhawar / H. Swale.”; (3)

“Pres. by / Impl. Bureau Ent. / 1915 - 164”; (4) “Physo.

/ fascipennis / Brun. Type ¢ / Det. E. Brunetti 1924”; (5)

“BMNH(E)# / 249047”: coll. NHML.

12 syntype of Physocephala fascipennis Brunetti,

1925: (1) “Syn - / type”; (2) “Nyasaland / Cholo / R. C.

Wood”; (3) “Pres. by / Imp. Bur. Ent. / Brit. Mus. / 1924

- 306”; (4) “Physo. / fascipennis / Brun. Type & / Det. E.

Brunetti 1924”; (5) “BMNH(E)# / 249048”; coll. NHML.

Additional material. BENIN: 19, vi.2002, Pénéssou-

lou, forest area [09°15’58.26"N 01°33’04.81"E], leg. G.

Goergen, coll. IITA; 1’, 28.i.2018, Togbota, leg. G. Go-

ergen, coll. IITA; DEMOCRATIC REPUBLIC OF CON-

GO: 14, iv.1913, Congo da Lemba, det. as P. fascipennis

by Brunetti 1925, leg. R. Mayné, coll. MRAC; 16), 1942,

Bas-Congo, Mayidi, leg. P. Van Eyen, coll. ISNB; 19,

1945, ditto; ETHIOPIA: 19, 111.1969, Bahir Dar, leg.

Schauffele, coll. SMNS; 2¢'¢', 30.-31.v.2015, Sof Umer

[6°54'N 40°51'E], 1200 m, leg. J. Halada, coll. CULSP;

KENYA: 16, xi.1948, Garissa-Bura, Tana River, det.

as D. abdominalis by Camras 1999, leg. van Someren,

coll. NHML [NHMUK010922084]; 16, 12.x.1998,

Morigat, leg. F. Kaplan, A. Freidberg, coll. TAUI; 19,

12. —26.11.2005, Nyanza Province, Ungoye Field Station

[0°36.91'S 34°05.52'E], 1147 m, leg. R. Copeland, coll.

NMKE; MALAWI: 16, 11.v.1916, Nyasaland, Ruo Val-

ley, 1000-2000 ft, des. as syntype of P. fascipennis, leg.

R. C. Wood, coll. NHML [NHMUK010922076]; MO-

ZAMBIQUE: 19, 19.—30.iv.2015, Sofala pr., Gorongosa

Park, small lake [18°56'39"E 34°26'35"E], 30 m, Mal-

aise trap, leg. M. Hauser, A. Rung, coll. PMHA; SOUTH

AFRICA: 16, i.2000, Mondl Forest, Hilton, KZN, leg.

Lyawb, coll. PMHA; 1 specimen, 1.-111.1927, Natal,

Weenen, det. as D. abdominalis by Krober 1930, leg.

H. P. Thommaset, coll. NHML [NHMUK010922090];

192, xii.1923, Natal, Weenen, det. as D. fascipennis

by Brunetti 1924, leg. H. P. Thommaset, coll. NHML

[NHMUK010922093]; 1 specimen, 2.x.1915, Umbilo,

Durban, Natal, des. as syntype of P. fascipennis, [col-

lector unknown], coll. NHML [NHMUK010922077];:

14, iv.1955, Cape Province, Grahamstown, det. as

D. fascipennis by Smith 1957, leg. Allison, coll. NHML

[NHMUK010922088]; 14, 26.xi.1958, Cape Province,

Grahamstown, leg. C. Jacot-Guillarmod, coll. AMGS;

14, 2.xii.1958, ditto; 12, 17.xi.1961, Cape Province,

Grahamstown, leg. E. McC Callan, coll. AMGS; 16,

©ZFMK

New conopid records from the Afrotropical Region (Diptera) — Part 3: Physocephalini 215

19.xu1.1971, Cape Province, Grahamstown, det. as D. ab-

dominalis by Camras 2000, leg. D. J. Greathead, coll.

NHML [NHMUK010922085]; 192, 26.11.1987, Cape

Province, Grahamstown, leg. P. E. Hulley, coll. AMGS;

14, 7.vii.2002, Cape Province, Grahamstown [33.17°S

26.31°E], leg. H. V. Lewis, coll. AMGS; 19, 15.1x.2006,

Cape Province, Grahamstown [33°23'S 26°29'E], leg. A.

McClure, coll. AMGS; 13, 4.iv.1981, Cape Province,

Grahamstown [33°19'S 26°31'E], leg. G. T. Lloyd, coll.

AMGS; 19, 10.i11.1981, Cape Province, Grahamstown,

200 Dept. window [33.19S 26.32E], leg. E. Nieman, coll.

AMGS; 264, 22.i1.1970, Cape Province, Grahamstown,

—— 7 ie! ra ea ae

7 ae Pe | ee, 2

- “= : ald ——

a ~ - .

Figs 1-5. Physocephala abdominalis (Krober, 1915). 1. Wing, dorsal view (4, Belmont Valley); 2. Tip of 3 abdomen, lateral view

(34, Belmont Valley); 3. Spoon-shaped structure of 9 postabdomen (9, Zambesi Valley); 4. Epandrium, dorsal view (d', Graham-

stown); 5. Postabdomen, lateral view (4, Grahamstown). ce = cercus; dp = distiphallus; ep = epandrium; pa = phallapodeme; hya =

hypandrial arm; psh — phallus sheath.

Bonn zoological Bulletin 68 (2): 209-273 ©ZFMK

216 Jens-Hermann Stuke & David K. Clements

Belmont Valley, on flowering Foeniculum vulgare Mill.,

leg. F. W. Gess, coll. AMGS; 26'3', 23.i.1970, Cape

Province, Grahamstown, Belmont Valley, on flowering

Foeniculum vulgare Mill., leg. F. W. Gess, coll. AMGS;

224, 26.1.1970, Cape Province, Grahamstown, Belmont

Valley, on flowering Foeniculum vulgare Mill., leg. F. W.

Gess, coll. AMGS; 13 19, 25.i1.1972, Cape Province,

Grahamstown, Belmont Valley, leg. F. W. Gess, coll.

AMGS; 1, 6.1.1977, Cape Province, Grahamstown, Hil-

ton [-33.310629 26.525595], on Acacia karroo flowers,

leg. D. W. Gees, coll. AMGS; 1, 2.xii.1979, Cape Prov-

ince, Grahamstown, Hilton [-33.310629 26.525595], leg.

F. W. Gees, S. K. Gees, coll. AMGS; 14, 6.—14.1.1972,

Cape Province, Grahamstown, Howison’s I'oort, leg. F.

W. Gess, coll. AMGS; 12, 19.-22.xi-1971, ditto; 14,

8 —9.1v.1979, Cape Province, Salt Vlei, Port Alfred, leg.

A. E. Mel. Collan, coll. AMGS; 13, 19.xi.1999, East-

ern Cape Province, 37 km nw of Steytlerville [33°11'S

24°10'E], 695 m, leg. M. Hauser, coll. PMHA; 16,

1.—10.11.1933, Eastern Cape Province, Katberg, det.

as D. abdominalis by Krober 1938, leg. R. E. Turner,

coll. NHML [NHMUK010922087]; 14, 1.-10.ii.1933,

Eastern Cape Province, Katberg, det. as D. abdom-

inalis by Krober 1938, leg. R. E. Turner, coll. NHML

[NHMUK010922089]; 19, 1—10.11.1933, Eastern Cape

Province, Katberg, det. as D. abdominalis by Krober 1938,

leg. R. E. Turner, coll. NHML [NHMUK010922092]:

192, 1—10.11.1933, Eastern Cape Province, Katberg, det.

as D. abdominalis by Krober 1938, leg. R. E. Turner,

coll. NHML [NHMUK010922094]; 14, 1.-10.ii.1933,

Eastern Cape Province, Katberg, det. as P. platycephala

by Krober, 1938, det. as P. rufitarsis by Camras 2000,

leg. R. E. Turner, coll. NHML [NHMUK010922157];

14, 26.ix.2010, Free State Province, Bloemfontein, 10

Jan Venter Street [29°06'12"S 26°08'42"E], leg. R. J.

Nuttall, coll. BMSA; 19, 26.ix.2010, Free State Prov-

ince, Bloemfontein, 10 Jan Venter Street [29°06'12"S

26°08'42"E], on window, leg. R. J. Nuttall, coll. BMSA;

14, xii.2009, Free State Province, Bloemfontein, Nation-

al Museum Library [29°6'54.35"S 26°13'9.72"E], leg. L.

Coetzee, coll. BMSA; 19, 13.xii.2012, KwaZulu-Natal

Province, Pietermaritzburg, Kwela Lodge, 940 m, leg. B.

Lechner, coll. PASS; 1, 22.-25.xi.2003, Mpumalanga

Province, 30 km ne Lydenbrug near Ohrig, leg. J. Hala-

da, coll. CULSP; 19, no date, Natal Province, Howick,

des. as syntype of P. fascipennis, leg. J. P. Gregor, coll.

NHML [NHMUK0O10922078]; 16, 14.i.1953, Natal

Province, St. Michaele, det. as D. fascipennis by Smith

1957, on flowers of Scutia myrtina, leg. E. McC Callan,

coll. NHML [NHMUK010922086]; 14, 28.iv.2002,

Western Cape Province, Nature Valley, 33 23 DC, leg.

L. de Wet, coll. AMGS; TANZANIA: 1, 10.v.1956,

Old Shinyanga, Block 9, det. as P. caenoneura by Emden

1950, leg. E. Burtt., coll. NHML [NHMUK010922143];

ZIMBABWE: 19, 13.11.1936, Salisbury [= Hara-

re] [17.84 31.05], leg. W. L. Williams, coll. NHML

[NHMUK010922091]; 13, 24.vii.1958, Salisbury [=

Harare] [17.84 31.05], leg. A. D. Graham, coll. AMGS;

12, 1.1987, Zambesi Valley, Rekometjie [16°10'S

29°25'E], leg. S. GuBmann, coll. ZFMK; UNKNOWN

LOCATION: 19, 27.xii.1937, [characters illegible], det.

as D. abdominalis by Krober 1938, leg. A. I. Bevis, coll.

NHML [NHMUK010922095].

P. kaplanae (Camras, 2001)

(Fig. 6)

Dacops kaplanae Camras 2001

Material. CENTRAL AFRICAN REPUBLIC: 19,

17.x11.2008, 45 km e Nola [03°40'N 16°26'E], 570 m,

leg. J. Halada, coll. CULSP; DEMOCRATIC REPUB-

LIC OF CONGO: 16), 1968, Tshuapa, Bamanya, leg. P.

Hulstaert, coll. MRAC; ETHIOPIA: 19, 4.xi.2018, Bahir

Dar [11.597292°N 37.355696°E], leg. G. Goergen, coll.

IITA; 19, 11.xi1.2014, Bonga, StraBensaum, 1955 m

[07°14'09"N 036°16'36"E], leg. H.-J. Fligel, coll. PHJF;

14, 20.ix.2012, Chencha, 1916 m [06°09'N 37°34'E],

leg. A. Pauly, coll. ISNB; 1, 1912, Harar, leg. S. V. Kris-

Fig. 6. Physocephala abdominalis (Krober, 1915). 1. Wing, dorsal view (3', Belmont Valley); 2. Tip of 4 abdomen, lateral view

(4, BelmoWing of Physocephala kaplanae (Camras, 2001), dorsal view (Monts de Christal).

Bonn zoological Bulletin 68 (2): 209-273

©ZFMK

New conopid records from the Afrotropical Region (Diptera) — Part 3: Physocephalini

tensen, coll. ZMHB; 19, 18.x.1957, near Axum, paratype

of P. kaplanae, det. as D. abdominalis by Smith 1969,

leg. D. J. Greathead, coll. NHML [NHMUK010922075];

GABON: 16, 11.-12.1i1.1990, Wolen-Ntem, Monts de

Christal, Tchimbelé, 600 m [00°37'N 10°24'E], leg. J. J.

Wieringa, coll. RMNH.

It is possible that P kaplanae is nothing more than a

darker form of P. abdominalis, although we are not aware

of any intermediates and therefore accept the validity of

this species.

Physocephala antiqua species-group

All of those species which Camras (2001) placed in his

maculipes and similis species-groups are herewith includ-

ed within the antiqua species-group. The segregation of

the Camras species-groups is difficult or impossible due

to the extreme variability of the characters used to sepa-

rate them, and it is therefore more convenient to combine

them. In general, members of the antiqua species-group

have a very typical Physocephala habitus lacking any

atypical characters other than an extremely short aris-

ta. Careful evaluation of the character combination set

out in Key 1 is necessary to distinguish members of this

group, however, especially males. Several species of this

group are restricted to the Afrotropical Region but oth-

ers have a wider distribution which reaches into southern

and central Europe, with some also reaching eastwards as

far as China and Mongolia. The species-group also con-

tains several Palaearctic species which are not currently

known from the Afrotropical Region.

Key 3 — Identification of the Physocephala antiqua

species-group.

1. Basal cell completely covered with microtrichia, and

usually brownish in colouration...........000.....000000. 2

— Basal cell almost completely (or at least centrally)

bare of microtrichia and hyaline ..........0.0....00.00. 4

2. Wings obviously reduced in length, not reaching

beyond ‘apex of tereite 33(PiG AT) ee cs cee etait ben

Ci et ee en eee oe P. brevipennis Camras, 1962

— Wing not obviously reduced, about as long as

abdomen or at least extending beyond apex of tergite

3. Cell r,,, completely brown (Fig. 21); subcosta light

yellow to yellowish-brown, obviously different

in colour from radial veins (Fig. 21); legs orange-

brown, lacking a blackish ring on hind femur or

darkened tarsi (Fig. 14); scutum orange-brown with

black central stripe and small two lateral black spots

which may sometimes fuse with mid-stripe (Fig. 17);

Bonn zoological Bulletin 68 (2): 209-273

217

scutellum orange-brown (Fig. 17); larger species,

wing length 6.3—7.6 mm....P. guillarmodi spec. nov.

— Cell r,,, hyaline apically (Fig. 36); subosta dark

brown, same as radial veins (Fig. 36); at least hind

femur with black markings, and apical tarsi black;

scutum and scutellum sometimes blackish; smaller

species, wing length 3.7—5.3 mm (based on 3 known

AIHOLOPICAly SPECIMENS ONIN) ose eee te eM oi ante’

$6 satire a Beste has tinallon Roath P. pusilla (Meigen, 1804)

4. Anepimeron with long setulae, about as long as

maximum width of tibiae; dusting stripe on pleura

distinct, narrow at anepisternum and broader when

reaching notopleuron (Fig. 34), obviously contrasting

with shining anterior half of anepisternum and

anepimeron (Fig. 34)............. P. ssymanki spec. nov.

— Anepimeron at most with short setulae; any dusting

stripe on pleura reaching notopleuron without

widening, and sometimes indistinct. .......0000.00.000. 5

5. 3 aristomeres, the basal one minute and easily

overlooked; scape usually about three times as long

as high (Fig. 28); gena black, and parafacia above

gena mainly yellow (Fig. 28); frons with black

longitudinal stripe reaching from vertex to base

of antennae (Fig. 26); black facial stripe divided

ventrally and reaching to mouth edge (Fig. 28);

abdomen longer (Fig. 25), ratio of combined dorsal

length of tergites 2+3 : dorsal length of tergite 4 in J

24a os} fy Fare onceediparioe tarda P._larvata (Speiser, 1911)

— 2aristomeres (e.g. Fig. 28); scape usually only about

twice as long as high; gena and frons lack black

markings; abdomen shorter, ratio of combined dorsal

length of tergites 2+3: dorsal length of tergite 4 in ¢

SID a EP scr Ja aie ERB shyt IS EET ITR sce Paeke lee 6

6. Wing completely hyaline (Fig. 12) ........0000. ee.

gras nd RV Alek la P. claripennis Becker, 1923

— Wing with distinct brown to blackish markings (Figs

ea EG hoe eee sy OPT Bi P. antiqua (Wiedemann, 1830)

P. antiqua (Wiedemann, 1830)

(Figs 7-10)

Conops antiqua Wiedemann 1830

= Conops maculipes Bigot, 1887 syn. nov.

= Conops interrupta Bezzi, 1901

= Conops erythraspis Bezzi, 1901

= Physocephala flavifacies Krober, 1915

= Physocephala limbata Krober, 1915

= Physocephala rubicunda Krober, 1915

= Physocephala madagascariensis Krober, 1915 syn. nov.

= Physocephala gracilia Krober, 1915 syn. nov.

= Physocephala decisa Brunetti, 1925

= Physocephala brevistylata Krober, 1931

= Physocephala minutissima Krober, 1933 syn. nov.

= Physocephala nigroscutellata Krober, 1933

©ZFMK

218 Jens-Hermann Stuke & David K. Clements

Primary type material examined. 9 holotype of Con-

ops maculipes Bigot, 1887: (1) “Holo - / type”; (2) “S.

Africa: / Cape of Good Hope. / ex coll. J. Bigot / ex

coll. G. H. Verrall / B.M. 1914-500 [strikethrough]”; (3)

“C. maculigera 2 / Cap. B. Hop.”; coll. NHML.

S holotype of Physocephala rubicunda Krober, 1915:

(1)’3055”; (2) “Type”; (3) “Physocephala 3 / rubicunda

Krb. / O. Krober det. 1914”; coll. ZMHB.

3 syntype of Physocephala decisa Brunetti 1925: (1)

“Syn- / type”; (2) “N. E. Rhodesia. / Upper Luangwa R.

/ 27 July 13 Aug. 1910. /S. A. Neave”; (3) “Physoceph-

ala / decisa / Brun. Type 6 / Det. E. Brunetti 1924”; (4)

“BMNH(EB)# / 249074”; coll. NHML.

9 syntype of Physocephala decisa Brunetti 1925: (1)

“Syn- / type”; (2) “N. E. Rhodesia. / Upper Luangwa R.

/ 27 July 13 Aug. 1910. /S. A. Neave”; (3) “Physoceph-

ala | decisa/ Brun. Type & / Det. E. Brunetti 1924”; (4)

“BMNH(E)# / 249077”; coll. NHML.

9 syntype of Physocephala decisa Brunetti 1925: (1)

“Syn- / type”; (2) “Rusapi / S. Rhodesia / 19.12.1920 /

Rhodesi / Museum”; (3) “Pres. by. / Imp. Bur. Ent. / Brit.

Mus. / 1924-306”; (4) “Physocephala / decisa / Brun.

/ Type & / Det. E. Brunetti 1924”; (5) “BMNH(E)# /

249075”; coll. NHML.

S holotype of Physocephala minutissima Krober,

1933: (1) “1” “Holo- / type”; (2) “Type”; (3) “8.6.16 /

Ruo / Nyasaland / 200 ft / R. C. Wood / 601.”; (4) “Pres.

by. / Imp. Inst. Ent. / Brit. Mus. / 1931-480”; (5) “Phys-

ocephala | minutissima / 3 Krb”; (6) “BMNH(E)# /

249071”; coll. NHML.

9 holotype of Physocephala nigroscutellata Krober

1933: (1) “Holo- / type”; (2) “Type”; (3) “Lady Grey /

1 Feb. 1924/R. I. Nel”; (4) “Pres. by / Imp. Inst. Ent. /

Brit. Mus. / 1932-143.”; (5) “Phsocephala / nigroscute-

Ila [sic] / 2° Krb / det. Kréber 1931”; (6) “~BMNH(E)# /

249072”; coll. NHML.

Additional material. BENIN: 19, ix.2006, Adohoun

[06°39’27.55"N 01°39°09.76"E], light trap, leg. G. Goer-

gen, coll. IITA; 14, v.2007, Athiémé [06°14‘20.00"N

01°40‘00.00"E], leg. G. Goergen, coll. IITA; 16,

vili.2006, Athiémé [06°14’20.00"N 01°40’00.00"E],

sweep netting, leg. G. Goergen, coll. IITA; 19, 14.

vi.2006, Calavi, sweep netting, leg. G. Goergen, coll.

IITA; 13), 7.viii.2006, Calavi, campus [06°26’15.00"N

02°19°42.00"E], sweep netting, leg. G. Goergen, coll.

IITA; 19, 25.x.2007, ditto, BURUNDI: 19, 24. viti.1957,

Kisenyi, leg. F. J. Francois, coll. ISNB; 19, 22.1.1950,

Rumonge, 780 m, det. as P. nigroscutellata by Janssen

1954, leg. F. J. Francois, coll. ISNB; 1@, 9.iii.1952,

Bubanza Province, 7 km s de Gihanga, 850 m, det. as

P. nigroscutellata by Janssen 1954, leg. F. J. Francois,

coll. BMSA; CAMEROON: 19, 24.iv.1914, Uamgebiet,

Bossum, leg. G. Tessmann, coll. ZMHB; 19, 12.v.1914,

Uamegebiet, Bossum, det. as P. interrupta by Krober, leg.

G. Tessmann, coll. ZMHB; CENTRAL AFRICAN RE-

Bonn zoological Bulletin 68 (2): 209-273

PUBLIC: 19, 14.v.2009, 45 km ssw Bamingui [07°15'N

20°03'E], leg. J. Halada, coll. CULSP; 14, 20.iv.2010,

Reserve Koukorou Bamingui [07°15'N 20°03'E], 440 m,

leg. J. Halada, coll. CULSP; DEMOCRATIC REPUB-

LIC OF CONGO: 1, i-11.1913, Congo da Lemba, det.

as P. decisa by Brunetti 1925, leg. R. Mayné, coll.

MRAC; ETHIOPIA: 1, 24.v.2015, 10 km nw Mega

[04°08'N 38°16'E], 1670 m, leg. J. Halada, coll. CULSP;

83.3, 11—13.v.2015, 20 km se Konsa [05°15'N 37°32'E],

850 m, leg. J. Halada, coll. CULSP; 14 299, 14.v.2015,

40 km w Konso [5°19'N 37°04'E], 600 m, leg. J. Halada,

coll. CULSP; 14, 10.v.2015, 45 km ne Arba Minch

[6°17'N 37°47'E], 1200 m, leg. J. Halada, coll. CULSP;

19, 21.iv.2016, Arsi [7°49'06"N 40°31'52"E], 886 m,

leg. J. Halada, coll. CULSP; 19, 22.v.2015, Wachile env.

[04°32'N 39°03’E], 1070 m, leg. J. Halada, coll. CULSP;

GAMBIA: 1, 4.xi.1977, Bakau at tropic Bungalow

[UTM 28PCK1790], swept in meadow rich in flowers, at

the beach, leg. Cederholm, Daniellson, Hammarstedt,

Hedqvist, Samuelsson, coll. MZLU; 19, 16.x.1999, Ki-

ang West National Park, headquarter, leg. W. Schacht,

coll. ZSMC; IVORY COAST: 19, 9.xi.1983, Badenous

Mbingué [9.50°N 5.50°W], leg. R. Summkeller, coll.

ZFMK; KENYA: 16, 27.iv.2008, Nguni, n of Ngomeni,

hand net, leg. M. Snizek, coll. PMME; 204, 29.iv.1995,

Tsavo East, leg. R. Copeland, coll. NMKE; 1¢, 10.

v1.1998, Tsavo East National Park, near Athi River

[2°37'S 38°22'E], leg. R. Copeland, coll. NMKE; 1,

22.v.2006, Coast Province, Tsavo East National Park,

near Galana River [UTM UTM 37 m 503352 9665277],

246 m, hand net, leg. P. Cerretti, D. Avesani, G. Carpan-

eto, G. Nardi, coll. PMME; 19°, 26—29.vi.1999, Eastern

Province, at Athi river [2°38.51'S 38°21.98'E], Malaise

trap, leg. R. Copeland, coll. ITA; 13’, 17.-31.viii.2005,

Nyanza Province, Ungoye, ICIPE Field Station

[0.61325°S 34.08908°E], 1127 m, leg. R. Copeland, coll.

NMKE; 16, 26.vi.—10.vii.2007, Rift Valley Province,

Sumburu Nature Reserve, near Ewaso Ng'iro River

[0.56797°N 37.53563°E], 874 m, Malaise trap, riverine

forest next to headquarter, leg. R. Copeland, coll. NMKE;

14, 12.-26.vi.2006, ditto; 14, 10.iv.1998, Samburu Dis-

trict, Samburu Serena Lodge, leg. R. Copeland, coll.

NMKE; LESOTHO: 1, 1.xi.1949, Mamathes

[-29.136617 27.845796], leg. C. Jacot-Guillarmod, coll.

AMGS; MADAGASCAR: 19, 8.x1.2003, Anteninde,

Sakahara [22°53'57"S 44°28'12"E], 395 m, leg. A. Ssy-

mank, coll. PASS; 1¢, v.1937, Bekily, leg. A. Seyrig,

coll. MNHN; 1.3, iv.1937, ditto; 14’, v.1937, ditto; 13

12, 1v.1942, Bekily, det. as P. madagascariensis by

Camras 1960, leg. A. Seyrig, coll. MRAC; 204, 9.

iv.1994, Berenty reserve 80 km w of Port-Dauphin

[25°00'S 46°18'E], leg. M. Wasbauer, coll. UCDC,

PMHA; 19, 6.x1.2003, Ihasofotsy-Flussufer, ca 10 km

wsw Ihosy [22°25'16"S 46°00'14"E], 1000 m, leg. A.

Ssymank, coll. PASS; 14), 19.iii.1994, Isalo, Analalava,

leg. A. Pauly, coll. MRAC; 19, 25.11—3.111.1968, Majun-

©ZFMK

New conopid records from the Afrotropical Region (Diptera) — Part 3: Physocephalini 219

gas. |., det. as P. madagascariensis by Camras 2000, leg.

K.M. G. & P. D., coll. NHML [NHMUK010922098];

14, 1.1992, Morarano-Chrome [17°45'S 47°59'E], leg.

A. Pauly, coll. MRAC; 1<, no date, Sambirano, leg. [?],

coll. ZMHB; 18, no date, Tananarivo, det. as P. mada-

gascariensis by Camras 1960, [collector unknown], coll.

MRAC; 19 1 specimen, 1.1952, Tzimbazaza-Tananarive,

det. as P. madagascariensis by Camras 1960, leg. R.

Benoist, coll, MRAC; 19, 28.vii—vi.vii.2001, Di-

ego-Suarez (Antsiranana), Sakalava Beach, malaise

across sandy trail in dwarf littoral forest [12°15'46"S

49°23'51"E], 10 m, leg. R. Harin'Hala, coll. CAS; 14,

41.2007, Mahayanga Province, 20 km nw Borinziny

[15°27.07'S 47°36.85'E], 37 m, hand netted in tropical

dry forest on white sand, leg. M. E. Irwin, F. D. Parker, R.

Harin'‘Hala, coll. CAS; 19, 1.xi.1993, Tamatave (Toama-

sina), Analmalotran Tamatave, leg. C. F. Kassebeer, coll.

PCFK; 19, 12.xi1.1991, Tuléar (Toliara), Ampanihy, leg.

A. Pauly, coll. MRAC; 19, 6.—16.1.2003, Tuléar (Toli-

ara), Andohaela National Park, Parcel II, Tsimela, mal-

aise trap in transitional forest, 175 m [24°56.21'S

46°37.60'E], leg. M. E. Irwin, F. D. Parker, R. Harin'Ha-

la, coll. CAS; 14), 12.iv.1968, Tuléar (Toliara), Bevilany,

300 m, det. as P. madagascariensis by Camras 2000, leg.

K.M. G. & P. D., coll. NHML [NHMUK010922097];

12, 8-18.vi.2002, Tuléar (Toliara), Beza Mahafaly Re-

serve, malaise in dry deciduous gallery forest, 165 m

[23°41.19'S 44°35.46'E], leg. R. Harin'Hala, M. E. Irwin,

coll. CAS; MALI: 19, 20.viii.1991, 10 km s Mopti, leg.

M. Schwarz, coll. PMHA; 1¢, 20.viii.1991, 10 km s

Mopti, sweep netting, leg. M. Schwarz, coll. PMHA; 13,

31.vu1.1991, 40 km sw Segou, sweep netting, leg. M.

Schwarz, coll. PMHA; MOZAMBIQUE: 244, 12.-20.

x1.2003, Manica Province, 45 km nw Chimoio, leg. J.

Halada, coll. CULSP; NAMIBIA: 19, 28.1.1993, 100 km

sw Rundu, leg. M. Schwarz, coll. PMHA; 299,

16.1.1993, 125 km sw Rundu, leg. M. Schwarz, coll.

PMHA; 299, 16.1.1993, ditto; 23.4, 4.111993, 15 kme

Swakopmund, sweep netting, leg. M. Schwarz, coll.

PMHA; 2'4, 29.iii.2000, 18 km se Stamprieton C15 to

Gochas [24.28°S 18.30°E], visiting pink flowers of Gale-

nia, leg. F. W. Gees, S. K. Gees, coll. AMGS; 224, 4.

11.1990, 30 km e Windhoek, sweep netting, leg. M.

Schwarz, coll. PMHA; 16, 12.iii.2014, 35 km w Gobabis

[22°23'S 18°39'E], 1480 m, leg. J. Halada, coll. CULSP;

14, 16.11.1990, 40 km w Witvlei, leg. M. Schwarz, coll.

PMHA; 14, 16.ii.1990, 40 km w Witvlei, sweep netting,

leg. M. Schwarz, coll. PMHA; 16, 11.iii—9.iv.1985,

6 km n Arandis, Damaraland [22°22'S 14°59'E], leg. J.

Irish, H. Rust, coll. SMWN; 19, 21.1.1993, 60 km E

Rundu, leg. M. Schwarz, coll. PMHA; 1¢, 13.ii.1990,

73 km s Mariental, sweep netting, leg. M. Schwarz, coll.

PMHA; 39°92, 25.ix.1997, e Oranjemund 28 km from

checkpoint to Sendelingsdrif [29.26°S 16.42°E], on yel-

low flowers of Deverradenudata (Viv.) Pfisterer & Podl.,

leg. F. W. Gees, S. K. Gees, coll. AMGS; 234, 20.

Bonn zoological Bulletin 68 (2): 209-273

x1.1995, Etosha National Park, 50 km ne Okaukuejo,

“Salvado” [18.9°S 16.5°E], leg. C. Schmid-Egger, coll.

PMHA; 19, 16—19.1.2007, Gobabeb, Kuiseb-Tal, at

light, leg. Mey, Ebert, coll. ZMHB; 19, 11.11.1993, Oka-

handja, leg. M. Schwarz, coll. PMHA; 19, 7.111.1999,

Karas Mountains, 6 km s on 201 from 26 [27.09°S

19.01°E], on yellow flowers of Vahlia capensis (L. f.)

Thunb., leg. F. W. Gees, S. K. Gees, coll. AMGS; 103,

25.1v.1972, Karasburg, Warmbad, SE 2818 Ba, [collector

unknown], coll. SMWN; 1, 17.-19.v.1978, Khowarib,

R., SE 1914 Ac, Kackoland, leg. S. Louw, M.-L. Penrith,

coll. SMWN; 16, 12.-14.iv.1974, Mukorob 14,

Namaland, SE 2918 Ac, [collector unknown], coll.

SMWN; 163, 15.iii.1997, near Aus on road to Helmering-

hausen [26.37°S 19.20°E], visiting white flowers of Psi-

locaulon goéareosum (Berger) Dinter & Schwantes, leg.

F. W. Gees, S. K. Gees, coll. AMGS; 1, 10.-13.ii.1972,

Otjikoko-Sud 61, Omaruru, SE 2116 Ad, [collector un-

known], coll. SMWN; 1, 4.—5.iii.1972, Plateau 38,

Luderitz, SE 2616 Cb, [collector unknown], coll. SMWN;

14, 27.iv.1972, Rotegab 95, Keetmanshoop, SE 2718

Ad, [collector unknown], coll. SMWN; 19, 11.1v.1998,

Swakop River bed on road to Goanikontes [22.41°S

14.35°E], on white flowers of Psilocaulon salicornioides

(Pax) Schwantes, leg. F W. Gees, S. K. Gees, coll.

AMGS; 16, 11.iv.1998, Swakop River bed on road to

Goanikontes [22.41°S 14.35°E], visiting deep pink flow-

ers of Gelenia papulosa (Eckl. & Zeyh.) Sond., leg. F. W.

Gees, S. K. Gees, coll. AMGS; 20'¢", 11.iv.1998, Swakop

River bed on road to Goanikontes [22.41°S 14.35°E],

visiting white flowers of Psilocaulon salicornioides

(Pax) Schwantes, leg. F W. Gees, S. K. Gees, coll.

AMGS; 16, 11.iv.1998, Swakop River bed on road to

Goanikontes [22.41°S 14.35°E], visiting yellow flowers

of Zygophyllum simplex, leg. F. W. Gees, S. K. Gees, coll.

AMGS; 19, 16.111.1999, Swakopmund, Swakop River at

bridge [22.42°S 14°32°E], on white flowers of Zygophyl-

lum stapffii Schinz, leg. F. W. Gees, S. K. Gees, coll.

AMGS; 16, 13.iv.2002, Walfisbay, Swakopmundvia

Dune 7 [22.55°S 14.36°E], leg. F. W. Gees, S. K. Gees,

coll. AMGS; 19, 20.—-23.x11.1974, Windhoek, Wasser-

valle1 [22°55'S 16°22'E], [collector unknown], coll.

SMWN; 19, 68.11.2001, Gobabis District, Somerkoms

S21 227015908: 1975.0 22" Fi. leo PALI Kark-Spriges.

coll. SMWN; 19, 20.11.2014, Harda Province, 15 km s

Rehobot [23°28'S 17°07'E], 1400 m, leg. J. Halada, coll.

CULSP; 224, 16.11.1990, Omaheke Region, Gobabis,

40 km w Witvlei, leg. M. Schwarz, coll. PMHA; 19,

25 —27.111.2003, Rundu District, Mile 46 [18°18'39"S

19°15'29"E], leg. A. H. Kirk-Spriggs, coll. SMWN;

SENEGAL: 16, 27.vi.2004, 60 km s Velingara, sweep

netting, leg. M. Halada, coll. CULSP; SOUTH AFRICA:

13 299, 8-12.ii.1982, Sandveld Nature Reserve, Hoop-

stad, SE 2725 Da, leg. Entomology Department, coll.

BMSA; 1, xii.1956, [?] “Snoot Rivien’, leg. Martin,

coll. AMGS; 16, 3.-8.x.1989, Cape Province, 15 kmn of

©ZFMK

220 Jens-Hermann Stuke & David K. Clements

Nieuwoudtville on road to Loeriesfontein, leg. F. W.

Gees, S. K. Gees, coll. AMGS; 19, 4.1.2010, Cape Prov-

ince, 25 km n Jansenville [32°49'S 24°44'E], 600 m, leg.

J. Halada, coll. CULSP; 19, xii.1922, Cape Province,

Aliwal North, det. as P. brevistylatus by Krober 1938,

leg. R. E. Turner, coll. NHML [NHMUK010922102];

192, 3.xi1.1986, Cape Province, Bloutoring [33°28'40"S

20°19'1S5"E], leg. F. W. Gess, coll. AMGS; 19, 24.

x1.1960, Cape Province, Grahamstown, leg. E. McC

Callan, coll. AMGS; 19, 27.x1.1981, Cape Province,

Grahamstown, [?] ”Olakwater”, leg. F. W. Gess, coll.

AMGS; 19, 13.1.1986, Cape Province, Grahamstown,

Clifton, leg. R. W. Gees, coll. AMGS; 19, 12.—30.1.1970,

Cape Province, Grahamstown, Hilton [-33.310629

26.525595], Malaise trap, leg. F. W. Gess, coll. AMGS;

14, 3.iii.1978, Cape Province, Grahamstown, Hilton

[-33.310629 26.525595], on flowers of Melolobium can-

dicans in sandpit, leg. F. W. Gess, coll. AMGS; 1, 3.

11.1978, Cape Province, Grahamstown, Hilton

[-33.310629 26.525595], leg. S. K. Gess, coll. AMGS;

14, 2.xii.1989, Cape Province, Grahamstown, Hilton

[-33.310629 26.525595], leg. F. W. Gees, S. K. Gees,

coll. AMGS; 19, 22.x.1981, Cape Province, Graham-

stown, Lynton, leg. F. W. Gees, S. K. Gees, coll. AMGS;

14, 12.11.1967, Cape Province, Grahamstown, Strowahn,

leg. C. Jacot-Guillarmod, coll. AMGS; 10, 19-24.

i1.191[?], Cape Province, Grahamstown, Table Farm,

leg. F. W. Gess, coll. AMGS; 14 19, 2-13.iii.2008,

Cape Province, Grahamstown, Three Chimneys Farm

[33°18.542'S 26°29.846'E], leg. A. H. Kirk-Spriggs, coll.

AMGS; 16, 11.x.1994, Cape Province, Hamaqual and

Kamieskroon / Sors Sors, leg. F. W. Gees, S. K. Gees,

coll. AMGS; 134, 17-21. & 24.ix.1995, Cape Province,

Richtersveld National Park, Koeroegabvlakte [28.11°S

17.03°E], visiting deep pink flowers of Hermbstaetia

glauca (Wendl.) Reichb. ex Steud., leg. F. W. Gess, S. K.

Gess., R. W. Gess, coll. AMGS; 1, 15.ix.1996, Cape

Province, Richtersveld, Pachtviel [28.33°S 16.34°E],

leg. F. W. Gess, S. K. Gess., R. W. Gess, coll. AMGS; 16,

16.x1.1994, Cape Province, Tierberg [33.10°S 22.16°E],

visiting cream flowers of Asclepias buchenaviana Schinz,

leg. F. W. Gees, S. K. Gees, coll. AMGS; 14, 26.xi—S.

x1.1987, Cape Province, Tierberg, research station

[33°7'42"S 22°16'24"E], leg. F. W. Gess, S. K. Gess., R.

W. Gess, coll. AMGS; 19, 26.xi—25.xii.1987, Cape

Province, Tierberg, study site [33°10'S 22°16'24"E], on

flowers of Asclepis buchenaviana Schinz, leg. F. W.

Gess, S. K. Gess., R. W. Gess, coll. AMGS; 63'4, 26.

x1—5.xu1.1987, Cape Province, Tierberg, study site

[33°10'S 22°16'24"E], visiting flowers of Asclepias bu-

chenaviana Schinz, leg. F. W. Gess, S. K. Gess., R. W.

Gess, coll. AMGS; 19, 1.-9-x11.1988, Cape Province,

Vanwyksfontein, 8 km w of Norvalspont, leg. R. W.

Gees, coll. AMGS; 19, 12.1.2004, Eastern Cape Prov-

ince, Grahamstown, [33°16'00"S 26°28'60”"E], 629 m,

leg. C. Hepburn, coll. AMGS; 1, 10.x.1992, Eastern

Bonn zoological Bulletin 68 (2): 209-273

Cape Province, Grahamstown, Hoowisonspoort, 3326

BC, leg. T. Doubel, coll. AMGS; 1, 28.ii.2004, Eastern

Cape Province, Thomas Baines Nature Reserve

[33°23'02'S 26°29'01"E], leg. C. C. Robertson, coll.

AMGS; 19, no date, Eastern Cape Province, Willow-

more, leg. Brauns, coll. PJHS; 14, xii.1907, Eastern

Cape Province, Willowmore, det. as P. interrupta, leg. H.

Brauns, coll. MRAC; 19, no date, Eastern Cape Prov-

ince, Willowmore, det. as P. interrupta, leg. H. Brauns,

coll. MRAC; 1, 25.i1.1902, Eastern Cape Province, Wil-

lowmore, leg. Brauns, coll. PJHS; 192, 25.—-29.x.2010,

Free State Province, Brandfort, Florisbad Res. Stat.

[28°46.039'S 26°04.234'E], leg. A. H. Kirk-Spriggs, coll.

BMSA; 299, 8—12.11.1982, Free State Province, Hoop-

stad, Sandfeld Nature Reserve, SE 2725 Da, leg. Ento-

mology Department, coll. BMSA; 19, 13.x11.1995,

Mpumalanga Province, Blyderevierspoort NP [24°39'S

30°50'E], leg. F. Koch, coll. ZMHB; 1, 1.x.1997,

Northern Cape, 60 km e Springbok [29.28°S 18.26°E],

visiting cream flowers of Asclepias buchenaviana Schinz,

leg. F. W. Gees, S. K. Gees, coll. AMGS; 1, 10.x.2000,

Northern Cape, Pachtvlei, e Alexander Bay [28.33°S

16°34'E], visiting yellow flowers of Lebeckia, leg. F. W.

Gees, S. K. Gees, coll. AMGS; 1’, 20.xi.1999, Western

Cape Province, 7 km n of Avontuur [33°40'S 23°09'E],

sweep netting, leg. M. Hauser, coll. PMHA; 19, 11.

iv.2006, Western Cape Province, Stellenbosch [33°55'S

18°51'E], leg. E. Mostert, coll. AMGS; SUDAN: 16,

14.v1.1996, Blue Nile, Wad Medani, leg. G. G. M. Schul-

ten, coll. RMNH; 1, 1935, Sangha prés Bandiagara,

leg. M. Griaule, coll, MNHN:; TANZANIA: 19,

16.1.2007, Pwani Province, 15 km e Utete [08°03'S

38°53’'E], 75 m, leg. J. Halada, coll. CULSP; TOGO:

22 9, 7.1v.2008, Mono riverside s Ahassomé [7°11'14"N

01°28'52"E], 90 m, leg. A. Ssymank, coll. PASS; ZAM-

BIA: 19, 27.vu—13.viii.1910, Upper Luangwa River,

des. as syntype of P. decisa, leg. S.A. Neave, coll. NHML

[NHMUK010922107]; 54:4, 20.v.—29.vii.1988, Zambe-

si Valley [16°18'S 30°16'E], leg. J. Weyrich, coll. ZFMK;

ZIMBABWE: 14499, 18.-19.xii.2011, 60 km nnw Bu-

lawayo [19°41'S 28°21'E], 1200 m, leg. J. Halada, coll.

CULSP; 19, 14.x11.1919, Bulawayo, des. as syntype of

P. decisa, [collector unknown], coll. NHML

[NHMUK010922108]; UNKNOWN LOCATION: 1°,

24.1.1928, “Rersolution”, leg. A. Walton, coll. AMGS;

19, 1893, “Shitlesia”, leg., coll. AMGS; 1, no date,

“T287”, [collector unknown], coll. ZFMK; 1, no date,

“T426”, [collector unknown], coll. ZFMK.

Stuke (2015b) concluded that P antigua is an out-

standingly variable species and as a result several new

synonyms were recognised in the Palaearctic Region.

While reviewing the large amount of material available

it became evident that most of the commonly occurring

Afrotropical Physocephala specimens also belong to the

antiqua species-group. It cannot be excluded that factors

©ZFMK

New conopid records from the Afrotropical Region (Diptera) — Part 3: Physocephalini 221

er >

ra. ’

"ag

Figs 7-10. Physocephala antiqua (Wiedemann, 1830). 7. Epandrium, dorsal view (4, Namaqualand and Kamieskroon); 8. Hypan-

drium and phallus, lateral view (4, Namaqualand and Kamieskroon); 9. Wing, darker form, ventral view (Q, Clifton, Graham-

stown); 10. Wing, paler form, ventral view (2, Badenou s Mbingué). dp = distiphallus; ea = ejaculatory apodeme; har = hypandrial

arms; hba = hypandrial bars; hbr = hypandrial bridge; pa = phallapodeme; po = postgonite; poe = postgonite evagination; psh =

phallus sheath.

such as DNA-sequencing, newly discovered morpholog-

ical characters or the host ranges of larvae may even-

tually necessitate the splitting of P antiqua as defined

herewith but at the present time the consistent separation

of distinct species within this highly variable taxon is not

possible using the morphological characters which are

Bonn zoological Bulletin 68 (2): 209-273

currently available, or which have been used in the past.

We could not find any consistent differences in the male

postabdomen or female theca within the various morphs,

for example, and the Afrotropical material is highly vari-

able in respect of characters such as wing pattern and

microtrichiation, size and shape of the vertex, dusting

©ZFMK

DpH

and setulae on the pleura, body size, and colouration of

the legs, abdomen and thorax. Dark-winged specimens

with the basal cell almost completely covered with mi-

crotrichia (although never completely covered!) appear

remarkably distinct, but we also found intermediates.

Such specimens could potentially be incorrectly assigned

to the newly described P. guillarmodi (see below) al-

though the dark brown subcosta of P. antiqua (light or-

ange-brown or yellow in P. gui/larmodi) and the more or

less black scutellum (orange in P. guillarmodi) are good

characters for separating these two species. Physoceph-

ala antiqua specimens with a shortened radial-medial

crossvein appear very similar to species of the microve-

na species-group, but always lack a distinct shining in-

dention at the hind margin of the eye and have a differ-

ent male postabdomen. Specimens in which the pleural

dusting stripe is indistinct, such as in old specimens or

specimens which have been wetted, could very easily be

misidentified as belonging to the vittata species-group.

Physocephala maculipes has previously been de-

scribed as a very variable species (Camras 2001). We

cannot find any differences to distinguish Afrotropical

specimens previously reported as P. maculipes, nor the

examined holotype, from Palaearctic specimens which

have been identified as P. antiqua. Therefore, Physoceph-

ala |Conops| maculipes Bigot, 1887 is herewith placed

as a junior synonym of Physocephala [Conops| antiqua

Jens-Hermann Stuke & David K. Clements

Wiedemann, 1830 (syn. nov.). Consequently, all previ-

ous synonyms of P. maculipes are also now placed as

synonyms of P. antiqua (see Stuke 2017a).

Physocephala madagascariensis Krober, 1915 is the

only species of the genus reported from Madagascar to

date. This is also described as being a very variable spe-

cies (Camras 1962a) and in our view cannot be distin-

guished from P. antiqua, and therefore it is also herewith

placed as a junior synonym to Physocephala [Conops |

antiqua Wiedemann, 1830 (syn. nov.). This is the only

species known from Madagascar, so there should be no

doubt concerning the identification of specimens.

Krober (1939) had previously assumed that his P. minu-

tissima was no more than a small, dark morph of P. in-

terrupta Bezzi, 1901 (= P. antiqua). We examined the

male holotype, hitherto the only known specimen of this

species, which is in very bad condition with both wings

and abdomen lost. We also have specimens of small and

dark Afrotropical P. antigua which fit completely with

the original description of P. minutissima, as well as with

the remaining parts of the holotype. Physocephala minu-

tissima Krober, 1933 is therefore also herewith placed

as a junior synonym to Physocephala [Conops] antiqua

Wiedemann, 1830 (syn. nov.).

Physocephala gracilia Krober, 1915 has so far only

been known from two syntype specimens. Camras (2001)

considered that P. gracilia might be synonym of P. mac-

Fig. 11. Habitus of Physocephala brevipennis Camras, 1962 (holotype).

Bonn zoological Bulletin 68 (2): 209-273

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New conopid records from the Afrotropical Region (Diptera) — Part 3: Physocephalini 223

ulipes Bigot, 1887 (= P. antiqua) and in fact almost all

the characters given both in the original description and

in the additional information provided by Camras (2001)

fall within the variability of P. antiqua. We also have oth-

er specimens to hand which fit completely with the orig-

inal description. The only anomalous characters in the

original description are the short proboscis, completely

black claws and shining tergites 1-3, which have not

been found to date in material of P. antiqua. These char-

acters were not mentioned in the key of Krober (1915),

however, nor in the subsequent comments of Camras

(2001), and are not evident in any the material which we

have to hand. One syntype is probably destroyed (Stuke

2017a) whilst the second is held at the Museo Civico di

Storia Naturale “Giacomo Doria” but was not available

for examination. In the absence of better information we

therefore believe the best current solution for this cryp-

tic taxon is to place Physocephala gracilia Krober, 1915

as a junior synonym of Physocephala |Conops]| antiqua

Wiedemann, 1830 (syn. nov.).

P. brevipennis Camras, 1962

(Fig. 11)

Physocephala brevipennis Camras, 1962

Primary type material examined. 3 holotype of Phys-

ocephala brevipennis Camras, 1962: (1) “Ngutu, / Zulu-

land / II-21-1951 /A. L. Capener”; (2) “Physocephala /

brevipennis / Camras”; (3) “Holotype 4 / Physocephala /

brevipennis / Camras”; (4) “FMNHINS / 3130492 / Field

Museum / pinned”; coll. FMNH.

The first impression when looking at this strange speci-

men is that it is perhaps an aberrant example of Physo-

cephala antiqua with exceedingly shortened wings. Until

further material becomes available, however, we have re-

tained this taxon.

P. claripennis Becker, 1923

(Fig. 12)

Physocephala claripennis Becker 1923

Primary type material examined. 3 holotype of Physo-

cephala claripennis Becker, 1923: (1) “El Obeid / 13.-14.

IN”; (2) “Aegypt. Sudan / Ebner, 1914”; (3) “Ph. clarip-

enis [sic] / Beck. / det. Becker”; (4) “Holotype / Physo-

cephala / claripennis / Becker / det. Camras, 2000”; (5)

“J. Skevington / Specimen # / 45425”; coll. NMW.

This may only be an extremely pale specimen of P. an-

tiqua with completely hyaline wings, but until further

material becomes available we have retained this taxon.

P. guillarmodi spec. nov.

(Figs 13-24)

urn: lsid:zoobank. org: act:8F911A66-CF 42-44 60-83 7F-708FA 7S8BLE9C

Holotype @&. (1) “Mamathes / Basutoland / 28-XTI-1959 /

C. Jacot / Guillarmod”; (2) “? Physocephala brevistylata

/ Krober 2”; (3) Holotypus / Physocephala guillarmodi

/ spec. nov. 2 / 2018”. Holotype is deposited in AMGS.

The specimen is pinned and in perfect condition.

Paratypes. BURUNDI: 19, 18.xi.1949, Bururi Prov-

ince, Bururi, 1950 m, Orée dela forét, leg. F. Francois,

coll. ISNB; LESOTHO: 16, 26.xii.1946, Bokong, leg.

A. Jacot Guillarmod, coll. AMGS; 19, 28.x11.1947,

Leribe, Hensley's Dam, leg. A. Jacot-Guillarmod, coll.

AMGS; 19, 16.xi1.1950, Mamathes, leg. C. Jacot-Guil-

larmod, coll. AMGS; 1, 28.xii.1959, ditto, SOUTH

AFRICA: 16, 3.-7.x.1988, Cape Province, Clanwilliam

Fig. 12. Wing of Physocephala claripennis Becker, 1923 (holotype).

Bonn zoological Bulletin 68 (2): 209-273

©ZFMK

224 Jens-Hermann Stuke & David K. Clements

Dam [32°11'30"S 18°53'42"E], on flowers of Aspala-

thus desertorum Bol., leg. F. W. Gees, S. K. Gees, coll.

AMGS; 16, 5.xii.1980, Cape Province, Grahamstown,

Hilton, Senecio, leg. D. W. Gees, coll. AMGS; 19, 8—

13.x.1987, Cape Province, Klein Alexandershoek, Clan-

william District [32°20'20"S 18°46'E], leg. F. W. Gees,

S. K. Gees, coll. AMGS; 14 19, 28.ix.1985, ditto; 14,

4 -—8.x.1994, Cape Province, Koornplanskloof 10 km s

Citrusdaal [32°40'S 19°01'E], 200-270 m, leg. R. Dan-

ielsson, coll. MZLU.

Description of holotype (female)

Length 9.6 mm; Wing-length 7.6 mm; Head-height

2.9 mm.

Head. Antenna orange-brown (Fig. 13). Arista very

short stylus-like, with 2 aristomeres situated at tip of first

flagellomere (Fig. 16). Both aristomeres minute, length

of apical aristomere shorter than height. Scape about

twice as long as maximum width, apically and lateral-

ly with black setae. Pedicel about five times longer than

maximum width, covered with black setae. Pedicel lack-

Figs 13-16. Physocephala guillarmodi spec. nov. (holotype). 13. Antenna, lateral view; 14. Habitus, lateral view; 15. Frons, dorsal

view; 16. Arista, lateral view

Bonn zoological Bulletin 68 (2): 209-273

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New conopid records from the Afrotropical Region (Diptera) — Part 3: Physocephalini 225

ing any ridge at base, and expanded towards apex. First

flagellomere long and conical, about two times as long as

high, pointed, and ventrally with an indistinct membra-

nous area. Lunule between base of antennae and ptilinal

suture distinctly developed, shorter than width of scape.

Eyes brown, lacking ommatrichia, facets all of about the

same size. Posterior margin of eye lacking any shining

indentation. Gena height / eye height (measurements tak-

en from head in lateral view) = 0.3. No ocellar tubercle,

no ocelli and no ocellar triangle evident. Frons (Fig. 15)

yellow, somewhat broader than long, concave, slightly

projecting above eyes posteriorly and lacking any setu-

lae. Anterior margin of frons concave. No frontofacial

spot. Frons with indistinct dusting all over, subshining

laterally. Vertex as broad as frons, separated from latter

by a more or less distinct ridge. Vertex shining, and in

posterior half covered with black setulae. Vertex apical-

ly with an indistinct depression. Ridge of vertex lacking

longitudinal grooves. Face yellow, with brown facial

keel. Face lacks dusting except for some very narrow

silver dusting along eye margin. Gena yellow, lacking

setae. Distinct facial grooves reaching mouth edge. Dis-

tinct facial carina reaching from base of antennae to a

distinctly broadened and outstanding frontoclypeal tu-

bercle. Ptilinal suture extending well beneath antennal

bases on either side. Oral cavity tapers dorsally. Postcra-

nium not obviously invaginated, yellow to light brown.

Whole postcranium lightly dusted, with dusting adjacent

to posterior eye margin obviously denser. Occiput and

postgena covered with long black setulae. Postgena not

widened and not delimited from occiput. Bottom por-

tion of postcranium clearly delimited and less setulose.

Proboscis reddish-brown, labellum black. Frontoclypeal

membrane long, light orange-brown and barely delim-

ited from orange-brown clypeus. Palps absent. Labium

distinctly longer than head-length, distinctly thickened

basally, anterior section completely fused and tubular.

Labrum not visible in type specimen. Labellum short,

completely divided, hardly broader than adjacent haust-

ellum, and covered with very short setulae only.

Thorax mainly orange-brown, with meduiotergite,

katepisternum and meron mainly black. Scutum with

large medial black spot and small lateral black spots (Fig.

17). Thorax grey dusted all over. Pleura as in Fig. 18.

Distinct dusting stripe on pleura reaching from middle

coxa to notopleuron. The dusting stripe depends on the

viewing angle and is best seen in dorsal view — care is

required as it may be hard to see with the wrong view-

ing angle (Fig. 18). Anterior half of anepimeron shining

to subshining. Presternum distinct, narrow. Basisternum

broad, ventrally narrowed to a point, and with scattered

setulae. Proepisternum lacks setae or setulae. Notopleu-

ron with a few slightly stronger setae. Postalar callus

with a few strong short setae flanked laterally by several

curved setulae. Katepisternum with 13—15 setae postero-

dorsally and no setae ventrally. Metakatepisternum lacks

Bonn zoological Bulletin 68 (2): 209-273

setae. Anepimeron with 8—10 small setulae. Mediotergite

convex, with minute and hardly visible black setae. Sub-

scutellum inconspicuous. Scutum covered with scattered

small black setulae and with no outstanding setae. Wing

as Fig. 21: Fore-margin tinged brown, with brown mem-

brane between subcosta and media M. Cell r,,. hyaline

apically and discal-medial cell dm brown basally. Veins

brown, subcosta paler brown than radial veins. Wing

mostly covered with microtrichia, but anal lobe partly

lacking microtrichia. Radial-medial crossvein rm small

but complete (Fig. 20). Basal-medial-cubital crossvein

incomplete. Radius R, and R,,, terminate close together

in costa, well beyond end of subcosta. Radius R,,. with

shallow, even curve in distal section which is directed to-

wards fore-edge of wing. Cell r,,. pedunculate, with vein

44+5

R,,,+M, distinctly longer than radial-medial crossvein.

Basal cell lacks any thickening. Cubital cell cup elongat-

ed, distinctly longer than vein Al+CuA,, and pointed dis-

tally (i.e. cubitus CuA, and anal vein A, meet at an acute

angle). Vein A,+CuA, not reaching hind margin of wing.

Cubital veins CuA, and basal-medial-cubital crossvein

distinctly separated. Upper and lower calypters yellow-

ish-white to brown, margin of upper calypter with short

white setulae. Alula slightly shorter than broad, lacking

setulae on posterior margin. Venae spuriae pronounced

in cell r,,., cubital cell cup and indistinctly in cubital

cell cua,. Haltere light yellowish-white with brown base

and reddish knob. Knob of haltere with brown setulae.

Legs orange with fore coxa mainly black. Legs with in-

conspicuous silver dusting to shining, hind coxa densely

silver-dusted. Posterior surfaces of fore and middle tib-

iae with obvious silver-dusted fields distally. Legs with

short, adpressed black setulae. Base of fore and middle

femora lacking denser black setulae. Areas of dense

black to brown setulae anteroventrally at tip of fore tibia,

and ventrally and posteriorly at tip of hind tibia. Middle

femur lacking a distinct row of regularly arranged setu-

lae. Hind femur lacks outstanding setulae. No preapical

setae dorsally and no setae ventrally on tibiae. Femora

ventrally lacking rows of short black setae. Coxae with

several setulae but lacking outstanding setae. Hind femur

slightly thickened in basal half. All tibiae thickened in

apical half. Each metatarsus with 2 stronger setae ven-

trally at base. Pulvilli yellowish-white. Claws brown,

with narrow black tips. Empodium light brown, about as

long as pulvilli.

Abdomen orange to dark brown, with diffuse dark

markings dorsally on tergites 1-4. Abdomen with short

black setulae all over, those on tergite 2 very small. 9 ab-

domen completely silver-dusted, although the impression

of dusting depends on viewing angle. Denser dusted hind

margins of tergites are not obvious. Tergites 1-3 fused

but remain distinct from each other. Maximum width of

abdomen at segment 4. Length : maximum width of terg-

ite 2 = 2.0; length : maximum width of tergite 3 = 1.9.

Tergite 5 and sternite 5 not completely fused laterally.

©ZFMK

226 Jens-Hermann Stuke & David K. Clements

Figs 17-21. Physocephala guillarmodi spec. nov. (holotype). 17. Scutum, dorsolateral view; 18. Pleura, lateral view; 19. Theca,

lateral view; 20. Radial-medial crossvein, dorsal view; 21. Wing, dorsal view.

Figs 22-23. Postabdomen Physocephala guillarmodi spec. nov. (<, Hilton, Grahamstown). 22. Epandrium, dorsal view; 23. Post-

gonite evagination, ventral view. ep = epandrium; dp = distiphallus; poe = postgonite evagination, tep = tooth on posterior margin

of epandrium; ce = cercus.

Bonn zoological Bulletin 68 (2): 209-273 ©ZFMK

New conopid records from the Afrotropical Region (Diptera) — Part 3: Physocephalini 227

Fig. 24. Distribution of P. guillarmodi spec. nov.

Shape of theca as shown in Fig. 19. Anterior surface of

theca apically with a few long setulae. Posterior surface

of theca almost completely covered with close-set long

blunt spicules, arranged in about 12 horizontal lines.

Tergite 7 bends distinctly ventrally, with an indistinct

longitudinal gap, and with a minute protruding tooth at

middle of posterior margin.

Female characters added from one dissected paratype

Sternites 1—2 fused. Sternites 3-4 not protruding ven-

trally, posterior parts of sternites inconspicuous. Tergites

3 and 4 lack evaginations. Sternite 5 anteriorly lacking

any elongation. Sternite 6 almost completely covered

Bonn zoological Bulletin 68 (2): 209-273

with long blunt spicules which are mainly arranged in

close-set, broken horizontal lines. Sternite 7 longer than

broad, anteriorly pointed, posterior margin with two

patches of dense long black setae. Ventral sclerotisation

fused with tergite 8 and therefore connecting at sides.

Tooth on tergite 8 distinct, its base elongated anteriorly.

Sternite 8 bulging posteriorly, covered with strong long

black setae and with obvious long setulae on posterior

margin. Paired cerci distinct. Sack-like ventral protrusion

of vagina with distinct annular sclerotisation. Opposite

the annular sclerotisation the ventral protrusion is hardly

sclerotised and is covered with short, broad microtrichia

giving the ventral protrusion a roughened surface. No ob-

©ZFMK

228

vious sclerotisation at base of either the accessory glands

or the spermathecal ducts. Sack-like ventral protrusion

of vagina lacks any additional sclerotisation. 2 pairs of

spherical spermathecae, the spermathecal ducts fusing

shortly after leaving the spermathecae. Spermathecal

ducts are sclerotised at the spermathecae, this sclerotised

part of the duct being sinuous.

Description of male

3 abdomen strongly dusted from posterior half of tergite

3 to tip (in anterior view) and almost shining laterally on

tergites 1-3. No obvious dusting at hind margin of terg-

ites. Tergite 1 with obvious black setulae laterally on bul-

bous lateral projections. Tergite 2 distinctly elongated,

about 5 times as long as width at posterior margin. Terg-

ite 2 lacking lateral tufts of setulae. Tergite 3 obviously

longer than broad and widened posteriorly, anteriorly

about 2.8 times as wide as posteriorly. Sternites 1, 4 and 5

present, sternites 2 and 3 reduced. Tergite 5 and sternite 5

separate. Sternite 4 almost square, minute and not clear-

ly delimited from membrane, with about 4 minute setu-

lae. Sternite 5 apically with small field of thick setae and

several long black setulae laterally. Protandrium broader

than epandrium and therefore projecting over it. Sternite

8 delimited from protandrium. Line of minute black setu-

lae at border of protandrium and sternite 8. Lateral edges

of protandrium fused ventrally by a narrow sclerotised

strip, which is not medially broadened. Paired cerci dis-

tinct, completely sclerotised and covered with scattered

black setulae. Epandrium as shown in Fig. 22, not fused

behind cerci but with a slightly sclerotised connection

on both sides of the epandrium. Within this connection

there is an elongated sclerotisation. Posterior margin of

epandrium with small black setulae, and an obvious short

black tooth with a broad blackish base. Epandrium dis-

tinctly concave laterally. No hypoproct evident and no

remains of surstyli recognised. No obvious strong black

setae nor long black setulae which would mark the base

of a surstylus. Subepandrial plate not sclerotised nor cov-

ered with microtrichia, and therefore not distinct. Dorsal

hypandrial bridge developed. No hypandrial lobe evi-

dent. Hypandrial bars fused distally, hypandrium ending

in a hypandrial arm. Hypandrial membrane almost ab-

sent, lacking microtrichia. Phallus sheath fused dorsally,

lacking any evagination or setulae. Postgonite distinct.

Postgonite evagination not sclerotised, and not projecting

above distiphallus. Lateral side of postgonite evagination

with indistinct microtrichia, the medially directed surface

with broad and overlapping microtrichia (Fig. 23). No

plate at postgonite evagination. Ring sclerite between

phallapodeme and distiphallus indistinct. No epiphallus

recognised. Distiphallus shorter than epandrium, entirely

covered with microtrichia, some of which are black and

arranged in lines. Distiphallus dorsally with large light-

ly sclerotized plate (Fig. 23) but lacking evaginations.

Phallapodeme longer than hypandrium arm. Ejaculatory

Bonn zoological Bulletin 68 (2): 209-273

Jens-Hermann Stuke & David K. Clements

apodeme elongated, lacking distinct attachment to sperm

sac.

Variability

Wing length 6.3—-7.6 mm. Basal aristomere sometimes

black apically. Face, gena, frons and postcranium some-

times more or less black in melanistic specimens. Facial

keel sometimes yellow. Extent of black marking on tho-

rax varies somewhat, and scutum sometimes completely

orange. Proepisternum may lack setulae. Wing some-

times more extensively brown, with discal-medial cell

dm almost completely brown and cubital cell cual brown

at base. Alula and cubital cell cup may partly lack mi-

crotrichia. All coxae sometimes obviously blackish. Hind

femur may have an obscure blackish or brown ring.

Diagnosis

Physocephala guillarmodi belongs to the antiqua spe-

cies-group as defined in Key 1. It is easily overlooked

amongst the more common and variable P. antiqua, but

can be differentiated from the latter by the brown bas-

al cell which is completely covered with microtrichia

(Fig. 20). With this character P. guillarmodi can only be

confused within the antiqua species-group with dark-

winged specimens of P. antiqua which may atypically

have the basal cell almost (but never completely) cov-

ered with microtrichia, or with P. pusilla. Physocephala

guillarmodi can be separated from both of these species

by its light yellow to yellowish-brown subosta which is

obviously different from the darker radial veins (Fig. 21)

and by the characteristic light brown thorax with three

black spots (which may be fused) on the scutum (Fig.

17). At a first glance this species resembles P. schmideg-

geri Stuke, 2017 which occurs in the Arabian Peninsula

(Stuke, 2017b) but P. guillarmodi is distinguished by the

complete dusting band which starts immediately above

the middle coxa and reaches up to the notopleuron with-

out becoming narrower. In the event that P. brevipennis is

eventually demonstrated to be an aberrant and misshaped

antiqua specimen (see above) this will also be distin-

guished from P. guillarmodi by its dark brown subcosta

which does not differ in colour from the radial veins.

Etymology

This species is named in honour of Charles Fréderic Jacot

Guillarmod (1912-1979), who collected several speci-

mens of this new species and whose collecting activities

made a huge contribution to knowledge of the Diptera of

South Africa.

Distribution

Physocephala guillarmodi is widely distributed in the

Afrotropical Region (Fig. 24).

P. larvata (Speiser, 1911)

(Figs 25—29)

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New conopid records from the Afrotropical Region (Diptera) — Part 3: Physocephalini 229

Conops (Physocephala) larvatus Speiser 1911

= Physocephala similis Krober 1915 (syn. nov.)

Material. DEMOCRATIC REPUBLIC OF CONGO:

14, 30.xii.1952, Nord-Kivu, Kinshasa, Vitshumbi, s Lake

Edward, leg. J. Verbeke, coll. ISNB; TANZANIA: 16,

1911, Marienhof [=Murutunguru], Ukerewe, leg. Con-

rads, coll. SMTD; UGANDA: 16, 12. viii. 1958, Uganda,

det. as P. similis by Camras 2000, leg. J. Bowden, coll.

NHML [NHMUK010922104].

Physocephala similis was distinguished from P. /arva-

ta by Camras (2001) within his similis species-group,

but the character which Camras used to identify similis

(“Face yellow’) contradicts the original description of

Krober (1915). Krober compared P. similis with P. lar-

vata and stated that P. similis “Ist P. larvata zum ver-

wechseln ahnlich, hat aber ganz andere Fligelzeich-

nung” [Is confused with the similar P /arvata, but has

quite a different wing pattern] although this difference

is not obvious when comparing the original descriptions

of P. similis and P. larvata. Both descriptions fit to the

wing shown in Fig. 27 and there remains no other differ-

ence between these two species. The depository of the

holotype is unknown (Stuke 2017a) and therefore Phys-

ocephala similis Krober, 1915 is placed as a junior syn-

onym of Physocephala |Conops| larvatus Speiser, 1911

(syn. nov.).

P. ssymanki spec. nov.

(Figs 30-35)

urn: lsid:zoobank. org: act: 7476796 B-0C67-44D8-AA85-1313BC97B38C

Holotype <. (1) “98/99/79”; (2) “visiting white fis /

Brownanthus kuntzei /(Schinz) Thlenf. & Bittrich / Aizoe-

ceae: Mesembryanthema”; (3) “Namibia / NW of Cape

Cross /21.44S 13.59E/ 14.11.1999 / F. W. and S. K. Gees;

(4) Holotypus / Physocephala ssymanki / spec. nov. /

2018”. Holotype is deposited in AMGS. The specimen is

pinned and in perfect condition.

Description of holotype (male)

Length 11.7 mm; Wing-length 7.0 mm; Head-height

2.6 mm.

Head. Antenna black, tip of pedicel and base of first

flagellomere brown. Arista stylus-like, with 2 aristomeres

situated at tip of first flagellomere (Fig. 32). Both ari-

stomeres minute, length of apical aristomere shorter than

height. Scape about twice as long as maximum width,

apically with black setae. Pedicel about six times longer

than maximum width, mainly dorsally and apically cov-

ered with black setae. Pedicel lacking any ridge at base,

and expanded towards apex. First flagellomere long and

conical, about 2.5 times as long as high, pointed, ventral-

ly lacking a membranous area. Lunule between base of

antennae and ptilinal suture distinct, shorter than width

Bonn zoological Bulletin 68 (2): 209-273

of scape. Eye brown, lacking ommatrichia, with facets

all of about the same size. Posterior margin of eye lack-

ing any shining indentation. Gena height / eye height

(measurements taken from head in lateral view) = 0.2.

No ocellar tubercle, no ocelli and no ocellar triangle evi-

dent. Frons (Fig. 31) broader than long, slightly concave,

slightly projecting above eyes posteriorly, and lacking

setulae. Anterior margin concave. Frons yellow, with no

frontofacial spot and with indistinct dusting all over. Ver-

tex as broad as frons, separated from latter by a more or

less distinct ridge. Vertex shining, and in posterior half

covered with black setulae; apically with an indistinct

depression. Ridge of vertex with minute longitudinal

grooves. Face yellow, lacking dusting except for some

silver dusting along eye margin. Gena yellow, lacking

setae. Distinct facial grooves reaching mouth edge, and

facial carina reaching from base of antennae to a distinct-

ly broadened and outstanding frontoclypeal tubercle.

Ptilinal suture extending well beneath antennal bases on

either side. Oral cavity tapers dorsally. Postcranium not

obviously invaginated, black to dark brown. Whole post-

cranium slightly dusted, with dusting adjacent to posteri-

or margin of eye obviously denser. Occiput and postgena

covered with black setulae. Postgena not widened and

not delimited from occiput. Bottom portion of postcra-

nium not delimited and not setulose. Proboscis black to

reddish-brown, labellum black. Frontoclypeal membrane

long, light orange-brown and hardly delimited from or-

ange-brown clypeus. Palps absent. Labium longer than

head-length in lateral view, distinctly thickened basally,

and with anterior section completely fused into a tube.

Labrum not visible in the holotype. Labellum short, com-

pletely divided, hardly broader than adjacent haustellum,

and covered with very short setulae only.

Thorax mainly black, with pleura black to brown. Post-

pronotum orange-brown. Thorax grey-dusted all over,

with distinct dust-stripe reaching from middle coxa to

notopleuron and becoming wider dorsally. Anterior half

of anepisternum and posterior half of anepimeron shining

(Fig. 34). Presternum distinct, broad. Basisternum broad,

ventrally narrowed to a point, lacking setae or setulae.

Proepisternum lacking setae or setulae. Scutum covered

with small black setae. Notopleuron with several stronger

setae. Postalar callus with a few strong, short black setae

above several curved setulae. Katepisternum with 2 se-

tae posterodorsally and no setae ventrally. Metakatepis-

ternum lacks setae. Anepimeron with 12 long setulae

about as long as maximum width of tibiae. Mediotergite

convex, with minute and barely visible black setae. Sub-

scutellum inconspicuous. Scutum covered with scattered

small black setae, and with no outstanding setae. Wing as

Fig. 35: Fore margin of wing tinged brown, with brown

membrane between subcosta and radius R,,, .. Cell r,,, hy-

aline apically and cell r,,, brown basally. Veins brown,

subcosta paler brown than radial veins. Wing mostly

covered with microtrichia, but with basal cell, basal me-

©ZFMK

230 Jens-Hermann Stuke & David K. Clements

Figs 25-29. Physocephala larvata (Speiser, 1911). 25. Habitus, lateral view (<, Vitshumbi); 26. Frons, dorsal view (3', Marienhof);

27. Wing, dorsal view (<, Vitshumbi); 28. Head, ventrolateral view (<', Marienhof); 29. Epandrium, dorsal view (<, Marienhof).

dial cell, base of discal-medial cell and base of cubital

cell cup lacking microtrichia. Radial-medial crossvein

small but complete. Basal-medial-cubital crossvein also

complete. Radius R, and R,,, terminate close together in

costa, well beyond insertion point of subcosta in costa.

Bonn zoological Bulletin 68 (2): 209-273

Radius R,,,, with shallow and even curve in distal section

directed towards fore-edge of wing. Cell r,, .pedunculate,

with vein R,,.+M, well expressed and distinctly longer

than radial-medial crossvein. Basal cell lacks any thick-

ening. Cubital cell cup elongated, distinctly longer than

©ZFMK

New conopid records from the Afrotropical Region (Diptera) — Part 3: Physocephalini 231

Figs 30-35. Physocephala ssymanki spec. nov. (holotype). 30. Habitus, lateral view; 31. Frons, dorsal view; 32. Antenna, lateral

view; 33. Basal cell, dorsal view; 34. Anepisternum, dorsolateral view; 35. Wing, dorsal view.

vein A,+CuA.,, and pointed distally (i.e. cubitus CuA, and

anal vein A, meet at an acute angle). Vein A,+CuA, not

reaching hind margin of wing. Cubital veins CuA, and

crossvein distinctly separated. Upper and lower calypters

yellowish-white to brown, margin of upper calypter

Bonn zoological Bulletin 68 (2): 209-273

with white setulae. Alula almost square, about as long as

broad, lacking setulae on posterior margin. Venae spuriae

pronounced in cell r,,. and cubital cell cup. Haltere white,

with light brown base. Knob of haltere with brown set-

ulae. Legs orange, with hind femur and hind tibia black

©ZFMK

232

in apical half. Legs with inconspicuous silver dusting or

shining, hind coxa densely silver-dusted. Posterior sur-

faces of fore and middle tibiae with obvious silver-dusted

fields distally. Legs generally with short, adpressed black

setulae. Base of fore and middle femora basally with

denser black setulae. Areas of dense black to brown set-

ulae anteroventrally on tip of fore tibia and ventrally and

posteriorly at tip of hind tibia. Middle femur lacking any

distinct row of regularly arranged setulae. Hind femur

lacking outstanding setulae. No preapical setae dorsally,

and no setae ventrally on tibiae. Femora ventrally lacking

rows of short black setae. Coxae with several setae, but

lacking any outstandingly long ones. Hind femur slight-

ly thickened in basal half. All tibiae thickened in apical

half. Each metatarsus with 0-1 stronger seta ventrally at

base. Pulvilli yellowish-white. Claws brown, with broad

black tips. Empodium light brown, and about as long as

pulvilli.

Abdomen orange to dark brown, with diffuse paler

markings. Abdomen with short black setulae all over. ¢

abdomen strongly dusted from posterior half of tergite

3 to tip (in anterior view) and almost shining laterally

on tergites 1-3. No obvious dusting at hind margin of

tergites. Tergites 1-3 fused but remain distinct. Abdomen

of holotype not dissected. Tergite 1 with obvious black

setulae on bulbous lateral projections. Tergite 2 elongat-

ed, about four times as long as smallest width, lacking

obvious lateral tufts of setulae. Tergite 3 slightly widened

posteriorly, about twice as wide posteriorly than ante-

riorly. Tergite 5 and sternite 5 separate. Sternite 5 with

scattered long black setulae and denser small black setu-

lae forming an indistinct field. Protandrium broader than

epandrium and projecting over it. Sternite 8 distinctly de-

limited from protandrium. Indistinct line of small black

setulae at border between sternite 8 and protandrium.

Paired cerci distinct, completely sclerotised and covered

with scattered setulae. Epandrium not fused behind cerc1,

but with a slightly sclerotised connection at both sides of

epandrium.

Diagnosis

Physocephala ssymanki belongs to the P. antiqua spe-

cies-group and can easily be recognised from the anepi-

meron, which has several characteristically long setulae

which are about as long as the maximum width of the

tibiae. The dusting stripe on the pleura obviously con-

trasts with the shining anepisternum and anepimeron,

and becomes wider dorsally (Fig. 34). The hind tibia and

femur are black in the apical two-thirds, contrasting with

the remaining orange-brown legs (Fig. 30).

Etymology

This species is dedicated to Axel Ssymank (Bonn) who

generously loaned numerous Conopidae specimens for

this project.

Bonn zoological Bulletin 68 (2): 209-273

Jens-Hermann Stuke & David K. Clements

Distribution

To date P. ssymanki is only known from the locus typicus

on the coast of southern Namibia.

P. pusilla (Meigen, 1804)

(Fig. 36)

Conops pusilla Meigen 1804

Material. CENTRAL AFRICAN REPUBLIC: 1¢ 299,

4.xi1.2010, 60 km w Banibo [3°50'N 16°44’E], 630 m,

leg. J. Halada, coll. CULSP.

Afrotropical specimens of this otherwise Palaearctic spe-

cies have previously caused confusion, and for a long

time were suspected of belonging to a different species.

Somewhat surprisingly, however, their morphological

characters fall within the variability of P pusilla as re-

cently defined by Stuke (2016). Physocephala pusilla

had not previously been recorded from the Afrotropi-

cal Region and was assumed to be a Palaearctic species

(Stuke 2017a). Together with P. antiqua, P. vittata and

Conops elegans Meigen, 1804, this is the fourth example

of a very variable and mainly Palaearctic species with a

distribution which extends into the Afrotropical Region.

Physocephala bimarginipennis species-group

Only one distinct Afrotropical species is included in this

group, as was also recognised by Camras (2001). Phys-

ocephala bimarginipennis was previously treated in the

genus Archiphysocephala Krober 1939, now synony-

mised. Identification using the characters given in Key 1

is straightforward.

P. bimarginipennis Karsch, 1887

(Figs 37-41)

Physocephala bimarginipennis Karsch 1887

= Physocephala gigantea Krober, 1936

Primary type material examined. 2 holotype of Phys-

ocephala bimarginipennis Karsch, 1887: (1) “Pungo An-

dongo / A. v. Homayer”; (2) “11031”; (3) “Physocephala

/ bimarginipennis / n. sp.”; (4) “Holotypus”; coll. ZMHB.

3 syntype of Physocephala gigantea Kroéber, 1936: (1)

“Ph. Type / gigantea’; (2) “Musée du Congo / Elisabeth-

ville / XI-1927 / Dr. M. Bequaert “; (3) “Physocephala

/ gigantea / Krb. / det. Kréber 1935”; (4) “R. DET / V

/ 2992”: (5) “Typus”; (6) “RMCA ENT / 000012173”;

coll. MRAC.

3 syntype of Physocephala gigantea Kroéber, 1936: (1)

“Paratype / Ph. gigantea’; (2) “Musée du Congo / Kivu:

Musingiro / 8-[X-1927 / Ch. Seydel”; (3) “Musinglro

[si] / 8 Sept. 1927 / Ch. Seydel”; (4) “Physocephala /

gigantea Krb. / det. Krober 1935”; (5) “R. DET / W /

2992”; coll. MRAC.

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New conopid records from the Afrotropical Region (Diptera) — Part 3: Physocephalini 233

Fig. 36. Habitus of Physocephala pusilla (Meigen, 1804) (3, w Banibo).

Additional material. DEMOCRATIC REPUBLIC OF

CONGO: 19, 8.vii.1937, Bambesa, det. as P. gigantea

by Janssens 1954, leg. J. Vrydagh, coll. MRAC; 19,

vi.1932, Eala, det. as P bimarginipennis by Camras

1962, leg. A. Corbisier, coll. MRAC; 19, 5.111.1954,

North Kivu Province, Lac Vert, forét route Goma — Sake,

leg. J. Verbeke, coll. ISNB; ETHIOPIA: 19, 4.xii.2014,

Bonga Waldsaum, 1628 m [07°11.87'N 036°15.13'E],

Bothriocline schimperi, leg. H.-J. Fligel, coll. PHJF; 12,

vil.2019, Bonga, Kafa Development Association Guest-

house [7°14’59.14”"N 36°15712.26”E], leg. T. Kirschey,

coll. PJHS; KENYA: 19, 16.x1.1967, Nairobi, ex Xylo-

copa flavorufa found dead 23.1x.1967 Green Office Colo-

ny, [collector unknown], coll. NHML; 1°, 9—23.v.2006,

Coast Province, Shimba Hills National Park, near artifi-

cial Ponds [4.22752°S 39.43197°E], 335 m, mixed grass-

shrub-land, Malaise trap, leg. R. Copeland, coll. NMKE;

SOUTH AFRICA: 19, 1.—3.1v.2001, Mpumalanga Prov-

ince, Blyderevierspoort NP [24°39'S 30°50'E], leg. D. J.

Greathead, coll. ZMHB; TOGO: 1, ii.2017, Kloto, for-

est area [6°57°31.66”N 0°34°29.75”E], leg. G. Goergen,

coll. ITA; 19, x11.2016, ditto; 19, x.2001, ditto; UGAN-

DA: 16, 22.vii. 1966, Kwanda, ex 3 Xylocapa rufa, leg.

D. J. Greathead, coll. NHML; 19, 12.111.2012, Western

Uganda, n Fort Portal Kiyura, forest edge [N302540

E004731], leg. Oehlke, coll. PASS.

Physocephala caenoneura species-group

Species belonging to this group were previously included

together with other species in the Pseudophysocephala

nigritarsis species-group of Camras (2001). This caused

some confusion because these species may have both a

distinct vena spuria which is well separated from radius

Bonn zoological Bulletin 68 (2): 209-273

R,,,, and short apical abdominal segments, thus poten-

tially appearing to fall within his Pseudophysocephala

vitripennis and Pseudophysocephala platycephala spe-

cles-groups respectively.

Females of the caenoneura species-group are easi-

ly recognised by the normally developed, usually ad-

pressed theca and short tergites 5 and 6. From outside

the caenoneura group, only P. microvena and P. pseu-

domicrovena share this combination of characters but

are easily recognised as belonging to the Physocephala

microvena species-group by the lack of a vena spuria in

cell r,,, and typically also by the reduced or absent radi-

al-medial crossvein. Males of caenoneura group can be

more difficult to assign, however, the elongated basal ari-

stomere and less modified wing venation being the only

characters which reliably distinguish them from males of

the microvena species-group. All members of this group

are restricted to the Afrotropical Region.

Key 4 -— Identification of the Physocephala caenoneura

species-group

1. Basal cell completely hyaline and almost completely

lacking microtrichia, or with only very short

microtrichia which are hardly visible and much less

obvious than those on the remaining wing (e.g. Fig.

53); base of cell r,,, hyaline; hind femur at most with

an indistinct dark spot ......000..0cccccccecccssececesseeeeenees 2

— Basal cell at least with some dark colouration and

sometimes completely black or brown, and almost

completely covered with microtrichia similar to

those on the remainder of the wing (e.g. Fig. 59);

cell r,,, completely brownish infuscated (e.g. Fig.

59); hind femur may have a distinct blackish ring

©ZFMK

Jens-Hermann Stuke & David K. Clements

which is sharply demarcated from the orange-brown

COLGUEALIO cas pecs seats eas lee eI hee ee meee es 3

Tarsi orange-brown, not contrasting with tibiae;

apical aristomere typically less than half length of

projection on basal aristomere (Fig. 48); antennae

completely orange-brown (Fig. 47); gena usually

brownish, contrasting with yellow face (Fig. 47); °

theca short in side view, distinctly shorter than wide

Fea eC Re P. braunsi (Kroéber, 1931)

Tars1 dark brown, contrasting with brown tibiae

(Fig. 51); apical aristomere more than half as long

as projection on basal aristomere (Fig. 50); antennae

black to brown, with a contrasting orange-brown

Bonn zoological Bulletin 68 (2): 209-273

Figs 37-41. Physocephala bimarginipennis Karsch, 1887 (4, Kawanda). 37. Face, frontal view; 38. Frons, dorsal view; 39. Arista,

lateral view; 40. Tip of ¢ abdomen, lateral view; 41. Wing, dorsal view.

scape (sometimes faded in old specimens) (Fig. 50);

gena usually yellow, like the face; 9 theca long,

almost as long as wide (Fig. 54)......0.000cccceeeee

US, Fea i, ee ere P. caenoneura Krober, 1939

Projection of basal aristomere elongated, about

twice as long as apical aristomere (Fig. 58); radial

veins dark brown (Fig. 59); basal cell only slightly

darkened, obviously paler than cell r,,, (Fig. 59)......

Pe, eee 8 A ee P. capensis Krober, 1931

Projection of basal aristomere not as elongated,

not much longer than apical aristomere (e.g. Figs

44, 61, 65); radial veins sometimes light orange-

brown; basal cell mostly as dark as cell r,,, (e.g. Fig.

©ZFMK

New conopid records from the Afrotropical Region (Diptera) — Part 3: Physocephalini 235

45); base of cell r,,,and usually apex of basal cell

hyaline, and therefore radial-medial crossvein within

ajbyaline:patch (erie Fie" OO)". ) ci soo. moet 4

4. Characteristic dark mark at junction of discal medial-

cubital crossvein and media (Fig. 60); @ theca

obviously short, with about 7 interrupted rows of

blackssetde::hcareaait 4s P. fenestralis (Kréber, 1939)

— Wing colouration different, lacking dark mark at

junction of discal medial-cubital crossvein and

media, nor with hyaline spot surrounding radial-

medial crossvein; 9 theca longer, with more than 15

interrupted rows of black setae (e.g. Figs 43, 64)... 5

5. At least base of media and subcosta light yellowish-

brown; scutum orange-brown, with three elongated

blackish spots (Fig. 63); katepisternum orange-

brown; 2 with black setulae on theca less dense, not

obscuring underlying surface (Fig. 64) ..........0.0.000.

Pie oe oh oe ee P. rufitarsis (Camras, 1962)

— Veins uniformly dark brown; scutum mainly blackish,

with no distinct spots (Fig. 42); katepisternum mainly

black; ° with black setulae on theca denser, largely

obscuring underlying surface (Fig. 43) ........000.0..000...

Daca EMME ARe as LP ia P. afenestralis (Camras, 2001)

P. afenestralis (Camras, 2001)

(Figs 42-45)

Pseudophysocephala afenestralis Camras 2001

Material. SOUTH AFRICA: 19, 20.—24.x1.2014, Kwa-

Zulu-Natal Province, Royal Natal National Park, Gudu

Bush falls [28°40.925'S 28°55.778'E], 1628 m, Malaise

trap in indigenous afromontane forest, leg. A. H. Kirk-

Spriggs & E. Letsobe, coll. BMSA.

P. braunsi (Kréber, 1931)

(Figs 46-49)

Physocephala braunsi Krober 1931

Material. SOUTH AFRICA: 1, 15.iv.1948, Marieps

Mountain, det. as P. braunsi by Camras 2000, leg. G. van

Son., coll. FMNH; 16, xi.1933, Cape Province, Swel-

lendam, det. as P. platycephala by Krober 1938, det.

as P. braunsi by Camras 2000, leg. R. E. Turner, coll.

NHML [NHMUK010922151]; ZIMBABWE: 19,

7.xu1.1976, Salisbury [= Harare] [17.84 31.05], leg. P. E.

Hulley, coll. AMGS.

We are not convinced that the characters used to dis-

tinguish P. braunsi from the very similar P. caenoneu-

ra merit separate species status, but the few specimens

which are available can be segregated using the charac-

ters given in Key 4.

Bonn zoological Bulletin 68 (2): 209-273

P. caenoneura (Krober, 1939)

(Figs 50-54)

Pseudophysocephala caenoneura Krober 1939

= Pseudophysocephala stylata Krober 1939

Primary type material examined. 9 holotype of Pseu-

dophysocephala caenoneura Krober 1939: (1) “Type”;

(2) “Nyassa-See / Langenburg / 26. VII. —8. VIII. 98 / Ful-

leborn S.”; (3) “Typus”; (4) “Pseudophysocephala / styl-

ata Kb. / det. Krober 1938”; (5) “Zool. Mus. / Berlin’;

coll. ZMHB.

1 4 holotype of Pseudophysocephala stylata Krober

1939: (1) “Type”; (2) “Nyassa-See / Langenburg / 1.—26.

VII.98 / Fulleborn S.”; (3) “Typus”; (4) “Pseudophyso-

cephala / stylata Krb.”; (5) “Zool. Mus. / Berlin’; (6)

“Pseudophysocephala / caenoneura Krober / det. Cam-

ras 1999”: coll. ZMHB.

Additional material. KENYA: 19, 2.1i1.1993, II-

rad, w Nairobi, 1900 m, leg. B. Merz, coll. TAU; 18,

10.viit.2004, near Karura Forest, Nairobi [1°14.829’S

36°48.953’E], leg. R. Copeland, coll. NMKE; TAN-

ZANIA: 14 19, 4.ii.1977, Amani, East Usambara,

1000 m, det. as P. caenoneura by Camras 1999, leg.

H. Enghoff, O. Lomholdt, O. Martin, coll. FMNH; 13,

10.111.1963, Kilimajaru, Weruweru, det. as P. caenoneu-

ra by Camras 1999, leg. D. J. Greathead, coll. NHML

[NHMUK010922144]; 14, 20.iv.1966, plains e Arusha,

det. as P. caenoneura by Camras 1999, leg. D. J. Great-

head, coll. NHML [NHMUK010922145]6.

P. capensis Kroéber, 1931

(Figs 55-59)

Physocephala capensis Krober 1931

Material. SOUTH AFRICA: 1, 25.ii.2016, Barberton,

Mpumalanga [25°10'18"S 30°56'45"E], 1400 m, leg.

A. Ssymank, coll. PASS; 19, 4.-7.xi.1931, Cape Prov-

ince, George, det. as P. capensis by Krober, leg. H. P.

Thommaset, coll. NHML [NHMUK010922135]; 19,

23.1x.2007, Eastern Cape Province, Zuurberg [33°21'S

25°44'E], leg. C. Turnbull, coll. AMGS; 19, 15.x11.2012,

KwaZulu-Natal Province, Kwela Lodge, Pietermaritz-

burg [29°29'34"S 30°21'40"E], 930 m, leg. G. Stahls,

coll. PASS; 1, 15.x.2003, Western Cape Province, Hu-

mansdorp [34°1'60S 24°46'0E], 114 m, leg. C. Hepburn,

coll. AMGS.

P. fenestralis (Kriéber, 1939)

(Figs 60-62)

Pseudophysocephala fenestralis Krober 1939

Primary type material examined. 1 ¢ syntype of

Pseudophysocephala_ fenestralis Krober 1939: (1)

“Type”; (2) “Nyassa-See / Langenburg / 14.V.1999/

Fulleborn S.”; (3) “Typus”; (4) “Pseudophysocepha-

©ZFMK

236 Jens-Hermann Stuke & David K. Clements

Figs 42-45. Physocephala afenestralis (Camras, 2001) (2, Royal Natal National park). 42. Frons and scutum, anterodorsal view;

43. Theca, ventral view; 44. Arista, lateral view; 45. Wing, dorsal view.

la | fenestralis Krb. /det. Krober 1938”; coll. ZMHB. 19 syntype of Pseudophysocephala fenestralis Krober

1939: (1) “Syn- / type”; (2) “George, Cape Province, /

27.VI.-1.VII.1920.”; (3) “S. Africa / R. E. Turner. / 1920-

Bonn zoological Bulletin 68 (2): 209-273 ©ZFMK

New conopid records from the Afrotropical Region (Diptera) — Part 3: Physocephalini 237

AGS eer PSN asise

pe nine hee

Figs 46-49. Physocephala braunsi Krober, 2001 (4, Marieps Mnt.). 46. Habitus, lateral view, 47. Head, lateral view, 48. Ari-

stomeres, lateral view; 49. Wing, ventral view.

318.”; (4) “Pseudophysocephala / fenestralis, Krb. / ex-

amined & det. / O. Krober, 1938.”; (5) “Pseudophyso-

cephala / fenestralis / Krb”; coll. NHML.

Bonn zoological Bulletin 68 (2): 209-273

Additional material. TANZANIA: 19, 20.viii.1980,

Mt. Rungwe, sw, 1900 m, det. as P. fenestralis by Cam-

ras 2000, leg. M. Stoltze, N. Scharff, coll. FMNH; 19,

16.11.1952, Njombe, 6000-6500 ft, det. as P. fenes-

©ZFMK

238 Jens-Hermann Stuke & David K. Clements

Figs 50-54. Physocephala caenoneura (Krober, 1939) (2, Nairobi). 50. Antenna, lateral view; 51. Leg with tarsi, dorsal view; 52.

Frons, dorsal view; 53. Wing, dorsal view; 54. Theca, posterior view.

tralis by Camras 1999, leg. W. Peters, coll. NHML

[NHMUK010922142].

P. rufitarsis (Camras, 1962)

(Figs 63-65)

Pseudophysocephala rufitarsis Camras 1962

Primary type material examined. 9 holotype of Pseu-

dophysocephala rufitarsis Camras 1962: (1) “Holo- /

type”; (2) “S. Africa. / R. E. Turner / Brit. Mus. / 1933-

108.”; (3) “Holotype 9 / Pseudophysocephala / rufitar-

sis / Camras”; (4) “Pseudophysocephala / platycepha-

la Loew / examined & det. / O. Krober. 1938”; (5) “E.

Cape Prov. / Katberg. / 15—30.1.1933.”; (6) “BMNH(E)#

/ 249149”: coll. NHML

Additional material. SOUTH AFRICA: 1, 1.1953, Na-

tal, Hilton Road, det. as P. rufitarsis by Camras 2000, leg. P.

Graham, coll. NHML[NHMUKO010922159]; 14, xi.1930,

Cape Province, Somerset East, det. as P. platycephala by

Krober 1938, det. as P. rufitarsis by Camras 2000, leg.

R. E. Turner, coll. NHML [NHMUK010922162]; 224,

19.x.1994, Cape Province, Tsitsikama National Park,

Storms River Pass [33°59’'S 23°55'E], leg. R. Daniels-

son, coll. MZLU; 19, 1.-10.1932, Eastern Cape Prov-

ince, Katberg, det. as P. platycephala by Krober 1938;

Bonn zoological Bulletin 68 (2): 209-273

det. as P. rufitarsis by Camras, 2000, leg. R. E. Turner,

coll. NHML [NHMUK010922189]; 192, 110.1932,

Eastern Cape Province, Katberg, det. as P. platycephala

by Krober 1938; det. as P. rufitarsis by Camras, 2000,

leg. R. E. Turner, coll. NHML [NHMUK010922190];

14, 11-18.ii.1933, Eastern Cape Province, Katberg,

det. as P. platycephala by Krober, 1938, det. as P. ruf-

itarsis by Camras 2000, leg. R. E. Turner, coll. NHML

[NHMUKO010922158]; 14, 1.—10.ii.1933, Eastern Cape

Province, Katberg, det. as P. platycephala by Krober 1938,

det. as P. rufitarsis by Camras 2000, leg. R. E. Turner,

coll. NHML [NHMUK010922160]; 14, 1.—12.iii.1933,

Eastern Cape Province, Katberg, det. as P. platycephala

by Krober 1938, det. as P. rufitarsis by Camras 2000, leg.

R. E. Turner, coll. NHML [NHMUK010922161]; 19,

15.—30.1.1933, Eastern Cape Province, Ratberg, det. as

P. platycephala by Krober 1938, det. as P. rufitarsis by

Camras 2000, leg. R. E. Turner, coll. FMNH.

Physocephala halterata species-group

Camras (1962b) originally placed P. congoensis and

P. halterata in his Physocephala halterata species-group

and P. acroschista — very probably misidentified — in

his Pseudophysocephala vitripennis group. This spe-

cles-group is easily recognised by the velvety black

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New conopid records from the Afrotropical Region (Diptera) — Part 3: Physocephalini 239

Figs 55—59. Physocephala capensis Krober, 1931. 55. Theca, lateral view (2, Zuurberg); 56. Theca, posterior view (9, Zuurberg);

57. 2 abdomen, dorsal view (9, Zuurberg); 58. Arista, lateral view (2, Zuurberg); 59. Wing, dorsal view (2, Kwela lodge).

halteres. Members of the Physocephala halterata spe-

cies-group can be identified using Key 5 below. All mem-

bers of this species-group are restricted to the Afrotrop-

ical Region.

Key 5 — Identification of the Physocephala halterata

species-group

1. Frons orange-brown, lacking any distinct blackish

marking (Fig. 68); gena and facial groove orange to

Bonn zoological Bulletin 68 (2): 209-273

black (Fig. 67); at least postpronotum orange-brown;

femora orange-brown....P. halterata Brunetti, 1925

Frons with obvious blackish marking, or completely

black (e.g. Fig. 66); gena and facial groove more

or less black (e.g. Fig. 66); thorax including

postpronotum black to dark brown; femora black .. 2

Frons almost completely black (Fig. 66); gena brown

TOIDlaG kien: en soa8 alee P. congoensis Kroéber, 1936

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240 Jens-Hermann Stuke & David K. Clements

Figs 60-62. Physocephala fenestralis (Krober, 1939) (holotype). 60. Wing, dorsal view; 61. Arista, lateral view; 62. Wing, vena

spuria in cell r,, ., lateral view.

4+5?

— Frons black in basal half, and with a black medial

line reaching the lunule; gena yellow, with small

brown markings......... P. acroschista (Speiser, 1911)

P. acroschista (Speiser, 1911)

Conops (Physocephala) acroschistus Speiser 1911

Primary type material examined. ¢ holotype of Con-

ops acroschista Speiser, 1911: (1) “Holo- / TYPE”; (2)

“Musée du Congo / Kasai: Kondué / E. Luja”; (3) “R.

DET / B / 1477”; (4) “Type! / Conops (Physocephala)

/ acroschistus m. / P. Speiser det.”/; (5) “RMCA ENT /

000012172”; coll. MRAC.

The holotype reported here fits in all aspects to the origi-

nal description and there is no reason to assume that it is

not the type specimen, although according to the original

description this should be in the Museum d’ Histoire Nat-

uralle du Grand-Duchy de Luxembourg (MGDL). Cam-

ras (2001) reported concerning a specimen in the MGDL

as follows: “The unidentified holotype of acroschista

was finally found by J. M. Guinet and he informed me

that Luja was the collector.” The specimen reported by

Camras (2001) was a female, however, whereas in the

original description it was explicitly stated that the type

was a male, and a male is described. Evidently there are

more than one specimen bearing the same labels, which

Bonn zoological Bulletin 68 (2): 209-273

led to the misinterpretation of the female specimen in the

MGDL. Camras placed the species in his Pseudophys-

ocephala vitripennis species-group, having presumably

misinterpreted the species due to this wrongly labelled

specimen. The synonymy of P. intermedia Krober, 1936

and P. acroschista which was introduced by Camras

(2001) is therefore invalid and P. acroschista should be

treated as valid species (status rev.: valid species).

P. congoensis Kriber, 1936

(Fig. 66)

Physocephala halterata Brun. var. congoénsis Krober

1936

Primary type material examined. ¢ syntype of Phys-

ocephala halterata Brun. var. congoénsis Krober, 1936:

(1) “Ph. Type ¢ / halterata var / congoensis”; (2) “Musée

du Congo / Bambesa / 15-[X-1933 / H. J. Bredo”; (3)

“Physocephala ¢ / halterata Brun. / var. congoensis /

Krb / det. Kréber 1935”; (4) “R. DET” / P / 2992”; (5)

“Typus”; (6) “RMCA ENT / 000012174”; coll. MRAC.

9 syntype of Physocephala halterata Brun. var. con-

goénsis Kroéber, 1936: (1) “Ph. Type & / halterata var /

congoensis”’; (2) “Musée du Congo / Bambesa / 25-[X-

1933 / H. J. Bredo”; (3) “Physocephala ° / halterata

Brun. / v. congoensis Krb / det. Krober 1935”; (4) “R.

DET” / Q/ 2992”; (5) “Typus”; coll. MRAC.

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New conopid records from the Afrotropical Region (Diptera) — Part 3: Physocephalini 241

Figs 63-65. Physocephala rufitarsis (Camras, 1962) (2, Katberg). 63. Head and scutum, dorsal view; 64. Theca, posterior view;

65. Antenna, lateral view.

Additional material. BURUNDI: 13° 19, 15.1.1950,

Bururi, 1900 m, leg. F. J. Francois, coll. ISNB; 19,

26.v.1949, Bururi Province, Bururi, 1950 m, leg. F. J.

Francois, coll. ISNB; 13, 15.v.1949, Bururi Province,

Bururi, 2050 m, leg. F. J. Francois, coll. ISNB; DEM-

OCRATIC REPUBLIC OF CONGO: 19, 21-27.

vill.1931, Tshibinda [-0.287 28.776], det. as P. con-

goensis by Camras 2000, leg. J. Ogilvie, coll. NHML

[NHMUK010922105]; 14, 5.111949, North Kivu

Province, Gishari, Territory Masisi, 2300 m, leg. F. J.

Francois, coll. ISNB; EQUATORIAL GUINEA: 198,

15.-31.1.1907, Uelleburg, Benitogebiet, det. as P. congo-

ensis by Camras 2000, leg. G. Tessmann, coll. ZMHB.

Bonn zoological Bulletin 68 (2): 209-273

Physocephala congoensis is very similar to P. halterata

and P. acroschista, and all three species could be inter-

preted as colour morphs of a single species. As Camras

(2001) pointed out before, however, the few specimens

which are available can be consistently segregated using

the characters given in Key 5.

P. halterata Brunetti, 1925

(Figs 67—70)

Physocephala halterata Brunetti 1925

Primary type material examined. 9 holotype of Phys-

ocephala halterata Brunetti 1925: (1) “Holo- / type”; (2)

“S. Africa.”; (3) “Umbilo / 17.X.15”; (4) “Pres. by / Imp.

Bur. Ent. / Brit. Mus. / 1924-306.”; (5) “Durb / Mus”; (6)

©ZFMK

242 Jens-Hermann Stuke & David K. Clements

Fig. 66. Head of Physocephala congoensis Krober, 1936, anterolateral view (<, Bururi).

“Physocephala / halterata / Brun Type 2”; (7) BMN-

H(E)#/ 249078; coll. NHML

Additional Material. BURUNDI: 19, 23.i1v.1955,

Gitega [=Kitega], 1700 m, leg. F. J. Francois, coll.

ISNB; DEMOCRATIC REPUBLIC OF CONGO: 18,

6.111.1948, Katanga Province, Mukana, Lusinga, 1810 m,

leg. G. F. de Witte, coll. ISNB; 19, 15.11.1948, ditto;

ETHIOPIA: 19, 4.xi1.2014, Bonga Waldsaum, 1628 m

[07°11.87'N 036°15.13'E], Bothriocline schimperi, \eg.

H.-J. Flugel, coll. PHJF; KENYA: 19°, 25.1x.2005, Ker-

icho junction [0°23'S 35°16'E], 1530 m, leg. L. Fried-

man, coll. TAU; 14, 23.v.1910, Muguga, det. as P. hal-

terata by Camras 2000, [collector unknown], coll. NHML

[NHMUK010922109]; 19, vii.1937, Nairobi, det. as

P. halterata by Camras 2000, leg. V. G. L. van Someren,

coll. NHML [NHMUK010922121]; 1, vii.1937, Nairo-

bi, leg. van Someren, coll. NHML [NHMUK010922111];

LESOTHO: 16, 30.xii.1950, Mamathes [-29.136617

27.845796], leg. C. Jacot-Guillarmod, coll. AMGS; 19,

4.xi.1951, ditto; 234’, 9.xi.1951, ditto; 14, 11.xi.1951,

ditto; 229, 3.111952, ditto; 14, 15.11.1952, ditto; 19,

11.xi.1952, ditto; 24'3', 23.xi.1952, ditto; 14’, 14.xi.1954,

ditto; SOUTH AFRICA: 1, 1906, Transvaal, leg. A. J.

Cholmley, coll. NHML [NHMUK010922113]; 19, 6—

7.1.1993, Cape Province, Cape of Good Hope Nat. Res.,

leg. F. Koch, coll. ZMHB; 1, 12.xi.1952, Cape Province,

Bonn zoological Bulletin 68 (2): 209-273

Grahamstown, leg. C. Jacot-Guillarmod, coll. AMGS;

14, 23.i1.1970, Cape Province, Grahamstown, Belmont

Valley, on flowering Foeniculum vulgare Mill., leg. C.

Jacot-Guillarmod, coll. AMGS; 1’, i.1922, Cape Prov-

ince, Mossel Bay, det. as P. halterata by Krober 1938,

leg. R. E. Turner, coll. NHML [NHMUK010922114]:

12, 1v.1933, Cape Province, Mossel Bay, det. as hal-

terata by Krober 1938, leg. R. E. Turner, coll. NHML

[NHMUK010922117]; 19, 1—14.x1.1921, Cape Prov-

ince, Mossel Bay, det. as P. halterata by Krober 1938,

leg. R. E. Turner, coll. NHML [NHMUK010922120];

192, 22.11.1939, Cape Province, Mossel Bay, det. as

P. halterata by Camras 1962, leg. R. E. Turner, coll.

NHML [NHMUK010922124]; 14, 29.i1—5.ii.1924,

Eastern Cape Province, Port St. John, Bondoland, det.

as P. halterata by Krober 1938, leg. R. E. Turner, coll.

NHML [NHMUK010922115]; 19, 6.—25.11.1924, East-

ern Cape Province, Port St. John, Bondoland, det. as

P. halterata by Krober 1938, leg. R. E. Turner, coll.

NHML [NHMUKO010922119]; 19, 11.1927, Free State

Province, Harrismith, det. as P. halterata by Krober 1938,

leg. R. E. Turner, coll. NHML [NHMUK010922118]:

12, 10.x.2004, Gauteng Province, Benoni [26°08'S

28°23'E], leg. C. Midgley, coll. AMGS; 1, ii.1897, Na-

tal Province, Karkloof, [collector unknown], coll. NHML

[NHMUK010922110]; 19, 25—31.111.2001, Northern

Province, Lekgalameetse Reserve [24°12'S 30°20'E],

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New conopid records from the Afrotropical Region (Diptera) — Part 3: Physocephalini 243

Figs 67-70. Physocephala halterata Brunetti, 1925 (<4, Mamathes). 67. Habitus, lateral view; 68. Frons, dorsal view; 69. Haltere,

lateral view; 70. Wing, dorsal view.

Bonn zoological Bulletin 68 (2): 209-273 ©ZFMK

244

leg. F. Koch, coll. ZMHB; TANZANIA: 1, end of

xi1.1898 to end of 1 1899, Nyasasee, Langenburg [?=

Tukuyu], det. as P. congoensis by Camras 2000, leg. S.

Fulleborn, coll. ZMHB; 19, 24.x1.1963, Iringa Region,

Ulete, det. as halterata by Camras 2000, leg. D. J. Great-

head, coll. NHML [NHMUK010922122]; UGANDA:

14, 13.11.1966, Lake Nabugabo, leg. D. J. Greathead,

coll. NHML [NHMUK010922112]; ZIMBABWE: 16,

10.1926, Chirinda Forest, det. as P. halterata by Bry-

ant, leg. G. Arnold, coll. NHML [NHMUK010922116]:

12, 16—18.111.2001, Vumba Mountain, Botanical Gar-

den [19°07'S 32°47'E], leg. F. Koch, coll. ZMHB; UN-

KNOWN LOCATION: 19, 25.v.1901, [characters il-

legible], det. as P. halterata by Krober 1938, [collector

unknown], coll. NHML [NHMUK010922123].

Physocephala microvena species-group

This species-group combines some of the species which

were variously placed by Camras (2001) in his Pseudo-

physocephala_nigritarsis, Pseudophysocephala_platy-

cephala and Physocephala_ simplex species-groups.

Members of this group cannot be defined by single char-

acters but by a combination of characters as set out in

Key 1. A reduced or absent radial-medial crossvein, and

therefore either a reduced vena spuria or a vena spuria

that lies very close to the radius R,,, ., is typical, however.

Almost always there is no hyaline wing membrane be-

tween the vena spuria and radius R,, . and the aristomeres

are elongated, with the apical aristomere usually tower-

ing over the basal aristomere. The group contains some

very difficult species, several of which can readily be

confused with members from other species-groups, and

more material is necessary in order to understand the full

variability of the species. All members of this group are

restricted to the Afrotropical Region.

Key 6 — Identification of the Physocephala microvena

species-group

1. Face with deep black stripe extending from antennal

base to mouth edge, clearly separated from yellow

lateral part of facial groove (e.g. Fig. 92); hind femur

with a wide blackish ring in apical half, fore and

middle femora with distinct black spots (e.g. Fig.

91); dusting stripe on pleura not evident; vertex

almost entirely covered with long setulae and with

a roughened surface, although lacking longitudinal

grooves; margin of scutum and scutellum light

orange to brown (e.g. Fig. 91); cell r,,, completely

BROT (C50 MIMO SOS): es Eales alse teh laety, Aaa tt 2

— Face sometimes black but never with such a distinct

and deep black stripe; femora without such black

markings; other characters variable but never in the

ADOVCACOMDINATM ONE. cules Ay hawks neues Enea 3

Bonn zoological Bulletin 68 (2): 209-273

Jens-Hermann Stuke & David K. Clements

Discal-medial-cubital crossvein not completely

surrounded by brown colouration (Fig. 93); radial-

medial crossvein usually absent, radius and media

fused (Fig. 93); scape and pedicel light orange (Fig.

91); tarsi concolourous with tibiae, orange-brown

(Fig. 91); proboscis completely orange to brown,

lacking contrasting black labellum (Fig. 91); scutum

anteriorly with submedial pair of dull lines caused by

microtrichia; 4 tergites 3-5 densely dusted (anterior

view), tergite 2 densely dusted (posterior view) and

with a complete dusting band at its hind margin; 3

epandrium with isolated black tooth submedially on

posterior Margin... P. kroeberi nom. nov.

Discal-medial-cubital crossvein completely

surrounded by brown colouration; radial-medial

crossvein developed, radius and media therefore

separated; scape and pedicel mainly black to

dark brown; tarsi distinctly darker than tibiae;

proboscis black in apical half; scutum anteriorly

lacking submedial pair of dull lines caused by

microtrichia; 3 tergite 3 only lightly dusted, tergite

2 lacking dusting band at hind margin; no obvious

black tooth submedially on posterior margin of 3

SPAM riMI weer re: P. discalis (Camras, 1962)

3S narrow base of abdomen (from tergite 1 to end

of parallel-sided anterior part of tergite 3, Fig. 75)

only about half as long as the broad apical part of the

abdomen; 2 unknown....... P. brevipetiola (Camras,

1962)

Narrow base of abdomen (as measured above, e.g.

Figs 78, 94) more than half as long as the broad

apical part Of The ABC OMIM. rc9 eas Pasecreee Avena 4

S abdominal segments short, hind margin of tergite

3 almost as long as lateral margin (e.g. Fig. 78); ¢

tergites 4-5 densely dusted (anterior view); tergite 3

sometimes densely dusted (usually in posterior view)

and has a complete dusting band at its hind margin

(e.g. Fig. 78); 2 abdomen obviously narrows apically

in dorsal view (e.g. Fig. 76); Q theca small, not

projecting far ventrally (e.g. Fig. 77); 6 epandrium

with isolated black tooth submedially on posterior

margin (e.g. Fig. 82); epandrium in dorsal view as

e.g. Fig. 82; radial-medial crossvein sometimes short

but is always distinct; costa and subcosta usually

orange-brown; cell r,,, completely brown (e.g. Fig.

83); proboscis orange to brown, with contrasting

black labellum; legs including tarsi orange-brown;

face with more or less large brown to black mark;

gena typically light brown, darker than yellow

parafacia; frons with a more or less obvious brown

mark reaching from vertex to base of antennae;

scutum with three fused or separated blackish spots,

margin of scutum and scutellum always light orange

(e.g. Fig. 81); scutum anteriorly may have submedial

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New conopid records from the Afrotropical Region (Diptera) — Part 3: Physocephalini 245

dull lines caused by microtrichia (e.g. Fig. 81); where

such lines occur, the distance between them 1s almost

as wide as the lines themselves .............0...0cccccceee. 5

S abdominal segments usually longer, hind margin

of tergite 3 shorter than lateral margin (e. g. Figs

94, 106); 3 tergites 34 hardly to distinctly dusted,

tergite 2 at most indistinctly dusted and may lack

a complete dusting band at its hind margin (e. g.

Figs 94, 106); 2 abdomen not obviously narrowing

apically in dorsal view (e.g. Fig. 95); 9 theca always

larger, and usually projecting further ventrally (e.g.

Fig. 96); 4 epandrium lacking black tooth, or with

tooth situated on an elongated blackish keel on

posterior margin (e.g. Fig. 101); epandrium in dorsal

view sometimes different; radial-medial crossvein

sometimes completely absent (e.g. Fig. 101); costa

and subcosta sometimes dark brown like other veins;

cell r,,, may be hyaline in apical third; proboscis

sometimes completely brown to black, lacking a

distinctly darker labellum; legs and tarsi sometimes

completely black, or with obvious black markings;

face sometimes completely yellow to orange-

brown; gena sometimes yellow or orange-brown

like parafacia; frons may have no dark marking, or

is sometimes almost completely black to brown;

margins of scutum and scutellum sometimes black

or dark brown; scutum anteriorly with or without

submedial dull lines caused by microtrichia. Where

such lines occur these are only separated very

NAO WINACC ae OD )a. ro We ae ewe ical ee 6

Three separate blackish spots on scutum (Figs 103,

105); 9 theca about as broad as width of tergite 7

(Fig. 102), posterior surface with strong black setulae

less dense so that the underlying surface is visible,

ventral margin and posterior surface with long setae

irregularly arranged; 9 postabdomen elongated in

side view, apical tergites tapered in dorsal view (Figs

LOZ SOA cer re ee P. platycephala (Loew, 1853)

Black spots on scutum more or less fused (Fig.

81); 2 theca narrower than width of tergite 7 (Fig.

77), posterior surface with strong black setulae

densely packed so as to obscure underlying surface,

ventrolateral margin with long setae regularly

arranged in a line, about the same distance from each

other; 2 postabdomen not obviously elongated in

side view, apical tergites sharply narrowed in dorsal

view (Figs 76-77) .......... P. constricta Kriéber, 1915

Vertex usually with distinct longitudinal grooves

and always lacking distinct dense punctiform

indentations (cf Fig. 87); cell r,,, completely brown

(e.g. Fig. 107); radial-medial crossvein sometimes

completely absent, and radius and media fused (e.g.

Fig. 107); frons with a delimited black to light brown

marking from vertex to base of antennae, becoming

Bonn zoological Bulletin 68 (2): 209-273

narrower anteriorly; 2 theca short and broad, field of

black setae on sternite 4 about three times as broad as

long (e.g. Figs 96, 97); 9 tergite 6 obviously shorter

than tergite 4 in side view (e.g. Fig. 96)..........0....... o

Vertex usually lacking longitudinal grooves but with

distinct dense punctiform indentations (e.g. Fig. 87);

cell r,,, sometimes hyaline apically (e.g. Fig. 89);

radial-medial crossvein usually present (although

may be very short) and therefore media and radius

not touching (e.g. Fig. 89); frons sometimes almost

completely darkened; 2 theca longer and narrower,

field of black setae of sternite 4 at most about two

times as broad as long; 9 tergite 6 obviously longer

than tergite 4 in side view (e.g. Figs 73, 88)........... 8

Dusting stripe on pleura more or less distinct,

reaching from middle coxa to notopleuron; margin of

scutum and scutellum orange and distinctly delimited

from blackish central scutum; scutum anteriorly with

submedial dull lines caused by microtrichia (Fig.

99); abdomen partly orange-brown to light brown,

grading into dark brown or blackish areas (Figs 94,

95); tergites 3 and 4 with distinct dusted hind margins

or strongly dusted all over, tergite 5 and protandrium

(4) or tergite 6 (2) completely dusted (all in dorsal

view, Fig. 94, 95); 4 epandrium with distinct black

tooth standing on an elongated blackish keel on

POSLEMOR Manet GOTO 1) ee ren cee ec Ree

bee Me ARATE tn ne, P. microvena Brunetti, 1925

Dusting stripe on pleura absent or only poorly

developed; margin of scutum and_ scutellum

completely black or dark brown, and not obviously

delimited from black centre (Fig. 106); scutum

anteriorly lacking submedial dull lines caused by

microtrichia; abdomen uniformly blackish to dark

brown (Fig. 106); tergite 3 may have an indistinctly

dusted hind margin, abdomen otherwise shining

to only very lightly dusted; ¢ epandrium not

examined: *.., 202 P. pseudomicrovena Kriber, 1939

Smaller species, wing length less than 7 mm............

Ic Re TT Rado P. digitata (Speiser, mae

Larger species, wing length about 8 mm ................

Face yellow, without dark spot (Fig. 72); gena

sometimes completely brown, in all specimens

examined there 1s at least some brownish colouration;

pleura lightly dusted all over but without obvious

dense dusting; scutum anteriorly without submedial

dull lines caused by microtrichia; tergite 3 lacking

large silver dusting spots in posterior lateral corners;

discal-medial-cubital crossvein more or less brown

infuscated (Figs 71, 74); 2 theca exceptionally long

GHG BS Jars Fae ea a em P. abyssinica Kriber, 1915

Face with dark spot; gena sometimes completely

yellow, at most with some brown colouration but

©ZFMK

246 Jens-Hermann Stuke & David K. Clements

Figs 71-74. Physocephala abyssinica Kroéber, 1915.71. 9 habitus, lateral view (3, n Vumba); 72. 9 head, lateral view (¢, n Vum-

ba); 73. @ theca, lateral view (Q, Albertville); 74. Wing, dorsal view (3', n Vumba).

Bonn zoological Bulletin 68 (2): 209-273 ©ZFMK

New conopid records from the Afrotropical Region (Diptera) — Part 3: Physocephalini

never completely brown; katepisternum strongly

silver dusted in posterolateral corner; scutum

anteriorly with submedial dull lines caused by

microtrichia; tergite 3 with large silver dusted spots

in posterior lateral corners; discal-medial-cubital

crossvein not brown infuscated in the specimens

examined; 9 theca not exceptionally long................

Sige Adel ubhentire San anekseoun P. maculifacies Camras, 2001

P. abyssinica Krober, 1915

(Figs 71-74)

Physocephala abyssinica Krober 1915

= Physocephala fumivena Camras 2001 (Syn. nov.)

= Physocephala longitheca Camras 2001 (syn. nov.)

= Physocephala atronata Camras, 2001 (syn. nov.)

Primary type material examined. 9 holotype of Phys-

ocephala longitheca Camras, 2001: (1) “Angola (A37) /

5 mls. NE. Negola / 25.11.1972”; (2) “Southern / African

Exp. / B. M. 1972 -1”; (3) “Holotype ° / Physocephala /

longitheca / Camras”; coll. NHML.

S holotype of Physocephala atronata Camras, 2001:

(1) “W. Uganda / Kibale Forest / 12.x11.1971 —9.1.1972 /

R. L. Mason”; (2) “Holotype <4 / Pseudophysocephala |

atronata / Camras”’; coll. NHML.

Additional material. DEMOCRATIC REPUBLIC OF

CONGO: 299, 1.-20.1.1919, Albertville [= Kalemi],

780 m, det. as P. “? abyssinica” by Brunetti 1925, leg.

R. Mayné, coll. MRAC; 13, x.1932, Lulua, Kapan-

ga, det. as P. “? abyssinica Krb” by Krober 1935, leg.

F. G. Overlaet, coll. MRAC; MALAWI: 1, 4.x.1919,

Nyasaland, Cholo. [= Thyolo District], det. as P. “? abys-

sinica’ by Brunetti 1924, leg. R. C. Wood, coll. NHML

[NHMUK010922100]; UGANDA: 19, 21.1.1913,

West Mengo District, Entebbe, det. as P. “? abyssini-

ca’ by Brunetti 1924, leg. C. C. Gowdey, coll. NHML

[NHMUK010922101]; ZIMBABWE: 16, 2.1.1966, n

Vumba [= Bvumba] [-19.093432 32.740631], paratype

of P. fumivena, leg. D. Cookson, coll. FMNH.

Physocephala abyssinica is easily recognised in the fe-

male by the very long theca (Fig. 73) but is difficult to

identify in the male. Camras (2001) failed to recognize

that females of his newly described P. /ongitheca were in

fact conspecific with males previously identified as P. ab-

yssinica. Instead he described the males again as P. fumi-

vena — a species in which only males are known. The

only character given by Camras (2001) to distinguish

P. fumivena and P. longitheca is the wing colouration,

with a dark infuscated discal-medial-cubital crossvein

being present in P. fumivena. It is clear that this char-

acter is very variable, however, in which case there is

no other published character left to distinguish these two

species either from each other or from P. abysinica, and

we cannot find any such characters in the material we

Bonn zoological Bulletin 68 (2): 209-273

247

have to hand, which includes the holotype of P. /ongithe-

ca and a paratype of P. fumivena. Therefore, Physoceph-

ala longitheca Camras 2001 and Physocephala fumive-

na Camras, 2001 are both placed as junior synonyms of

Physocephala abyssinica Krober, 1915 (syns. nov.).

In addition, Camras (2001) also characterised P. atro-

nata with the character “Haltere with black on club”.

In fact there is only slight darkening on the haltere of

the holotype, which is quite different from the velvety

black haltere of species in the P. halterata species-group.

The holotype of P. atronata otherwise fits very well with

specimens of P. abyssinica and therefore Physocephala

atronata Camras, 2001 is also herewith placed as a ju-

nior synonym to Physocephala abyssinica Krober, 1915

(syn. nov.).

P. brevipetiola (Camras, 1962)

(Fig. 75)

Pseudophysocephala brevipetiola Camras, 1962

Primary type material examined. 3 holotype of Pseu-

dophysocephala brevipetiola Camras, 1962: (1) “< / Ho-

lotypus”; (2) “Musé du Congo / Eala / II-1932 / H. J.

Bredo”; (3) “Holotype 3 / Pseudophysocephala / brevi-

petiola / Camras”,; (4) “RMCA ENT / 000012177”; coll.

MRAC.

Additional Material. GABON: 1, viii.1892, Libre-

ville, determined as a Physocephala nov. spec. by Kréber

1921, [collector unknown], coll. MLUH.

Physocephala brevipetiola is a rarely recorded species.

Only the two males mentioned above are known to date.

These are obviously well characterised by the unique ab-

domen shape, having a very narrow base of the abdomen

(“petiole” sensu Camras (1962), cf Fig. 75) which is ob-

viously much shorter than remaining bulbous abdomen

(“abdominal club” sensu Camras 1962).

P. constricta Kroéber, 1915

Physocephala constricta Krober 1915

= Physocephala ruficoxa Krober, 1933

(Figs 76-83)

Primary type material examined. 9 lectotype of Phys-

ocephala constricta Kréber 1915 designated by Camras

(2001): (1) “3053”; (2) “Type”; (3) “Physocephala & /

constricta Krb. / O. Krober det. 1914”; coll. ZMHB.

S holotype of Physocephala ruficoxa Kréber 1933: (1)

“Holo - / type”; (2) “Lady Grey / 3 Feb. 1925. /R. I.

Nel”; (3) “Pres. By / Imp. Inst. Ent. / Brit. Mus. / 1932-

143”; (4) “Physocephala / ruficoxa Krob / det. Krober

1930”; (5) “Type”; (6) “Pseudophysocephala / constric-

ta Krober / compared to type / det. Camras, 2000”; coll.

NHML.

©ZFMK

248 Jens-Hermann Stuke & David K. Clements

eg RT en ie

f reart,

“a

+ .

. ete.

-. a «so

Fig. 75. Abdomen of Physocephala brevipetiola (Camras, 1962), dorsal view (<, Libreville).

Additional material. LESOTHO: 1, 13.xi.1949,

Mamathes [-29.136617 27.845796], leg. C. Jacot-Guil-

larmod, coll. AMGS; 16, 7.xi.1951, Mamathes

[-29.136617 27.845796], leg. A. Jacot-Guillarmod,

coll. AMGS; 2'3', 3.ii.1952, Mamathes [-29.136617

27.845796], leg. C. Jacot-Guillarmod, coll. AMGS; 12,

11.11.1952, ditto; 14, 23.xi.1952, ditto; 16’, 15.iii.1951,

Quthing, det. as P. spec., interrupta group by Smith 1966,

leg. Brink, Rudebeck, coll. MZLU; 16), 12.iii.1951, Se-

balabala, leg. C. Jacot-Guillarmod, coll. AMGS; 1¢,

8 —14.1.1963, Maseru district, Bushmans Pass, Maloti

Mountains, 2125—2250 m, det. as P. lineifrons by Cam-

ras 1963, leg. B. R. Stuckenberg, P. J. Stuckenberg,

coll. FMNH; SOUTH AFRICA: 16), “3053” [? Cape],

paralectotype of P. constricta, leg., coll. ZMHB; 19,

111.1959, Cathedral Peak, Forestry reserve, Natal Drak-

ensberg Little Berg summits, themeda Grassland, 5500—

6000 ft, det. as P. constricta by Camras 1962, leg. B. R.

Stuckenberg, P. J. Stuckenberg, coll. PJHS; 10, 10.-12.

x1.2009, Cape Province, Harrismith Scotland farm

[27°58'59.5"S 29°37'09.8"E], Malaise trap, dense Leu-

cosedea dominated scrub, leg. A. H. Kirk-Spriggs, coll.

BMSA; 1, 1.—10.ii.1933, Eastern Cape Province, Kat-

berg, det. as P. constricta by Krober 1938, det. as P. con-

stricta by Camras 2000, leg. R. E. Turner, coll. NHML

[NHMUK010922174]; 14, 1.-10.ii.1933, Eastern Cape

Province, Katberg, det. as P. constricta by Krober 1938,

det. as P. constricta by Camras 2000, leg. R. E. Turner,

coll. NHML [NHMUK010922175]; 14, 1.-10.ii.1933,

Eastern Cape Province, Katberg, det. as P. constricta

by Krober 1938, det. as P. constricta by Camras 2000,

leg. R. E. Turner, coll. NHML [NHMUK010922177]:

14, 1-10.i1.1933, Eastern Cape Province, Katberg,

det. as P. constricta by Krober 1938, det. as P. con-

stricta by Camras 2000, leg. R. E. Turner, coll. NHML

[NHMUK010922178]; 19, 19.—26.11.1933, Eastern Cape

Province, Katberg, det. as P. ruficoxa by Kroéber 1938;

det. as P. constricta by Camras 2000, leg. R. E. Turner,

Bonn zoological Bulletin 68 (2): 209-273

coll. NHML [NHMUK010922173]; 1, 6.i1.1925, East-

ern Cape Province, Lady Grey, paratype of P. ruficoxa,

leg. R. I. Nel, coll. NHML [NHMUK010922180]; 16,

26.1.1960, Eastern Cape Province, Molteno, leg. M. D.

Anderson, coll. AMGS; 1, 4.-5.ii.1992, Eastern Cape

Province, Rhode, village area [30°48'S 27°58'E], 1820 m,

det. as P. constricta by Camras 2000, leg. Natal Muse-

um Expedition, coll. FMNH; 14, 22.i.1965, Free State

Province, Senekal, leg. D. J. Brother, coll. AMGS; 1

19, 12.11.2016, KwaZulu-Natal Province, Garden Castle

N. R., Sleeping Bea [29°44'56'S 29°10'35"E], 2190 m,

leg. RSA-team Stahls, coll. PASS; 14, 4.xii.2012, Kwa-

Zulu-Natal Province, Royal Natal National Park, day

visitor Ca [28°41'25.4"S 28°56'53.9"E], 1410 m, Para-

type, leg. A. Ssymank, coll. PASS; 19, 20.—21.xi.2003,

Mpumalanga Province, 20 km sw Lydenbrug, leg. J.

Halada, coll. CULSP; 14, iii.1933, Natal Province, Na-

tal, Nationalpark, det. as P. constricta by Camras 2000,

leg. J. Ogilvie, coll. NHML [NHMUK010922179]; 16,

13.1v.1963, Northwest Reg., Retiefs Kloof, Rustenburg,

leg. H. N. Empey, coll. PMHA; UNKNOWN LOCA-

TION: 16,, “3054”, det. as P. platycephala by Krober

1914, wrongly labelled as allotype of P platycephala,

leg., coll. ZMHB; 29 9,, “AcP 5470”, leg., coll. NHML;

12, no date, “AcP 5470”, [collector unknown], coll.

NHML; 16, 13.xi.1954, [characters illegible], det. as

P. constricta by Camras 2000, leg. K. G. V. Smith, coll.

NHML [NHMUK010922176].

Females of P. constricta are easily identified by the

unique shape of the abdomen and theca as described in

Key 6 (Figs 76, 77). Males are extremely difficult to di-

agnose, however, and can be confused with members of

the abdominalis, microvena, caenoneura and antiqua

species-groups. Males of P. constricta should be com-

pared very carefully, bearing in mind that not all speci-

mens can be reliably assigned to species.

©ZFMK

New conopid records from the Afrotropical Region (Diptera) — Part 3: Physocephalini 249

Figs 76-83. Physocephala constricta Krober, 1915.76. 2 abdomen, dorsal view (9, Cathedral peak); 77. Tip of 2 abdomen, lateral

view (@, Cathedral peak); 78. 3 abdomen, dorsal view (3', Harrismith Scotland farm); 79. Arista, lateral view (3', Harrismith Scot-

land farm), 80. Indentation on posterior margin of eye, lateral view (2, Cathedral peak); 81. Scutum, dorsal view (3, Harrismith

Scotland farm); 82. Epandrium, dorsal view (<4, Mamathes); 83. Wing, ventral view (2, Cathedral peak).

P. digitata (Speiser, 1909) = Physocephala simplex Krober, 1915 (syn. nov.)

(Figs 84-90) = Physocephala ugandae Krober, 1915 (syn. nov.)

Conops (Physocephala) digitatus Speiser 1909 = Physocephala nigricoxa Brunetti, 1925

Bonn zoological Bulletin 68 (2): 209-273 ©ZFMK

250 Jens-Hermann Stuke & David K. Clements

= Physocephala bequaertorum Camras, 1962 (syn. nov.)

= Physocephala lineifrons Camras, 1962 (syn. nov.)

= Physocephala ethiopica Camras, 1962 (syn. nov.)

Primary type material examined. ¢ holotype of Pseu-

dophysocephala ugandae Krober 1939: (1) “?Psiloceph-

ala [sic] / ugandae, Krb. / examined & det. / O. Krober,

1938.”; (2) “Psilocephala [sic] / ugandae / Krb”; (3)

“Uganda, / Kampala, / 6.[1X.1918. / C C. Gowdey.”; (4)

“Kampala, / Uganda / 6.1X.1918 / No. 6545”; (5) “Pres.

by / Imp. Bur. Ent. / Brit. Mus. / 1921-153”; (6) “Type”:

coll. NHML.

S holotype of Physocephala nigricoxa Brunetti, 1925:

(1) “Syn- type”; (2) “Kampala, / Uganda / 4.1X.1918/C.

C. / No. 5392 Gowdey”; (3) “Pres. by / Imp. Bur. Ent. /

Brit. Mus. / 1924-306.”; (4) “Physo. / nigrocoxae [sic]

/ Brun Type @ / Det. E. Brunetti 1924”; coll. NHML.

There is a second damaged specimen lacking both head

and abdomen that is designated as “Syntype”. This can-

not be identified but in any event, in the original descrip-

tion the specimen listed above as the holotype is explicit-

ly identified as the “Type”.

9 holotype of Physocephala ethiopica Camras, 1962,

pinned together with <4 paratype: (1) “Holo- / type”;

(2) “Allo- / type”; (3) “Ethiopia: / Dilla, / (Sidamo). /

iv.1948.”; (5) “Holotype 2 / Physocephala / ethiopica /

Camras”; (6) “Allotype 3”; (7) “K. M. Guichard / B. M.

1948-200.”; (8) “[characters illegible]”; (9) “BMNH(E)#

/ 249079 / BMNH(E)# / 249080”; coll. NHML.

Additional material. BURUNDI: 19, V.1956, Mugera,

leg. F. J. Francois, coll. ISNB; CAMEROON: 19,

30.vil.1920, Lolodorf, det. as P. simplex by Camras 1962,

leg. A. I. Good, coll. FMNH; 14, 8.—-10.viii.2013, Far-

North Reg., Mayo-Tsanga, Mogode-Cha [10°34.111'N

13°36.422'E], 1001 m, Malaise trap, degrated savanna

forest, leg. A. H. Kirk-Spriggs, coll. BMSA; DEMO-

CRATIC REPUBLIC OF CONGO: 26'3', 15.ix.1933,

Bambesa, det. as P. ugandae by Krober 1935, leg. J. V.

Leroy, coll. MRAC; 19, 16.i11.1948, Basoko, Yamabuki,

Div. 153, det. as P. simplex by Camras 1962, leg. P. L.

G. Benoit, coll. MRAC; 10, 27.iv.1914, Beni [0°30'N

29°30'E], det. as P. bequaertorum by Camras 2000, leg.

J. Bequaert, coll. FMNH; 19, 23.1.1927, Elisabethville

[Lubumbashi] [-11.664232 27.482626], det. as P. ugan-

dae by Krober 1935, leg. M. Bequaert, coll. MRAC;

14, 4.iii.1936, Kamogobe, Sud Masisi, det. as P. nigri-

coxa by Camras 1962, leg. L. Lippens, coll. MRAC; 10,

18.vu.1935, Kivu, Nyongera, pres Rutshuru, 1218 m, leg.

G. F. de Witte, coll. ISNB; 1, 10.vii.1935, Kivu, Ritshu-

ru, riv. Musugereza, 1100 m, leg. G. F. de Witte, coll.

ISNB; 1, 10.vii.1935, Kivu, Ritshuru, riv. Musugere-

za, 1100 m, det. as P. nigrocoxalis by Vanschuytbroeck

F950. leg. “Gerk sde-Witte,, coll. MRACHID. /)5.x 1957

Massif Ruwenzori, Bomboka pres., Kyandolire, 1650 m,

leg. P. Vanschuytbroeck & J. Kekenbosch, coll. MRAC;

Bonn zoological Bulletin 68 (2): 209-273

192, 18.x11.1957, Massif Ruwenzori, riv. Lume, moy-

enne, 1800 m, leg. P. Vanschuytbroeck, coll. MRAC;

192, xi.1937, Rutshuru, det. as P digitata by Camras

2000, leg. J. Ghesquiere, coll. FMNH; 1, no date, Ville

Malela, Chief Casende [5,40S 23,45E], det. as P. simplex

by Camras, leg. J. Bequaert, coll. FMNH; RWANDA:

14, 6.i1.1935, Ruhengeri, riv. Mugara-Kigombe, 1800—

1825 m, det. as P. nigrocoxalis by Vanschuytbroeck 1950,

leg. G. F. de Witte, coll. MRAC; TOGO: 16, viii.2015,

Kloto, forest area [6°57°31.66”N 0°34’29.75”E], leg.

G. Goergen, coll. ITA; 19, 11.2002, ditto; 19, 11.2017,

ditto; 14', viii.2008, ditto; 14, vi.2016, ditto; UGAN-

DA: 16), 22.iv.1966, 20 miles e Mubende, det. as P. ni-

gricoxa by Camras 2000; det. as P. digitata by Camras

2001, at Chickweed flowers, leg. D. J. Greathead, coll.

NHML [NHMUKO010922194]; 14, 14.v.1958, Bugisu

Bugusege, det. as P. nigricoxa by Camras 2000; det. as

P. digitata by Camras 2001, at Chickweed flowers, leg.

J. Bowden, coll. NHML [NHMUK010922195]; 1,

1.—5.xi1.1911, Bugoma Forest, Unyoro, 3700 ft, det. as

P. nigricoxa by Camras 2000; det. as P. digitata by Cam-

ras 2001, at Chickweed flowers, leg. S. A. Neave, coll.

NHML [NHMUK010922192]; 14, 21.iv.1927, Dwoli,

det. as P. nigricoxa by Camras 2000; det. as P. digita-

ta by Camras 2001, leg. H. Hargreaves, coll. NHML

[NHMUK010922193]; 14, 15.-17.iii.1912, Uganda

West, Kibale Region, surrounding Bigodi [N0029209

E3020022], 1400 m, leg. Oehlke, coll. PASS; 14, 15—

17.iii.2012, ditto, UGANDA / KENYA: 16, v., Elgon,

1700 m, leg. Lindblom, coll. NHRS; ZAMBIA: 703

12, 12.-15.1.2003, 45 km se Kitwe, leg. J. Halada, coll.

CULSP.

Physocephala digitata is a species which shows great

variation in colouration of the face (with or without

dark marking); frons (more or less uniformly yellow to

light brown, the latter especially in old specimens, and

with distinct black midline or almost completely black);

coxae (completely light brown to almost black); scutum

(completely black to orange-brown with black centre

only); and wing (e.g. cell r,,, completely brown or hy-

aline apically, subcosta light orange to dark brown). In

all of these variations we find different intermediates and

no one of these characters, or combination of characters,

can be used to consistently segregate species. Camras

(2001) did not mention any character in his key for the

simplex species-group which is not variable in P. digita-

ta. Out of the eleven species he included in his simplex

species-group key, only three species can be recognised

consistently (see Key 6, couplets 8—9) and these three are

still difficult to identify convincingly.

The original description of P. simplex falls within the

variation of P. digitata as do the specimens reported by

Camras (2001) as P. simplex. The holotype of P. simplex

is stored in the Museo Civico di Storia Naturale “Giaco-

mo Doria” (Stuke 2017) and was not available for exam-

©ZFMK

New conopid records from the Afrotropical Region (Diptera) — Part 3: Physocephalini 251

Figs 84-90. Physocephala digitata (Speiser, 1909). 84. Frons, light form, dorsal view (2, Mugera); 85. Frons, dark form, dorsal

view (@, Kloto); 86. Epandrium, dorsal view (4, sw Kitwe); 87. Vertex, dorsal view (2, Mugera); 88. Theca, lateral view (9,

Mugera); 89. Wing, light form, dorsal view (2, Mugera); 90. Wing, dark form, dorsal view (°, Kloto).

Bonn zoological Bulletin 68 (2): 209-273 ©ZFMK

252 Jens-Hermann Stuke & David K. Clements

ination. Physocephala simplex Krober, 1915 is, however,

placed as a junior synonym of Physocephala [Conops|

digitata (Speiser, 1909) (syn. nov.).

Most specimens of P. ugandae are easily recognised

by the wing pattern (Fig. 89) with cell r,,, hyaline apical-

ly, and the frons yellow with a distinct black T-marking

(Fig. 84). There are, however, less commonly encoun-

tered forms typified by a completely dark cell r,,, and/or

either a completely dark or completely pale frons, togeth-

er with intermediates. Since this variation does not al-

low P. ugandae to be distinguished from P. digitata with

any consistency, Physocephala ugandae Krober, 1915 is

herewith placed as a junior synonym of Physocephala

[Conops] digitata (Speiser, 1909) (syn. nov.).

The original description and all of the key characters

used by Camras (1962b) to identify P bequaertorum fall

within the variation of P. digitata. One available headless

specimen in FMNH identified by Camras as P. bequaer-

torum also fits to P. digitata. Physocephala bequaerto-

rum Camras, 1962 is therefore placed as a junior syn-

onym of Physocephala |Conops| digitata (Speiser, 1909)

(Syn. nov.).

The type material of P. ethiopica falls within the varia-

tion of P. digitata. Physocephala ethiopica Camras, 1962

is therefore placed as a junior synonym of Physocephala

[Conops] digitata (Speiser, 1909) (syn. nov.).

The original description of P. /ineifrons compares it

with P. bequaertorum, stating that the two differ only

in the colouration of the gena and frons. The holotype

is deposited in the Natal Museum (KwaZulu-Natal, Pi-

etermaritzburg) and was not available for examination,

but the original description of P. Jineifrons falls within

the variation of P. digitata and therefore Physocephala

lineifrons Camras, 1962 is also placed as a junior syn-

onym of Physocephala |Conops| digitata (Speiser, 1909)

(Syn. nov.).

P. discalis (Camras, 1962)

Pseudophysocephala discalis Camras 1962

Material. UGANDA: 12, vii.1945, Bwamba Valley, det.

as P. discalis by Camras 2000, leg. van Someren, coll.

NHML [NHMUK010922150].

The interpretation of this species is based only upon the

above specimen in the NHML which was identified by

Camras as P. discalis following his comparison with the

holotype of that species at the USNM.

P. kroeberi nom. nov.

(Figs 91-93)

= annulipes (Krober 1939), junior secondary homonym

of Conops annulipes Wiedemann in Meigen 1824

Primary type material examined. ¢ holotype of Pseu-

dophysocephala annulipes Krober 1939: (1) “Holo- /

Bonn zoological Bulletin 68 (2): 209-273

type”; (2) “Kinangop / below bamboo / forest 8500 ft’;

(3) “Kenya: / Aberdare Range / x.1934. / B. M. E. Afr.

Exp. / B. M. 1935-203”; (4) “Pseudophysocephala / an-

nulipes. Krb / examined & det. / O. Kroéber 1938”; (5)

“Pseodophysoce | phala annuli- / pes Krb”; (6) [blank

red label]; coll. NHML.

Additional material. KENYA: 1, 22.ix.1990, Mount

Kenya, ~ 10.000 ft, sweep netting, leg. R. Copeland, coll.

IITA; 19, 1-11.1946, N. W. Mau Forest, 8000-1 0000ft,

det. as P. annulipes by Camras 1962, leg. H. P. Thomma-

set, coll. NHML [NHMUK010922136].

Physocephala kroeberi nom. nov. is a new name for

Pseudophysocephala annulipes Krober 1939 — Krober

(1939): 389; type-locality: “Kenya, Aberdare Range”;

HT ¢ [BMNH] - which is a junior secondary homonym

of Conops annulipes Wiedemann in Meigen 1824: 135;

type-locality: “Wahrscheinlich aus Oesterreich”; HT 3

[depository unknown, Evenhuis 1997] — available, in-

valid: junior synonym of Physocephala pusilla (Meigen,

1804).

Physocephala kroeberi is easily recognised by the dis-

tinctive black facial marking (Fig. 92), obvious markings

on the femora (Fig. 91), and wing pattern with the dis-

cal-medial-cubital crossvein not completely surrounded

by brown colouration (Fig. 93). It would be informative

to see more material to confirm that this is not merely

an extreme variation of P. platycephala or P. microvena,

however.

P. maculifacies Camras, 2001

Physocephala maculifacies Camras 2001

Primary type material examined. ¢ holotype of Physo-

cephala maculifacies Camras, 2001: (1) “Angola (A36) /

Chianga / 21 —24.111.1972”; (2) “Southern / African Exp.

/B.M. 1972 -1”; (3) “Holotype 3 / Physocephala / mac-

ulifacies / Camras”’; coll. NHML.

Additional material. ANGOLA: 19, 21.—24.111.1972,

Chianga, paratype of P maculifacies, leg. British Mu-

seum Expedition, coll. NHML [NHMUK010922096];

TANZANIA: 16, 10.iii.1963, Kilimajaru, Weruweru,

det. as P. abyssinica by Camras 2000, leg. D. J. Great-

head, coll. NHML [NHMUK010922099].

A difficult species for which more material is necessary

to test the stability of the identification characters given

in Key 6.

P. microvena Brunetti, 1925

(Figs 94-101)

Physocephala microvena Brunetti 1925

= Pseudophysocephala _ nigritarsis

(Syn. nov.)

Kroéber 1939

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New conopid records from the Afrotropical Region (Diptera) — Part 3: Physocephalini 253

Figs 91-93. Physocephala kroeberi nom. nov. (3, Mt. Kenya). 91. Habitus, lateral view, 92. Face, anterior view; 93. Wing, dorsal

view.

Primary type material examined. | 9 syntype of Phys-

ocephala microvena Brunetti 1925: (1) “Syn- / type”; (2)

“H. S. Stannus / Zomba. / Nyasaland.”; (3) “Pres. by /

Imp. Bur. Ent. / Brit. Mus. / 1924-306”; (4) “Physo. /

microvena / Brun Type & / Det. E. Brunetti 1924”; (5)

“BMNH(E)# / 249157”; coll. NHML.

14 syntype of Physocephala microvena Brunetti 1925:

(1) “Syn- / type”; (2) “H. S. Stannus / Zomba. / Nyasa-

land.”; (3) “Pres. by / Imp. Bur. Ent. / Brit. Mus. / 1924-

306”; (4) “Physo. / microvena / Brun / Type & / Det. E.

Brunetti 1924”; (5) “BMNH(E)# / 249156”; coll. NHML.

14 syntype of Physocephala microvena Brunetti 1925:

(1) “Syn- / type”; (2) “H. S. Stannus / Zomba. / Nyasa-

land.”; (3) “Pres. by / Imp. Bur. Ent. / Brit. Mus. / 1924-

Bonn zoological Bulletin 68 (2): 209-273

306”; (4) “Physo. / microvena / Brun /Cotype '/ Det. E.

Brunetti 1924”; (5) “BMNH(E)# / 249158”; coll. NHML.

1° syntype of Physocephala microvena Brunetti 1925:

(1) “Syn- / type”; (2) “H. S. Stannus / Zomba. / Nyas-

aland.”; (3) “Physo. / microvena / Brun / Cotype 3 /

Det. E. Brunetti 1924”; (4) “BMNH(E)# / 249158”; coll.

NHML.

1° syntype of Physocephala microvena Brunetti 1925:

(1) “Syn- / type”; (2) “H. S. Stannus / Zomba. / Nyasa-

land.”; (3) “Pres. by /Imp. Bur. Ent. / Brit. Mus. / 1924-

306”; (4) “Physocephala / microvena Brun. / 4. Cotype.

/ (Det. E. Brunetti 1924)”; (5) “BMNH(E)# / 249160”;

coll. NHML.

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254

9 holotype of Pseudophysocephala nigritarsis Krober

1939: (1) “Holo - / type”; (2) “Type”; (3) “Fort Portal /

Nyakasura / 24.1.1935 / J. W. Edwards”; (4) “Uganda: /

Ruwenzori Range / xii.1934-1.1935./B. M. E. Afr. Exp. /

B. M. 1935-203”; (5) “Pseudophysocephala / nigritarsis,

Krb. / examined & det. / O. Krober, 1938.”; (6) “Pseodo-

pDhysoce- | phala nigri- / tarsis Krb”; (7) “BMNH(E)# /

249147”; coll. NHML.

Additional material. BENIN: 19, 1.xu.2011, Lokoli,

swamp forest [07°03’40.00"N 02°15’50.00"E], leg. G.

Goergen, coll. IITA; BURUNDI: 299°, 512.111.1953,

Bururi, 1800-2000 m [-3.949 29.623], det. as P. nigri-

tarsis by Camras 1962, leg. P. Basilewsky, coll. FMNH,

MRAC; 19, 4.vi.1952, Colline Muramba, Bugoni, Terr.

de Muhinga [-2.930 30.357], 1400 m, det. as P. nigritar-

sis by Janssen 1955, leg. F. J. Francois, coll. ISNB; CAM-

EROON: 1, no date, Neu Kamerun, det. as P. nigri-

tarsis by Camras 2000, leg. G. Tessmann, coll. ZMHB;

DEMOCRATIC REPUBLIC OF CONGO: 19, no date,

Elisabethville [Lubumbashi] [-11.664232 27.482626],

det. as P. microvena by Camras 1962, leg. M. Bequaert,

coll. FMNH; 19, 10.xi.1928, Ituri, Blukwar, syntype of

P. curta but not conspecific with designated lectotype,

leg. A. Collart, coll. MRAC; 19, 12.iv.1953, Kivu, Kiten-

ga, pres riv. Ruzizi [-6.896 25.975], leg. J. Verbeke, coll.

ISNB; 19, 9.iv.1945, Rumangabo, riv. Bugombwa, det.

as P. gigantea by Janssens 1950, leg. G. F. de Witte, coll.

MRAC; 19, 29.11.1954, Secteur Tshiaberimu, riv. Kaliv-

ina, affl. Talia Nord, 2350 m [-0.13 29.42], 2350 m, leg.

P. Vanschuytbroeck & H. Synave, coll. MRAC; 14, 21-—

27.vii.1931, Tshibinda [-0.287 28.776], det. as P. nigri-

tarsis by Camras 1962, leg. J. Ogilvie, coll. FMNH;

KENYA: 1, 12.iii.1993, 5 km e Kaimosi, se Kakamega

[0.125 34.892], 1750 m, leg. B. Merz, coll. TAUI; 16,

10.x.1998, Bungoma [0.5645 34.558], leg. F. Kaplan,

A. Freidberg, coll. TAUL; 23'3', 12. -13.1.1996, Bungo-

ma [0.5645 34.558], leg. I. Yarom, A. Freidberg, coll.

TAUE; 19, 8.x.1998, Gilgil [-0.500043 36.326122], leg.

F. Kaplan, A. Freidberg, coll. TAUI; 34.4 19, 10.x.1998,

Kakamega [0.281 34.753], leg. F. Kaplan, A. Freidberg,

coll. TAUI; 14', 78.11.2011, Kakamega Forest, 1586 m

[0°13'37.2"N 34°52'49 8"E], 1586 m, leg. M. Mei, P.

Ceretti, D. Whitmore, coll. PMME; 14, 1—10.ii.2002,

Kakamega Isecheno Nature Reserve [0.24'N 34.86'E],

1800 m, leg. Okeka, coll. PMHA; 1, 11.-20.i.2003,

ditto; 14’, iv.1969, Karen nr Nairobi, leg. van Someren,

coll. NHML [NHMUK010922149]; 19, 9.x.1998, Keri-

cho [-0.368726 35.281219], leg. F. Kaplan, A. Freidberg,

coll. TAUI; 13), 11.—-12.iv.1957, Molo, Mau Escarpment,

2150-2200 m, det. as P. nigritarsis by Camras 1962, leg.

P. Basilewsky, N. Leleup, coll. MRAC; 1¢, ix.1939,

Naivasha, det. as P. nigritarsis by Camras 1999, leg. H.

J. A. Turner, coll. NHML [NHMUK010922138]; 19,

ix.1939, Naivasha, det. as P. nigritarsis by Camras 2000,

leg. H. J. A. Turner, coll. NHML [NHMUK010922141];

Bonn zoological Bulletin 68 (2): 209-273

Jens-Hermann Stuke & David K. Clements

14, 12.11.1993, near Kaimosi, se Kakamega [0.125606

34.844837], 1750 m, leg. Muhangani, coll. TAUI; 19,

16.x1.—22.x11.1998, Rift Valley Province, Mpala research

Station [00°19'N 36°53'E], Malaise trap, leg. S. Miller,

coll. NMKE; 1), 2.-9.i.2000, Western Province, Ka-

kamega Forest, Malaise trap [0°14.13'N 34°51.87'E], leg.

R. Copeland, coll. NUKE; UGANDA: 16, 22.iv.1966,

20 miles e Mubende, leg. D. J. Greathead, coll. NHML

[NHMUK010922147]; 13, 19.x.1958, Kawanda, det.

as P. nigritarsis by Camras 2000, [collector unknown],

coll NHML [NHMUK010922137]; 1 specimen,

x1.1934-1.1935, Ruwenzori Range, det. as P. nigritarsis

by Camras 2000, leg. B. M. E. Africa Expedition, coll.

NHML [NHMUK010922140]; 14, 20.iii.2012, Rwenzo-

ri Gebirge, Nyakalengila [N002100 E300149], 1705 m,

leg. Oehlke, coll. PASS; 1, 16.xi.1949, Shangugu

[= Cyangugu] [-2.483333 28.896667], 1460 m, det. as

P. platycephala by Janssen 1954, leg. F. J. Francois, coll.

ISNB; ZIMBABWE: 19, 2.—-6.xii.2015, Chirinda Forest,

Mt. Selinda [20°25'S 32°43'E], 1000 m, leg. J. Halada,

coll. CULSP; 14, 18.vi.1964, n Vumba [= Bvumba]

[-19.093432 32.740631], det. as P. microvena Camras

2000, leg. D. Cookson, coll. FMNH; 2007, 4.vii.1964,

n Vumba [= Bvumba] [-19.093432 32.740631], det. as

P. microvena Camras 2000, leg. D. Cookson, coll. FMNH;

192, 18.vii.1964, n Vumba [= Bvumba] [-19.093432

32.740631], det. as P. microvena by Camras 2000, leg. D.

Cookson, coll. FMNH; 19, 8.x.1964, n Vumba [= Bvum-

ba] [-19.093432 32.74063 1], det. as P. microvena by Cam-

ras 2000, leg. D. Cookson, coll. FMNH; 1, 20.11.1965,

n Vumba [= Bvumba] [-19.093432 32.740631], det.

as P. microvena Camras 2000, leg. D. Cookson, coll.

FMNH; 19, 8.v.1965, n Vumba [= Bvumba] [-19.093432

32.740631], det. as P. microvena by Camras 1999, leg. D.

Cookson, coll. FMNH; 19, 8.v.1965, n Vumba [= Bvum-

ba] [-19.093432 32.740631], det. as P microvena by

Camras 2000, leg. D. Cookson, coll. FMNH; 19, v.1932,

Xmas Pas, Umtali [= Christmas Pass near Mutare], leg. J.

Ogilvie, coll. NHML [NHMUK010922146]; 1, v.1932,

ditto [NHMUK010922148].

Most P. microvena specimens characteristically have

veins M and R,,. fused, thus lacking any radial-medial

crossvein (Figs 98, 100). A minority of specimens do

have a very short radial-medial crossvein present, how-

ever, and a reduced crossvein rm may also occur occa-

sionally in specimens of other species, causing complica-

tions in identification. Females of P. microvena are easily

identified by the short, broad theca (Fig. 97) and short

abdominal segments (Fig. 96), with confusion only really

likely with P. pseudomicrovena, which may be conspe-

cific. Conversely, males of P. microvena are extremely

difficult to identify reliably and could be easily confused

with males of other species-groups and also with males

of P. constricta where these also have a reduced radi-

al-medial crossvein. Where present, the distance between

©ZFMK

New conopid records from the Afrotropical Region (Diptera) — Part 3: Physocephalini 255

_ ,

= ata fe Si

Figs 94-101. Physocephala microvena Brunetti, 1925. 94. 3 abdomen, dorsal view (4, n Vumba); 95. 2 abdomen, dorsal view

(2, n Vumba); 96. 9 abdomen, lateral view (2, n Vumba); 97. 2 theca, ventral view (2, n Vumba); 98. Wing, dorsal view (9,

n Vumba); 99. Scutum with dusting stripes, anterodorsal view (<', n Vumba); 100. Completely reduced radial-medial crossvein,

dorsal view (2, n Vumba); 101. 3 epandrium, dorsal view (4, Bumgoma).

Bonn zoological Bulletin 68 (2): 209-273 ©ZFMK

256 Jens-Hermann Stuke & David K. Clements

Figs 102-105. Physocephala platycephala (Loew, 1853). 102: tip of 2 abdomen, lateral view (2, Worcester); 103. 2 scutum, dor-

sal view (2, Worcester); 104. Tip of 2 abdomen, dorsal view (2, Worcester); 105. 3 scutum, dorsal view (@, Clanwilliam dam).

the two short dark submedial stripes in the microtrichial

dusting on the anterior scutum may be the best character

for separating P. constricta and P. microvena, these being

separated by a wider medial dusted stripe in the former

(Fig. 81) than in the latter (Fig. 99).

As mentioned by Camras (2001) there are several inter-

mediates between typical P. microvena and P. nigritarsis.

Therefore, Pseudophysocephala nigritarsis Krober, 1939

is herewith placed as a junior synonym of Physocephala

microvena Brunetti, 1925 (syn. nov.).

P. platycephala (Loew, 1853)

Conops platycephalus Loew 1853

(Figs 102-105)

Material. SOUTH AFRICA: 16, 3-7.x.1988, Cape

Province, Clanwilliam Dam [32°11'30"S 18°53'42"E],

on flowers of Aspalathus desertorum Bol., leg. F. W.

Gees, S. K. Gees, coll. AMGS; 19, 1x.1928, Cape Prov-

ince, Worcester, det. as Physocephala spec. by Krober

1938, det. as P. platycephala by Camras 2000, leg. R. E.

Turner, coll. NHML [NHMUK010922130],.

Bonn zoological Bulletin 68 (2): 209-273

While the female of P. platycephala is easily recognised

by the obvious shape of the postabdomen and theca (Figs

102, 104), males are more problematic. We doubt that

males can be reliably identified using the colour pattern

of the scutum as proposed by Camras (2001) and as de-

scribed in Key 6. Figs 103 & 105 show that the black

spots on the scutum are less distinctly separated in males

(Fig. 105) compared to females (Fig. 103), and may per-

haps sometimes become fused. Perhaps males of this rare

species — only two female specimens are known — have

therefore previously been misidentified as other species.

P. pseudomicrovena Krober, 1939

(Figs 106-107)

Physocephala pseudomicrovena Krober 1939

Primary type material examined. ¢ lectotype of Phys-

ocephala pseudomicrovena Krober, 1939 designated

by Camras (1962): (1) “< / Holotypus”; (2) “Musée du

Congo / Tanganyka-Moero : / Nyunzu - I - II - 1934/ De

Saeger”; (3) “R. DET / O / 2992”; (4) “Physocephala 3

/ microvena Brun. / var. / det. Krober 1935”; (5) “RMCA

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New conopid records from the Afrotropical Region (Diptera) — Part 3: Physocephalini 257

Figs 106-107. Physocephala pseudomicrovena Krober, 1939. 106. Habitus, dorsolateral view (<4, Athiémé); 107. Wing, dorsal

view (4, Athiémé).

ENT / 000012181”; (7) “Type 3 / Physocephala / pseu-

domicrovena / Krober”’; (7) “designated by / Camras,

1962”; coll. MRAC.

Additional material. BENIN: 16, viii.2006, Athiémé

[06°14’20.00"N 01°40°00.00"E], leg. G. Goergen,

coll. IITA; 2929, viii.2006, Athiémé [06°14’20.00"N

01°40’00.00"E], sweep netting, leg. G. Goergen, coll.

IITA; 19, v.2006, Lokossa, sweep netting, leg. G. Goer-

gen, coll. IITA.

Bonn zoological Bulletin 68 (2): 209-273

The lectotype is in poor condition, with the antennae

missing and a broken abdomen almost completely cov-

ered with glue. All of the characters which can be seen fit

well to the concept of the species as given in Key 6, how-

ever. Neverthess, this species is only poorly differentiat-

ed from P. microvena, and it may easily be no more than

a dark morph of that species. On the other hand, all of

the specimens available to us can be distinguished with

the characters given in Key 6 and therefore the species 1s

here accepted as valid.

©ZFMK

258

Physocephala pilitarsis species-group

This group is identical to the Pseudophysocephala pili-

tarsis species-group of Camras (2001). Contrary to Cam-

ras (2001), however, it is not straightforwardly identified

because the main character he used — a large hyaline

area between the vena spuria and media in cell r,,.— also

occurs in some other Physocephala species-groups. Fe-

males belonging to this group are relatively easy to rec-

ognise by the theca, and the chaetotaxy and shape of the

tarsi (Key 1). Males are more difficult to identify, how-

ever, and only careful examination of several characters

in combination will rule out placement in other groups.

In addition, species identification of males is so far based

mainly on colouration characters, which may be variable.

There has not been enough material available to begin

looking at the male genitalia. All members of this spe-

cies-group are restricted to the Afrotropical Region.

Key 7 — Identification of the Physocephala pilitarsis

species-group

1. Reddish-brown species, scutum with three black

stripes, scutellum reddish-brown, abdomen mainly

reddish-brown; & fore tarsi wide and bare, lacking

any long setulae which are curled apically [based on

the original description only] ........00.0.0000cc eee

pean Ts eel N re eR Be ee P. rufa (Camras, 2001)

— Black to brown species, scutum mainly black,

lacking distinct stripes, scutellum black, abdomen

with at most tergites 1-3 orange-brown; @ fore tarsi

often with unusual characters such as curled setulae

2. Basal cell and basal medial cell dark and completely

covered with microtrichia (Fig. 118); radial-medial

crossvein sometimes quite short and the hyaline

area posterior to vena spuria in cell r,,, sometimes

indistinct (Fig. 118); 9 fore tarsus and middle tarsus

with long setulae which are curled apically (Figs 114,

116); 2 theca triangular and adpressed to abdomen

(Fig. 115); 2 sternite 6 with broad field of black

setae which is concave posteriorly (Fig. 115)...........

Shap TE ae ed P. pilitarsis Kréber, 1936

— Basal cell and basal medial cell hyaline, and at least

partly lacking microtrichia; radial-medial crossvein

sometimes quite short but the hyaline area posterior

to vena spuria in cell r, , is always distinct; 2 fore

tarsus may have long setulae but these are not curled

apically; 2 theca and @ sternite 6 different............ 3

3., [Pulvallrspmeslike (FIG 1 20) boos 2. ccter lead delice ls

Boh eee a Et ede P. spinipes (Camras, 2001)

— Pulvilli not spine-like, but normally developed ...... 4

4. Cells r, and r,,, completely dark brown, costal cell

never hyaline but sometimes paler than cell r, (Fig.

Bonn zoological Bulletin 68 (2): 209-273

Jens-Hermann Stuke & David K. Clements

113); 2 fore tarsus and middle tarsus not obvious

widened and lacking outstandingly long setulae (Fig.

109); 2 hind tibia with extremely short tarsomeres

(Figs 111, 112); 9 theca shown in Fig. 110..............

Dh cM, beer ne P. nitida (Kroéber, 1915)

— Cells r, and r,,, more or less hyaline, never both

completely tinged dark brown, costal cell sometimes

hyaline; 9 fore tarsus with or without long setulae; 9

hind tibia, hind tasomeres and theca different......... 5

5. Costal cell pigmented but slightly paler than cell

r; cells r,,, and r,,. apically hyaline, with base

contrasting darker; 9 fore and middle tarsi obvious

shorter than tibiae; fore tarsus with characteristic long

setulae: at apex with one pair of forward-directed

and strongly incurved setulae, and basally with two

pairs of strongly back-curved setulae; Q sternite 6

with a broad field of black, dispersed setae...............

sO ee te Rete ead P. tetratarsata (Camras, 2001)

— Wing with different markings; 2 fore and middle

tarsi different; 2 sternite 6 different... 6

6. Cell r, completely dark brown, cell r,,, mostly dark

brown and only slightly paler apically; abdomen

completely black; 9 theca with two patches of 6-7

strong black setae; sternite 6 with two distinct lateral

fields of dense black setae and on each sits a pair

of very close-set, long and strong black setae; 9

fore tarsus not obviously widened, and lacking long

setulae which equal the tarsus width........................

PR 44. ee te P. basilewskyi (Camras, 1962)

— Cell r, mainly hyaline, cell r,,, at most slightly

infuscated dark brown (Fig. 125); tergites 2-3 at

least partly orange-brown (Fig. 122); 9 theca and

sternite 6 lacking any fields of setae or long strong

setae, but with a large plate which has sparse fine

setulae (Fig. 124); 9 fore tarsus obviously widened

and with long black setulae which equal the tarsus

width (Fig. 123)............ P. vitripennis Curran, 1928

P. basilewskyi (Camras, 1962)

Pseudophysocephala basilewskyi Camras 1962

Primary type material examined. 2 holotype of Pseu-

dophysocephala basilewskyi Camras, 1962: (1) “9 / Ho-

lotypus”; (2) “Coll. Mus Congo / Kib.-Ituri : Irumu / 18/

XI-1952 / P. Basilewsky”; (3) “Holotype 2 / Pseudo-

physocephala / basilewskyi / Camras”; (4) “RMCA ENT

/ 000012176”; coll. MRAC.

P. nitida (Kroéber, 1915)

(Figs 108-113)

Conops nitidus Krober 1915

Primary type material examined. ¢ holotype of Con-

ops nitidus Krober 1915f: (1) “Togo / Bismarckburg / 2. —

©ZFMK

New conopid records from the Afrotropical Region (Diptera) — Part 3: Physocephalini 259

113

Figs 108-113. Physocephala nitida (Krober, 1915) (Q, Kloto forest). 108. 2 abdomen, dorsal view; 109. 2 fore tarsi, dorsal view;

110. & theca, ventral view; 111. & hind tibia and hind tarsi, lateral view; 112. 9 hind tarsi, dorsal view; 113. Wing, dorsal view.

18.vi.93 / L. Conradt S.”; (2) “Conops & / nitidus Krob/

O. Krober det. 1914”: (3) “Cotype”; (4) “Zool. Mus. Ber-

lin’; (5) “Pseudophysocephala / nitidus / Krober / det.

Camras, 2000”; coll. ZMHB.

Additional material. BENIN: 16', x.2008, Ahozon, for-

est area [06°22’57.59"N 02°9715.92"E], leg. G. Goergen,

coll. IITA; 19, 26.11.2013, Dangbo, sacred forest, leg.

G. Goergen, coll. IITA; KENYA: 10, 12.-26.ii.2005,

Nyanza Province, Ungoye Field Station [0°36.91'S

34°05.52'E], 1147 m, leg. R. Copeland, coll. NMKE;

Bonn zoological Bulletin 68 (2): 209-273

14, 27.ii—6.iii.2005, ditto; TANZANIA: 19, 1.vi-4.

vui.1893, Bismarckburg [= Kasanga], det. as P. nitida by

Kr6ober 1914 and Camras 2000, wrongly interpreted as

female syntype, leg. L. Conradt S., coll. ZMHB; TOGO:

14, 26.i1.2016, Dzobégan, monastery [7°14’22.07"N

0°41>58.64"E], leg. G. Goergen, coll. IITA; 1, ix.2006,

Kloto, forest area [6°57’°31.66"N 0°34)29.75"E], leg. G.

Goergen, coll. IITA; 14, viii.2008, ditto; 14, 1.2005, dit-

to; 12, vi.2008, ditto; 12, x.2016, ditto.

©ZFMK

260 Jens-Hermann Stuke & David K. Clements

Figs 114-118. Physocephala pilitarsis Krober, 1936 (2, Kakamega forest). 114. 2 fore tarsus, ventral view; 115. 2 postabdomen,

ventral view; 116. 2 middle tarsus, lateroventral view; 117. Radial-medial crossvein, dorsal view; 118. Wing, dorsal view.

P. pilitarsis Kroéber, 1936

(Figs 114-118)

Physocephala pilitarsis Krober 1936

= Pseudophysocephala ugandae Krober 1939, junior

homonym of Physocephala ugandae Krober, 1915

Primary type material examined. 2 holotype of Phys-

ocephala pilitarsis Krober, 1936: (1) “Ph. Type / pilitar-

sis’; (2) “Musée du Congo / Ubangi: Nzali / 3/4-H-1932/

H. J. Brédo”; (3) “R. DET / N / 2992”: (4) “Physoceph-

ala / pilitarsis Krb. / det. Krober 1935”; (5) “Typus”; (6)

“RMCA ENT / 000012180”; coll. MRAC.

S holotype of Pseudophysocephala ugandae Krober,

1939: (1) “Holo- / type”; (2) “Mpanga Forest / c 4,000

ft / F. W. Edwards”; (3) “Uganda: / Ruwenzori Range. /

Bonn zoological Bulletin 68 (2): 209-273

xil.1934-1.1935./B. M. E. Afr. Exp. / B: M. 1935-203.”;

(4) “Pseudophysocephala / ugandae Krob. / examined &

det. / O. Krober 1938.”; (5) “Pseudophyso- / cephala

ugandae / Krb”; (6) “Type”; (7) “Pseudophysocephala

/ pilitarsis / Krober / det. Camras, 2000”; coll. NHML.

Additional material. CENTRAL AFRICAN REPUB-

LIC: 14, 14.vi.2009, 150 km nww Mbaiki [04°05'N

17°02'E], 620 m, leg. J. Halada, coll. CULSP; 19,

2.v.2010, 20 km nne Mbaiki [03°04’'N 18°00'E], 450 m,

leg. J. Halada, coll. CULSP; 1¢, 9.xi.2012, 40 km e of

Bambio [3°60'N 17°12'E], 500 m, leg. J. Halada, coll.

CULSP; 1, 9.xii.2008, 50km ne Bambio [03°59'N

17°11'E], 450 m, leg. J. Halada, coll. CULSP; 34'3,

4.xi1.2010, 60 km w Banibo [3°50'N 16°44'E], 630 m,

©ZFMK

New conopid records from the Afrotropical Region (Diptera) — Part 3: Physocephalini 261

leg. J. Halada, coll. CULSP; 14, 24--28.xi.2010, 70 km

nne Bangui [04°57'N 18°46'E], 445 m, leg. J. Halada,

coll. CULSP; DEMOCRATIC REPUBLIC OF CONGO:

14, xii.1923, Katompe, Katanga, det. as “? abyssinica”

by Brunetti 1925, det. as P. pilitarsis by Camras 1962,

leg. Bequaert, coll. MRAC; 16, viii.1932, Lulua, Kapan-

ga, det. as P. “abyssinica Krb var.” by Krober 1935, leg.

F. G. Overlaet, coll. MRAC; 1, xi.1932, Lulua, Kapan-

ga, det. as P. “abyssinica Krb var.” by Krober 1935, leg.

F. G. Overlaet, coll. MRAC; 1, 1.iv.1937, Terr. Lisala,

det. as P. pilitarsis by Camras 1962, leg. Leontovich, coll.

MRAC; EQUATORIAL GUINEA: 19, 16.—31.viil.1906,

Alén, Benitogebiet, det. as P. pilitarsis by Camras 2000,

leg. G. Tessmann, coll. ZMHB; 19, 1.—15.x1.1906, Alén,

Benitogebiet, det. as P. pilitarsis by Camras 2000, leg. G.

Tessmann, coll. ZMHB; 224, vi—viii.1908, Uelleburg,

det. as P. pilitarsis by Camras 2000, leg. G. Tessmann,

coll. ZMHB; KENYA: 1, 13.-27.viii.2006, Western

Province, Kakamega Forest near Rondo Guest House

[0.22767°N 34.88533°E], 1630 m, Malaise trap set

across small permanent stream, leg. R. Copeland, coll.

NMKE; 19, 3.—-17.xii.2006, ditto; 19, 13.—27.viii.2006,

ditto; 14, 12—19.iii.2000, Western Province, Kakame-

ga Forest, Malaise trap [0°14.13'N 34°51.87'E], leg. R.

Copeland, coll. NMKE; 1¢, 28.viii—4.ix.1999, ditto;

14, 2—9.1.2000, ditto; 19, 20—-27.xi.1999, ditto; 299,

17.-24.vii1.1999, ditto; 12, 13.-20.11.2000, ditto; 299,

11.-18.1x.1999, ditto; 229, 3.—10.vii.1999, ditto; 19,

21.—28.v.2000, ditto; UGANDA: 1, vi.1912, Kawanda,

det. as P. pilitarsis by Camras 2000, leg. D. J. Greathead,

coll. NHML [NHMUK010922154]; 14, 2.viii.1962, Ma-

bira Forest, det. as P. pilitarsis by Camras 2000, leg. D. J.

Greathead, coll. NHML [NHMUK010922155].

P. rufa (Camras, 2001)

Pseudophysocephala rufa Camras 2001

No material of this species has been seen by us. Only the

female holotype is known, recorded from Malawi (Mu-

lanje Mountain, Nr. Likabula) at 1500m.

P. spinipes (Camras, 2001)

(Figs 119-121)

Pseudophysocephala spinipes Camras 2001

Primary type material examined. 9 holotype of Pseu-

dophysocephala spinipes Camras 2001: (1) “Tanganyika

/ Amani / 1957 / J. G. Halcrow / in thick bush”; (2) “C.

I. E. Coll. / NO 15485”; (3) “Pseudophysocephala / sp.

? / Det. R. L. Coe. 1958”; (4) “Pres. by / Com Inst Ent /

B M 1969-3”; (5) “Holotype 2 / Pseudophysocephala /

spinipes / Camras”; coll. NHML.

Additional material. KENYA: 19, 23.1x.2005, Ka-

kamega Forest, Yala River [0°13'N 34°53'E], 1450 m,

leg. A. Freidberg, coll. TAUI.

Bonn zoological Bulletin 68 (2): 209-273

P. tetratarsata (Camras, 2001)

Pseudophysocephala tetratarsata Camras 2001

Primary type material examined. 9 holotype of Pseu-

dophysocephala tetratarsata Camras 2001: (1) “Yaoundé

Cam. / Nkolbisson / Nonvll. X 1974”; (2) “parasitizing

6°63). Gla“ (4) ClIsE, Coll: A. 78117; (5) “Pres

by / Comm Inst Ent / B M 1975-1”; (6) “Holotype & /

Pseudophysocephala / tetratarsata / Camras”; (7) “Pseu-

dophysocephala / tetratarsata’ and on underside of this

label “Pseudophyso. / BM. Mus. From. / Yaounde Cam”;

coll. NHML.

Below the holotype is a large label with additional infor-

mation: “Following and ovipositing on bees which were

collecting sap flowing from Scolyid entrance holes in

felled trees. Each was following a bee flying immediate-

ly behind it. When a Physocephala managed to approach

closely a bee the fly deposited an egg on its body with an

extreme rapidity and I could establish later that the egg

remained attached to the tegument of the bee. Dr. Gui-

do Nonveiller in letter 6.4.76 to K. M. Harris. C. I. E.”

[Commonwealth Institute of Entomology].

P. vitripennis Curran, 1928

(Figs 122—125)

Physocephala vitripennis Curran 1928

= Physocephala intermedia Krober 1936 (syn. nov.)

= Conops bouvieri Séguy 1936 (Syn. nov.)

= Pseudophysocephala meii Camras 2001 (syn. nov.)

= Pseudophysocephala acroschista sensu Camras (2001),

nec Speiser, 1911

Primary type material examined. 3 holotype of Phys-

ocephala intermedia Krober, 1936: (1) “Ph. Type ¢

/ intermedia’, (2) “Musée du Congo / Ubangi: Nzali /

3/4-II-1932/ H. J. Brédo”; (3) “R. DET / Z / 2992”; (4)

“Physocephala 3 / intermedia Krb. / det. Krober 1935”;

(5) “Type”; (6) “RMCA ENT / 000012179”; coll. MRAC.

2 holotype, 3 paratype (pinned on one needle) of

Pseudophysocephala meii Camras 2001: (1) “Repub-

lique de Guinea PNHNS / 10°16’43”N 10°26’02”W /

Faranen, Sidakoro, mangor, / 29.XII.1995, leg. M. Mei”:

(2) “Holotype 9° and 3 / Pseudophysocephala | meii /

Camras”; (3) [glued male abdomen]; (4) “Pseudophyso-

cephala / cf vitripennis Curran / M. Mei det. 1997”; (5)

“Pseudophysoc / mei’; coll. FMNH.

Additional material. BENIN: 14, 20.iii.2011, Niaou-

li, forest area [06°44’3.08"N 02°8’2.59"E], leg. G. Go-

ergen, coll. IITA; 16’, 2.ii.2014, Niaouli, forest area

[06°44’°3.08"N 02°8’2.59"E], sweepnetting, leg. G.

Goergen, coll. IITA; 244, xi.2016, Sérou, forest area

[09°40’03.00"N 01°41°50.00"E], leg. G. Goergen, coll.

IITA; DEMOCRATIC REPUBLIC OF CONGO: 1 spec-

imen, 18.x.1945, Tshuapa, Flandria, det. as P. intermedia

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262

Jens-Hermann Stuke & David K. Clements

Figs 119-121. Physocephala spinipes (Camras, 2001) (2°, Kakamega forest). 119. 9 hind tibia and tarsi, lateral view; 120. Middle

tarsi, dorsal view; 121. Wing, dorsal view.

by Camras 1962, leg. P. Hulstaert, coll. MRAC; 24'3', no

date, Ville Malela, Chief Casende [5,40S 23,45E], det.

as P. acroschista by Camras 2000, det. as P. intermedia

by Camras 1962, leg. J. Bequaert, coll. FMNH; GHA-

NA: 16, 9.ix.1947, Kumasi, paratype of P. meii, leg. J.

Bowden, coll. NHML [NHMUK010922153]; NIGE-

RIA: 23'3, xii.1998, Ibadan, IITA, sweeping net in for-

est, leg. G. Goergen, coll. IITA; 19, 30.1.1988, Ibadan,

IITA, leg. G. G. M. Schulten, coll. RMNH; 19, 3.11.2000,

Ibadan, IITA, on mud at forest margin, leg. G. Goergen,

coll. IITA; SIERRA LEONE: 1¢ 19, 28.xi.1993, 5 km

w Kabala, swept along road [09°35'N 11°35'W], leg.

L. Cederholm, R. Danielsson & R. Hall, coll. MZLU;

UGANDA: 19, 16.1.1966, nr. Entebbe, paratype of

Bonn zoological Bulletin 68 (2): 209-273

P. meii, attacking Apis, leg. D. J. Greathead, coll. NHML

[NHMUK010922152].

Camras (1962b, 2001) was the only author to distinguish

four very similar species of the pilitarsis group: P. vit-

ripennis (Curran, 1928), P. intermedia (Krober, 1936) (as

P. acroschista sensu Camras (2001) nec Speiser, 1911),

P. bouvieri (Séguy, 1936) and P. meii Camras, 2001. He

used characters of wing colouration, relative length of

tarsomeres and extent of dusting on the postpronotum to

segregate these, but all of these characters are variable

and the four taxa intergrade. There is little consistent dif-

ference in the female theca or tarsi between these taxa.

The holotypes of P. intermedia and P. meii were avail-

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New conopid records from the Afrotropical Region (Diptera) — Part 3: Physocephalini 263

Figs 122-125. Physocephala vitripennis (Curran, 1928). 122. 9 abdomen, dorsal view (9, w Kabala); 123. 9 fore tarsi, dorsal view

(2, w Kabala); 124. 9 theca, ventral view (2, w Kabala); 125. Wing, dorsal view (9, Ibadan).

able to us but the holotype of P. bowvieri, which should

be in Muséum National d’Histoire Naturelle, Paris, but

which was not included in the list of material available

there (GBIF 2019), could not be investigated. These four

species are herewith treated as synonyms: Physocepha-

la vitripennis Curran, 1928 = Physocephala intermedia

Krober, 1936 (syn. nov.) = Physocephala [Conops]| bou-

vieri Séguy, 1936 (syn. nov.) = Physocephala |Pseudo-

physocephala| meii Camras, 2001 (syn. nov.).

Physocephala pubescens species-group

Our pubescens-group equates to the Pseudophysoceph-

ala pubescens species-group of Camras (2001). This

group is easily recognised by the distinct setae on the

mediotergite and several other subtle characters as sum-

marised in Key 1. Key 8 below, and the interpretation of

the species presented here, is based mainly on the key

and descriptions of Camras (2001). All members of this

group are restricted to the Afrotropical Region.

Bonn zoological Bulletin 68 (2): 209-273

Key 8 — Identification of the Physocephala pubescens

species-group

1. Frons with numerous short setulae (e.g. Fig. 129), or

least laterally with some setulae additional to those

on vertex; scutum and mediotergite with short setae

only, shorter than diameter of a tibia (e.g. Fig. 148).

— Frons lacking setulae, setulae only present on vertex;

scutum and mediotergite with long setae, longer than

diameter of a tibia (e.g. Fig. 141)... 5

2. Tibiae lacking or with only very indistinct rows of

black setulae (e.g. Fig. 127); setulae on frons and

face obvious and widely distributed, some of them

inside the facial grooves and on the facial ridges (e.g.

Pigsel Ze ply ese cee he ee ct eR at 3

— Tibiae dorsally with distinct rows of black setulae

(e.g. Fig. 140); no setulae in facial grooves or on

jC Teses i @ U0 [Sieh te Aaa Re See hf SOR ORER RE 4

3. Very long, fine and dense setulae all over; long

setulae on hind femur are especially obvious. Long

setulae all over frons (longer than the diameter of

hind tibia), and on abdomen.P hirta (Kroéber, 1939)

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264

— Vestiture of fine setulae much less long and dense.

Very few setulae anywhere longer than diameter of

hingktibig.ss! = 57.8 eam P. barbata (Camras, 2001)

4. Maximum length of apical aristomere about twice

maximum length of projection of basal aristomere

(Krober 1936: 272, Fig 132); setulae on frons

indistinct and confined to outermost lateral margins

of frons (Fig. 131); 2 theca semi-circular, with an

elongated field of dense black setae, the anterior and

posterior margins of which are almost parallel .........

ray ss CO rds P. caenostylata Kréber, 1936

— Maximum length of apical aristomere less than twice

maximum length of projection of basal aristomere

(Fig 149); setulae on frons more widely distributed,

usually not only confined to outermost lateral margins

of frons (Fig. 147); 9 theca narrow, triangular, with

rounded central field of dense black setae (Fig. 150)

Bocce dee Ricca ote Ate P. pubescens Brunetti, 1925

5. Tibiae lacking dorsal lines of black setulae; scutum

orange-brown, with three black stripes; 9 unknown

[based on the original description only]....................

Se Se yee ake sie OP One (0S Be P. alinea (Camras, 2001)

— Tibiae with distinct lines of black setulae (e.g. Fig.

140); scutum completely black centrally 0.0.0.0... 6

6. Basal aristomere projection shorter than apical

aristomere (Fig. 137); setulae on vertex denser and

therefore surface of vertex appears very roughened.

Anterior part of vertex with a distinct area bare of

setulae; setulae on vertex shorter, forming a level-

topped pile in lateral view, the longest setulae on

vertex being shorter than scape (Fig. 136); 2 theca

with crescentric apical field of long, blunt black

setae, the longitudinal height of the field being

distinctly greater than width (Fig. 139).......000000000..

ig dn PM der. Pe P. goergeni spec. nov.

— Basal aristomere projection about same length as

apical aristomere (Fig. 145); anterior part of vertex

which lacks setulae indistinct and much smaller,

and with a distinct depression (Fig. 143); setulae

on vertex less dense and therefore surface of vertex

appears less roughened (Fig. 143); setulae on vertex

longer, becoming taller at rear in lateral view, the

longest setulae on vertex about as long as scape; 9

theca with a much broader field of black setae (Fig.

LAG) stom Fe ees Pee P. nigrita (Camras, 1962)

P. alinea (Camras, 2001)

Pseudophysocephala alinea Camras 2001

No material of this species has been seen by us. Only

the male holotype is known, from Tanzania (Usambara

Mountains, Amani) at 1000m.

Bonn zoological Bulletin 68 (2): 209-273

Jens-Hermann Stuke & David K. Clements

P. barbata (Camras, 2001)

(Figs 126-130)

Pseudophysocephala barbata Camras 2001

Material. SOUTH AFRICA: 1, 10.-22.xii.1930,

Cape Province, Somerset East, paratype of P. barbata,

det. as P. constricta by Krober 1938, leg. R. E. Turn-

er, coll. NHML [NHMUK010922128]; ZIMBABWE:

243, 28.x.1965, n Vumba [= Bvumba] [-19.093432

32.740631], paratype of P. barbata, leg. D. Cookson,

coll. FMNH.

Physocephala barbata \ooks like P. hirta but with shorter

setulae. Only the female holotype of P. hirta is known

whilst only males are known of P. barbata, leading to the

suspicion that they may be conspecific. It would be use-

ful to find the opposite sex of one or other of these spe-

cies to exclude the possibility of sexual dimorphism such

as that observed in other species of the pubescens group.

P. caenostylata Kroéber, 1936

(Figs 131-133)

Physocephala caenostylatus Krober 1936

Primary type material examined. 2 holotype of Phys-

ocephala caenostylatus Krober, 1936: (1) “Ph. Type & /

caenostylus [sic]; (2) “Musée du Congo / Kibali-Ituri :

Kilo / V-1930/G. du Soleil”; (3) “R. DET /U/2992”; (4)

“Physocephala & / caenostylus [sic]/ Krb / det. Krober

1935”; (5) “Typus”; (6) “RMCA ENT / 000012178”;

coll. MRAC.

Additional material. KENYA: 19, 7.x.2013, S. Masai

Reserve, des. as syntype of P. pubescens, leg. T. J. An-

derson, coll. NHML [NHMUK010922163]; TOGO: 16,

vi.2008, Kloto, forest area [6°57’31.66"N 0°34)29.75"E],

leg. G. Goergen, coll. IITA.

Whilst identification of female P. caenostylata is straight-

forward males are very difficult. We are not convinced

that all specimens can be safely distinguished from those

of P. pubescens.

P. goergeni spec. nov.

(Figs 134-142)

urn. lsid:zoobank.org:act:A4F67D5A-00A2-45 1D-8B10-59F 1CA272ECO0

Holotype &. (1) “Togo / Kloto /forest area / Feb. 2018 /

Col.: G. Goergen”; (2) “Holotypus / Physocephala goer-

geni / spec. nov. & / det. Stuke 2019”. Holotype is depos-

ited in ZMHB. The specimen 1s pinned and in very good

condition.

Description of holotype (female)

Length 10.8 mm; Wing-length 7.0 mm; Head-height

2.8 mm.

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New conopid records from the Afrotropical Region (Diptera) — Part 3: Physocephalini 265

Figs 126-130. Physocephala barbata (Camras, 2001) (3', paratype n Vumba). 126. Habitus, lateral view; 127. Middle tibia, dorsal

view; 128. Face, anterolateral view; 129. Frons, dorsal view; 130. Arista, lateral view.

266

Jens-Hermann Stuke & David K. Clements

Figs 131-133. Physocephala caenostylata Krober, 1936 (3, Kloto). 131. Frons, dorsal view; 132. Arista, lateral view; 133. Middle

tibia, dorsal view.

Head. Antenna orange-brown, pedicel and tip of first

flagellomere blackish-brown. Arista stylus-like, with 2

aristomeres situated at tip of first flagellomere (Fig. 137).

Basal aristomere with projection about as long as api-

cal aristomere. Scape about four times longer than max-

imum width, apically and ventrally with setae. Pedicel

about six times longer than maximum width, apically and

most of dorsal surface covered with black setulae. Ped-

icel lacking any ridge at base, expanded towards apex.

First flagellomere long and conical, about three times as

long as high, pointed, ventrally with indistinct membra-

nous area. Lunule between base of antennae and ptilinal

suture distinct, slightly longer than width of scape. Eye

brown, lacking ommatrichia, facets all of about the same

size. Posterior margin of eye with distinct shining inden-

tation. Gena-height / eye-height (measurements taken

from head in lateral view) = 0.1. No ocellar tubercle, no

ocelli and no ocellar triangle evident. Frons (Fig. 135)

longer than broad, slightly concave, not projecting above

eyes posteriorly, lacking any setulae. Anterior margin of

frons slightly concave. Frons yellow with broad brown

midstripe. No frontofacial spot. Frons indistinctly dusted

all over. Vertex as broad as frons, separated from latter

by laterally distinct and medially indistinct ridge. Dense

cover of black setulae on vertex forming a more or less

level-topped pile in lateral view (Fig. 136). Vertex api-

cally with triangular field which lacks setulae or any

distinct depression. Face yellow with central brown to

blackish spot. Facial grooves with indistinct dusting, nar-

rowly dusted along eye margin. Gena yellow, generally

lacking setae, although with several setae below postcra-

nium. Distinct facial grooves reaching mouth edge, fa-

cial carina reaching from base of antennae to a distinctly

broadened and outstanding frontoclypeal tubercle. Ptili-

nal suture stretching on either side well beneath antennal

bases. Oral cavity tapers dorsally. Postcranium not obvi-

ously invaginated, black dorsally and yellow ventrally.

Bonn zoological Bulletin 68 (2): 209-273

Whole postcranium slightly dusted, with no obviously

denser dusting adjacent to posterior margin of eye. Oc-

ciput and postgena covered with black setulae. Postge-

na not widened and not delimited from occiput. Bottom

portion of postcranium distinctly separated-off and not

setulose. Proboscis reddish-brown basally and black api-

cally, labellum blackish-brown. Frontoclypeal membrane

long, light orange-brown and hardly delimited from or-

ange-brown clypeus. Palps absent. Labium longer than

head-length in lateral view, thickened basally, anterior

section completely fused into a tube. Labrum as long as

labium but very narrow. Labellum short, completely di-

vided, hardly broader than adjacent haustellum, and cov-

ered with very short setulae.

Thorax mainly black, with pleura black to brown.

Postpronotum orange-brown. Thorax evenly grey-dusted

all over, lacking any distinctly denser dusting or shining

areas. Presternum distinct, broad. Basisternum broad, not

narrowed ventrallyh to a point, lacking setae or setulae.

Proepisternum also lacking setae or setulae. Scutum cov-

ered with black setae. Notopleuron and postalarcallus

with several stronger setae. Postalarcallus lacking any

curved setulae beneath the black setae. Katepisternum

with 10-15 setae posterodorsally, 1-2 setae medially,

and no setae ventrally. Metakatepisternum, anepisternum

and anepimeron lacking setae or setulae. Mediotergite

convex, covered with strong black setae which can be

as long as maximum diameter of hind femur. Subscute-

Ilum inconspicuous. Scutellum densely covered with

black setae and with 3 outstanding larger setae on pos-

terior margin. Wing as Fig. 142: Fore-margin of wing

tinged brown, with brownish membrane between costa

and media, brown basal-medial cell and slightly brown-

ish hind-margin of discal-medial cell. Hyaline membrane

between media and vena spuria in cell r,,.. Veins brown

to black. Wing completely covered with microtrichia.

Radial-medial crossvein short but complete. Basal-me-

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New conopid records from the Afrotropical Region (Diptera) — Part 3: Physocephalini 267

Figs 134-137. Physocephala goergeni spec. nov. (2, holotype). 134. Habitus, lateral view; 135. Frons, dorsal view; 136. Lev-

el-topped pile of setulae on vertex, lateral view; 137. Arista, lateral view.

dial-cubital crossvein incomplete. Radius R, and R,,,

terminate close together in costa, well beyond end of

subcosta. Radius R,,. with shallow, even curve in dis-

tal section directed towards fore-edge of wing. Cell r,,.

pedunculate, vein R,,.+M, well distinctly longer than

radial-medial crossvein. Cubital cell cup elongated, lon-

Bonn zoological Bulletin 68 (2): 209-273

ger than vein A,+CuA,, and pointed distally (i.e. cubitus

CuA, and anal vein A, meet at an acute angle). Cubital

veins CuA, and Basal-medial-cubital crossvein separat-

ed. Upper and lower calypters yellowish-white to brown,

upper calypter with black setulae on margin. Alula broad

(distinctly broader than long), lacking setulae on posteri-

©ZFMK

268 Jens-Hermann Stuke & David K. Clements

Figs 138-142. Physocephala goergeni spec. nov. (2, holotype). 138. 9 theca, lateral view; 139. 9 theca, posterior view; 140.

Middle tibia, dorsal view; 141. Mediotergite, posterior view; 142. Wing, dorsal view.

or margin. Venae spuriae pronounced in cell r,, ., cubital

cells cup and cual, and discal-medial cell. Haltere white,

with light brown base. Knob of haltere with black setulae.

Legs orange with black to dark brown tarsi. Legs with in-

conspicuous silver dusting, coxae densely silver-dusted.

Posterior surfaces of fore and middle tibiae with obvious

silver-dusted fields distally. Legs generally with short,

adpressed black setulae. Base of fore and middle femo-

ra lacking denser black setulae basally. Areas with dense

black to brown setulae anteroventrally on tip of fore tibia,

and ventrally and posteriorly on tip of hind tibia. Middle

femur lacking a distinct row of regularly arranged setu-

lae. Hind femur ventrally with several long thin setulae.

No preapical setae dorsally on tibiae. No setae ventral-

Bonn zoological Bulletin 68 (2): 209-273

ly on tibiae but all tibiae dorsally with distinct line of

densely-arranged small black setulae (Fig. 140). Femo-

ra ventrally lacking rows of short black setae, but with

indistinct lines of setulae. Coxae with several setae but

none outstandingly long. Hind femur slightly thickened

in basal half. Each metatarsus with 1—2 stronger seta ven-

trally on base. Pulvilli yellowish-white. Claws brown,

with broad black tips. Empodium light brown, about as

long as pulvilli.

Abdomen dark brown to black, with theca and later-

al margins of tergites 2-3 orange-brown. Abdomen with

dense black setulae all over, those on tergite 2 less dense

and obviously longer. Abdomen entirely somewhat sil-

ver, brown or golden-dusted, more strongly so at hind

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New conopid records from the Afrotropical Region (Diptera) — Part 3: Physocephalini 269

margins of tergites 1-3. Tergite 8 shining. Tergites 1-3

fused and hardly distinguishable from each other. Max-

imum width of abdomen at segment 3. Sternites cannot

be seen due the ventrally overlapping tergites. Tergite 5

and sternite 5 not completely fused laterally. Shape of

theca as Figs 138 & 139. Anterior surface of the theca

with long black setulae. Posterior surface apically with

crescentric field of long, blunt, close-set black spicules

which stand very close together, not arranged in rows.

Diagnosis

Physocephala goergeni 1s easily identified as member of

the Physocephala pubescens species-group by the long

setae on the mediotergite (Fig. 141). Within this group it

belongs among those species which lack any setulae on

the frons. The shape of the field of thick black setae on

the theca distinguishes 9 P. goergeni immediately from

P. nigrita (Fig. 139 vs. Fig. 146). Additional important

characters include the level-topped pile of setulae on the

vertex (Fig. 136) and the shorter basal aristomere (cf Fig.

137 vs. Fig. 145).

Etymology

This species is dedicated to Georg Goergen (Ibadan, Ni-

geria) who collected the holotype, and who appears to

be the first entomologist to collect Conopidae in Benin

and Togo.

Distribution

To date only the locus typicus in Togo is known. The

sampling site is situated at 6°57°31.66"N 0°34)29.75"E.

P. hirta (Kroéber, 1939)

Pseudophysocephala hirta Krober 1939

Primary type material examined. 9 holotype of Pseu-

dophysocephala hirta Krober 1939: (1) “Holo-/ type”; (2)

“Type”; (3) “E. Cape Prov. / Katberg. / 1.—10.11.1933.”;

(4) “S. Africa. / R. E. Turner. / Brit. Mus. / 1933-139.”;

(5) “Pseudophysocephala / hirta Krober / examined &

det. /O. Krober, 1938.”; coll. NHML

See comment under Physocephala barbata (Camras,

2001), above.

P. nigrita (Camras, 1962)

(Figs 143-146)

Pseudophysocephala nigrita Camras 1962

= Pseudophysocephala brevivertex Camras

(Syn. nov.)

2001

Material. CAMEROON: 19, = 17—19.viii.2003,

Northwest Reg., Mezam, Bafut village [06°05.026'N

10°07.442'E], 1060 m, leg. A. H. Kirk-Spriggs, coll.

BMSA; KENYA: 29, 4.—-11.xii.1999, Western Province,

Kakamega Forest, Malaise trap [0°14.13'N 34°51.87'E],

Bonn zoological Bulletin 68 (2): 209-273

leg. R. Copeland, coll. NMKE; 19, 16.—23.viii.2000, dit-

to; TOGO: 12, ii.2017, Kloto, forest area [6°57’31.66"N

0°34)29.75"E], leg. G. Goergen, coll. IITA; 19, viii.2015,

ditto; UGANDA: 12, vii.—viii.1946, Bwamba, leg. van

Someren, coll. NHML [NHMUK010922185]; 19, vii—

viii. 1946, ditto [NHMUK010922187]; 14, 16.xii.1934,

Kampala, flying against bees, leg. T. W. Chorley, coll.

NHML [NHMUKO010922191]; 14, 18.ii.1966, Lake

Nabugabo, det. as P. nigrita by Camras 2000, leg. D. J.

Greathead, coll. NHML [NHMUK010922156].

The main character given to distinguish P. brevivertex in

the original description of Camras (2001) was a “short

vertex”. This character is difficult to assess, however, be-

cause the vertex may be tightly curved and of variable

shape in the material to hand. Apically the vertex has an

area which is not covered with setulae and which may

have a depression, but the shape, size and extent of any

depression can vary considerably, giving a different im-

pression of the vertex shape. There is no other character

available to distinguish P. brevivertex from P. nigrita, and

therefore Physocephala |Pseudophysocephala| breviver-

tex Camras, 2001 is herewith placed as a junior synonym

of Physocephala | Pseudophysocephala| nigrita Camras,

1962 (syn. nov.).

P. pubescens Brunetti, 1925

Physocephala pubescens Brunetti 1925

= Physocephala curta Krober 1936

(Figs 147-151)

Primary type material examined. 3 lectotype of Phys-

ocephala pubescens Brunetti 1925 herewith designated:

(1) “Syn - / type”; (2) “23.9.16 / Limbe / Nyasaland. /

4000' (R. C. W.)/ R. b. Wood / 684.”; (3) “Physo. / pu-

bescens / Brun Type 4 / Det. E. Brunetti 1924”; (4) “Lec-

totypus / Physocephala / pubescens 3 / Brunetti, 1925 /

des. Stuke 2019”; NHML.

9 lectotype of Physocephala curta Kroéber, 1936 here-

with designated: (1) “Type / Ph. curta 2”; (2) “Musée

du Congo / Elisabethville / 1927 / Dr. M. Bequaert”;

(3) “Physocephala / curta Krb. / det. Krober 1935”; (4)

“R. DET / L / 2992”: (5) “Type”; (6) “RMCA ENT /

000012182”; coll. MRAC.

Additional material. DEMOCRATIC REPUBLIC OF

CONGO: 1, no date, Elisabethville [Lubumbashi]

[-11.664232 27.482626], det. as P. pubescens by Cam-

ras 1962, leg. M. Bequaert, coll. FMNH; KENYA: 19,

11.1932, Kijabe, det. as P pubescens by Camras 1962,

leg. van Someren, coll. NHML [NHMUK010922169]:

12, 20.11.1949, Nairobi, det. as P. pubescens by Camras

1962, leg. G. Salt, coll. NHML [NHMUK010922168];

14, 9.iv.1949, Nairobi, det. as P. pubescens by Camras,

2000, leg. G. Salt, coll. NHML [NHMUK010922188];

19, 3.vi1.1994, Nairobi, leg. R. Copeland, coll. NMKE;

©ZFMK

270 Jens-Hermann Stuke & David K. Clements

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Figs 143-146. Physocephala nigrita Camras (1962). 143. Frons, dorsal view (3, Bwamba); 144. Mediotergite, posterior view (9,

Bafut village); 145. Arista, lateral view (°, Bafut village); 146. 2 theca, ventral view; (°, Bafut village).

14, 12.viii.1966, Nairobi, 5400 ft, det. as P. pubes-

cens by Smith, 1967, leg. G. R. C. van Someren, coll.

NHML [NHMUK010922186]; 13, 2.i1.1997, Nairobi,

Kebete, 1800 m, leg. T. Romig, coll. PMHA; MALA-

WI: 16, 17.iii.1913, Nyasaland, Mlanje [= Mulanje],

des. as syntype of P. pubescens, leg. S. A. Neave, coll.

NHML [NHMUKO010922166]; 1, 6.xi.1912, Nyas-

aland, Mt. Mlanje [= Mulanje Massif], des. as syn-

type of P pubescens, leg. S. A. Neave, coll. NHML

[NHMUK010922165]; 1 specimen, 1.x.1913, Nyas-

aland, Mt. Mlanje [= Mulanje Massif], des. as syn-

type of P pubescens, leg. S. A. Neave, coll. NHML

[NHMUK010922164]; UNKNOWN LOCATION: 1°,

iv.1941, “Mtowambo”, det. as P. pubescens by Camras

1962, leg., coll. FMNH; 19, viii.1943, [characters illeg-

ible], det. as P pubescens by Camras 1962, leg. H. J. A.

Turner, coll. NHML [NHMUK010922167].

The material held under ‘Physocephala pubescens* at

the NHML includes four syntypes of P. pubescens and

two syntypes of P. curta. The former comprises a mixture

of P. pubescens and P. caenostylata Krober, 1936. It is

Bonn zoological Bulletin 68 (2): 209-273

therefore necessary to designate a lectotype for P. pubes-

cens and we herewith select a male syntype from Limbe

which agrees with the species concept as here defined

in order to prevent any change in the past usage of this

name.

Stuke (2017a) incorrectly stated that Smith (1980) had

synonymized P. curta with P. pubescens when in fact this

was done by Camras (1962). Re-examination of the two

P. curta syntypes held at NHML concluded that these

also comprise multiple species. The female syntype from

Elisabethville is identical with P. pubescens, and in order

to prevent further confusion we herewith designate this

specimen as the lectotype of P. curta. The second syn-

type belongs to P. microvena Brunetti, 1925 and bears

the labels: (1) “Paratype / Ph. curta 2”; (2) “Musée du

Congo / Ituri: Blukwa / 10 - XI - 1928 /A. Collart’; (3)

“Cotype”; (4) “Physocephala & / curta Krb. / det. Kréber

1935”: (5) “R. DET / M/ 2992”; coll. MRAC. This spec-

imen has, unusually, a distinctly developed radial-medial

crossvein and could therefore easily be confused with

other species.

©ZFMK

New conopid records from the Afrotropical Region (Diptera) — Part 3: Physocephalini 271

Figs 147-151. Physocephala pubescens Brunetti 1925. 147. Frons, dorsal view (<, Nairobi); 148. Mediotergite, posterior view

(4, Nairobi); 149. Arista, lateral view (4, Nairobi); 150. 9 theca, posteroventral view (9, Nairobi); 151. & theca, lateral view (9,

Nairobi).

Physocephala vittata species-group

This group contains only P. vittata, which Camras (2001)

included (as P. maculigera Krober, 1915) in his Phys-

ocephala maculipes species-group. The vittata group is

very similiar to the Physocephala antiqua species-group

due to its typical Physocephala habitus (as described in

Key 1) and short arista. Both species-groups can be dis-

tinguished by the characters given in Key 1, however. A

Bonn zoological Bulletin 68 (2): 209-273

second member of this species-group — P. schmideggeri

Stuke, 2017 — has been recorded from the Arabian Penin-

sula and could also potentially occur in the Afrotropical

Region. These species can be distinguished using Stuke

(2017b). Physocephala vittata is primarily a widespread

Palaearctic species which reaches as far as Mongolia and

China, as well as extending into the Afrotropical Region.

©ZFMK

242

P. vittata (Fabricius, 1794)

Conops vittata Fabricius 1794

Material. ERITREA: 19, 28.1x.1957, Taramna, det. as

P. maculigera by Camras 2000, leg. D. J. Greathead,

coll. NHML [NHMUK010922106]; ETHIOPIA: 19,

3.iv.2016, SNNPS State, Arba-Minch, Dorze [06°10'N

37°35'E], 2340 m, leg. J. Halada, coll. CULSP.

Physocephala, species not recognised

P. nigerrima Krober, 1915

Physocephala nigerrima Krober 1915

Physocephala nigerrima was only known from the fe-

male holotype, which is probably lost (Stuke 2017a).

Camras (2001) placed this species in his Physocepha-

la maculipes species-group although this interpretation

does not fit with the original description. Krober de-

scribed the arista as being pointed and slender, and the

lateral projection also, with both being of almost equal

length (“Griffel spitz und zart, Seitenfortsatz ebenfalls,

beide fast gleich lang”) and the antenna as long, very

slender and black, with the first flagellomere rusty brown

and the scape about three times as long as its width at the

base and slightly widened towards the tip (“Fuhler lang

und sehr schlank, schwarz, drittes Glied rostbraun. Erstes

Glied etwa dreimal so lang als unten breit, oben wenig

verbreitert.”). This species cannot readily be placed and

in the absence of any available material we therefore pro-

pose to classify Physocephala nigerrima Krober, 1915 as

an unrecognised species (Status rev.: nomen dubium)

until such time as new material appears.

Acknowledgements. We would like to thank all of the col-

leagues who have supported this work with their generous help:

Miroslav Bartak (Czech Republic, Prague), Eliana Buenaventu-

ra (Germany, Berlin), Pasquale Ciliberti (Netherlands, Leiden),

Robert Copeland (Kenya, Nairobi), Wouter Dekoninck (Bel-

gium, Brussels), Hans-Joachim Fliigel (Germany, Kniillwald),

Amnon Freidberg (Israel, Tel Aviv), Georg Goergen (Benin,

Calavi), Martin Hauser (USA, Sacramento), Christian F. Kasse-

beer (Germany, Damlos), Ashley H. Kirk-Spriggs (formerly

South Africa, Bloemfontein, now UK, London), Tom Kirschey

(Berlin), Maurizio Mei (Italy, Rome), Ximo Mengual (Ger-

many, Bonn), Frank Menzel (Germany, Mtincheberg), Crystal

Maier (USA, Chicago), Marc de Meyer (Belgium, Tervuren),

Wolfgang Schacht (Germany, |), Karla Schneider (Germany,

Halle), Peter Sehnal (Austria, Vienna), Axel Ssymank (Ger-

many, Bonn), Nigel Wyatt (UK, London) and Joachim Ziegler

(Germany, Berlin), all of whom provided material either from

their private collections or from collections under their care.

Martin Hauser in particular sorted through a huge amount of

Conopidae from Malaise trap samples and made it available for

our research. While visiting collections to look at Afrotropical

Conopidae we received generous help from Nigel Wyatt (UK,

London), Erica McAlister (UK, London), Sven Marotzke (Ger-

many, Berlin), Jenny Pohl (Germany, Berlin), Joachim Ziegler

(Germany, Berlin), Eliana Buenaventura (Germany, Berlin),

Bonn zoological Bulletin 68 (2): 209-273

Jens-Hermann Stuke & David K. Clements

Marc de Meyer (Belgium, Tervuren), Stéphane Hanot (Bel-

gium, Tervuren), Wouter Dekoninck (Belgium, Brussels) and

Frank Menzel (Germany, Muncheberg). With great skill, Nygel

Wyatt decoded illegible writing and characters on the labels of

many historical specimens.

REFERENCES

Borkent A (2018) The state of Phylogenetic Analysis: Narrow

Visions and Simple Answers — Examples from the Diptera

(Flies). Zootaxa 4374: 107-143

Camras S (1957) On some Conopidae (Dipt.) from Flores and

Sumba. Wissenschaftliche Ergebnisse der Sumba-Expedition

des Museums fiir Volkerkunde und des Naturhistorischen

Museums in Basel, 1949. Verhandlungen der naturforschen-

den Gesellschaft in Basel 68: 160—164

Camras S (1962a) The Conopidae of Madagascar (Diptera).

Mémoires de I’ Institut Scientifique de Madagascar, serie E

13: 179-187

Camras S (1962b) Records and descriptions of African Conopi-

dae (Diptera). Revue de Zoologie et de Botanique Africaines

66: 203-242

Camras S (1965) Family Conopidae. Pp. 625-632 in: Stone A,

Sabrosky CW, Wirth WW, Foote RH & Coulson JR (eds) A

catalog of the Diptera in America North of Mexico. Smitho-

nian Institution Press, Washington

Camras S (2001) Additional information on Afrotropical

Conopidae. Entomologist’s Monthly Magazine 137: 179-210

Cumming JM, Wood DM (2009) Adult Morphology and Ter-

minology. Pp. 9-50 in: Brown BV, Borkent A, Cumming JM,

Wood DM, Woodley NE & Zumbado MA (eds): Manual of

Central American Diptera. Volume 1: 1-714. NRC Research

Press, Ottawa.

GBIF (2019) Free and open access to biodiversity data.

https://www.gbif.org [last access 18.11.2019]

Gibson JF, Skevington JH (2013) Phylogeny and taxonomic

revision of all genera of Conopidae (Diptera) based on mor-

phological data. Zoological Journal of the Linnean Society

167: 43-81

Kotrba M (2000) 1.3 Morphology and terminology of the fe-

male postabdomen. Pp. 75—84 in: Papp L & Darvas B (eds):

Contributions to a Manual of Palearctic Diptera. Volume 1.

General and Applied Dipterology. Science Herald, Budapest.

Krober O (1915) Die afrikanischen Arten der Gattung Phys-

ocephala Schin. Archiv fiir Naturgeschichte, Abteilung A

80(11): 81-99

Krober O (1939) Beitrage zur Kenntnis der Conopiden. I. An-

nals and Magazine of Natural History 11(4): 362-395

Smith KGV (1980) 39. Family Conopidae. Pp. 511-517 in:

Crosskey RW (ed.): Catalogue of the Diptera of the Afro-

tropical Region. British Museum of Natural History, London.

Smith KGV, Cunningham-van Someren GR (1970) The identity

of Physocephala bimarginipennis Karsch (Diptera, Conopi-

dae) with notes on the immature stages and biology. Journal

of Natural History 4: 439-446

Smith KGV, Peterson BV (1987) 54. Conopidae. Pp. 749-756

in: McAlpine JF, Peterson BV, Shewell GE, Teskey HJ,

Vockeroth JR & Wood DM (eds): Manual of Nearctic Dip-

tera. Volume 2. Monograph of the Research Branch Agricul-

ture Canada 11: vi+ 658 pp; Ottawa.

Stuke J-H (2015a) New Conopid records from the Afrotropical

Region (Diptera). Part 1: Paramyopa Kréber, Pseudoconops

Camras, Stylogaster Macquart, Thecophora Rondani and Zo-

dion Latreille. Zootaxa 3963: 101-159

©ZFMK

New conopid records from the Afrotropical Region (Diptera) — Part 3: Physocephalini 273

Stuke J-H (2015b) A contribution to the knowledge of Physo-

cephala antiqua (Wiedemann) (Diptera: Conopidae). Studia

Dipterologica (Mtncheberg) 21: 209-220

Stuke J-H (2016): Taxonomic notes on West Palaearctic Conop-

idae (Diptera). Zootaxa 4178: 521-534

Stuke J-H (2017a): World Catalogue of Insects. Volume 15.

Conopidae (Diptera). E. J. Brill, Leiden, Boston

Bonn zoological Bulletin 68 (2): 209-273

Stuke J-H (2017b): Order Diptera, family Conopidae. Descrip-

tion of a new species of Physocephala Schiner. Arthropod

Fauna of the UAE 6: 613-620

Stuke J-H (in press): New conopid records from the Afrotrop-

ical Region (Diptera). Part 2: Conopinae excluding Physo-

cephalini. Israel Journal of Entomology

©ZFMK

BHL

Blank Page Digitally Inserted

Bonn zoological Bulletin 68 (2): 275-296

2019 - Thormann J. et al.

https://do1.org/10.20363/BZB-2019.68.2.275

ISSN 2190-7307

http://www.zoologicalbulletin.de

Report**

A prelude to the Caucasus Barcode of Life Platform (CaBOL):

Biodiversity Days in Georgia in 2018 and 2019

Jana Thormann’, Dirk Ahrens', Cort Anderson’, Jonas J. Astrin’’, Levan Mumladze’, Bjérn Rulik’,

David Tarkhnishvili', Marianne Espeland', Matthias Geiger', Nils Hein‘, Giorgi Iankoshvili’,

Elisabeth Karalashvili?, Ximo Mengual', Carsten Morkel*’, Marco T. Neiber®, Ralph S. Peters’,

André Reimann’, Axel Ssymank’®, Thomas Wesener', Joachim Ziegler? & Bernhard Misof'”

' Zoologisches Forschungsmuseum Alexander Koenig, Leibniz Institute for Animal Biodiversity, Center for Taxonomy and

Evolutionary Research, Adenauerallee 160, D-53113 Bonn, Germany

? Center of Biodiversity Studies, Institute of Ecology, Ilia State University, Cholokashvili ave.3/5, GE-0169 Tbilisi, Georgia

3 Institute of Zoology, Ilia State University, Cholokashvili ave.3/5, GE-0169 Tbilisi, Georgia

* School of Natural Sciences and Engineering Ilia State University, 3/5 Choloqashvili Ave, GE-Tbilisi 0162, Georgia

° Institute for Applied Entomology, Bartholomdusstrasse 24, D-37688 Beverungen

°Centrum fiir Naturkunde (CeNak) — Center of Natural History, Abteilung Biodiversitat der Tiere, Martin-Luther-King-Platz 3,

D-20146 Hamburg, Germany

’Senckenberg Naturhistorische Sammlungen Dresden, Museum fir Tierkunde, Konigsbriicker Landstrape 159,

D-01109 Dresden, Germany

° Falkenweg 6, D-53343 Wachtberg

? Museum fiir Naturkunde, Invalidenstrasse 43, D-10115 Berlin, Germany

* Corresponding authors: Email: j.astrin@leibniz-zfmk.de; b.misof@leibniz-zfmk.de

Abstract. Here we report on collaborative expeditions run by ISU (Ilia State University, Tbilisi), ZFMK (Zoologisches

Forschungsmuseum Alexander Koenig, Bonn) and external taxonomists in Georgia in 2018 and 2019 as part of the rene-

wed close cooperation between Germany and Caucasus countries aiming at the exploration of biodiversity in the Caucasus

region. The 2018 and 2019 field work campaigns serve as a starting point for a larger cooperative venture, planned to

start in 2020 — the Caucasus Barcode of Life Platform (CaBOL). To encourage participation by additional partners and

to build an active expert network, we here describe the so-far collected material and collection methods, provide detailed

information on collecting sites and the diversity of habitats and outline the goals of the planned CaBOL project. Building

on experiences from the German Barcode of Life (GBOL) initiative, our goal is to increase expertise and capacity in the

region for modern integrative taxonomy approaches.

Keywords. GBOL, GGBC, DNA barcoding, Kintrishi, Kazbeg1.

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INTRODUCTION

The Caucasus Region is an area situated between the

Black and the Caspian Sea, and is recognized as one of

the global biodiversity hotspots (Mittermeier et al. 2004;

Myers et al. 2000; Zazanashvili et al. 2004). Where West-

ern Asia meets Eastern Europe, Georgia is bounded to the

west by the Black Sea, to the north by Russia, to the south

by Turkey and Armenia, and to the southeast by Azerbai-

**T his report was not subject to peer review

jan. While making rapid strides towards modernization of

its research and educational capacity, Georgia still lacks

some key elements in both expertise and infrastructure.

Targeted investment in science and education in Georgia

is one of the major goals of our research collaboration,

and perhaps the best way to develop and augment capac-

ity for the essential biodiversity assessment in situ. It is

most effective in close cooperation with an experienced

partner. Germany and the Zoological Research Museum

Published: 11.12.2019

276

Alexander Koenig, Bonn (ZFMK) are leaders in biodi-

versity research within Europe, for example, coordinat-

ing the German Barcode of Life (GBOL) initiative that

has been successfully run for the last eight years and has

delivered an operative DNA barcode reference library for

German animals, plants and fungi (Geiger et al. 2016).

Based on this Germany-focused biodiversity research,

scientists from Georgia and Germany decided to launch a

collaborative initiative to adapt the structure and strategy

of the GBOL project to enable comprehensive biodiversi-

ty research in the species-rich Caucasus region. This pa-

per documents the joint field work done so far, and gives

an overview of the ongoing and future collaboration.

PREVIOUS PROJECTS AND TRANSFER OF DNA

BARCODING WORKFLOWS

Since 2012 ZFMK has led the GBOL project in which

several German research institutions successfully worked

together on a high-throughput sequencing pipeline to

create an extensive DNA barcode library (Hebert et al.

2003a, 2003b) of the German fauna, flora and fungi in

order to assess and access the national biodiversity (e.g.,

Raupach et al. 2014, Wesener et al. 2015, Astrin et al.

2016, Moriniere et al. 2017, Rulik et al. 2017). Applying

DNA barcodes for species identification and delimita-

tion requires a database of reference sequences. In order

to develop such a reference library, concerted efforts of

taxonomists, biodiversity data specialists, and experts on

molecular techniques are necessary. In 2017, the ZFMK

made the first steps for a knowledge transfer from the

GBOL project to Georgia, a country with a consider-

able need and potential for barcode-aided biodiversity

assessment (Mumladze et al. 2019). In September 2017,

a proposal for the development of a Georgian-German

biodiversity center (GGBC) was granted by the German

Federal Ministry of Education and Research (BMBF)

(project’s website: https://ggbc.eu/). The goal of the

GGBC is to establish an infrastructure and to provide

suitable training that allows the long-term development

of a DNA barcode reference database for Georgia.

The operational infrastructure of the GBOL initiative

is Supposed to serve as a model, guiding the development

of a comparable structure in Georgia. Such developments

of the relevant infrastructures are planned to go hand in

hand with taxonomic and methodological training of

Georgian students, species identification by taxonomic

experts, as well as molecular laboratory work. To this

end, a lively exchange of students and researchers has

already been established between ISU (Ilia State Univer-

sity, Tbilisi) and ZFMK.

Bonn zoological Bulletin 68 (2): 275-296

Jana Thormann et al.

THE KINTRISHI PROJECT: MALAISE TRAP

TRANSECT & BIODIVERSITY DAYS 2018

Within the framework of GGBC, the Kintrishi project

was the first pilot project launched by the ISU-ZFMK

collaborative team in April 2018. The project aimed to

investigate the biodiversity in the Kintrishi Protected

Areas and to start compiling a Georgian reference data-

base of DNA barcodes for selected taxa. Created in 1959,

the Kintrishi National Park is located in south-western

Georgia (41°75' N, 42°03' E) and currently encompasses

over 13,893 hectares (APA, 2019). Together with the ad-

jacent Mtirala National Park, Kintrishi is the most humid

area in the Caucasus region, with an annual precipitation

of around 2500 mm (Neidze, 2003). It is a Plio-Pleis-

tocene refugium featuring many relict and endemic ani-

mal and plant species (Kikvidze & Ohsawa 1999, Denk

et al. 2001, Milne & Abbott 2002, Shatilova et al. 2011,

Tarkhnishvili et al. 2011). The park ranges from 300 m

up to 2,500 maz.s.l. and includes pristine mountain humid

forests and subalpine/alpine habitats. Although the high

level of biodiversity in Kintrishi was recognized early in

the 20th century a validated catalog of animal and plant

taxa in Kintrishi and surroundings is still lacking (Gars-

tecki 2017).

In April 2018, Bjorn Rulik and Ameli Kirse from the

ZFMK spent two weeks in Georgia. Together with Le-

van Mumladze (ISU) and a group of Georgian students

they set up Malaise traps along an elevational gradient

in the Kintrishi gorge, from the lower reaches of the

Kintrishi River (400 m as.l.) up to alpine meadows

(2,500 m a.s.l.). Three trap replicates were placed at each

of the six selected elevation levels (1.e., 18 traps in total)

(Figs 1-3). The traps were emptied every two weeks by

a team of Georgian students: Eka Arsenashvili, Giorgi

Bakuradze, Giorgi Bananashvili, Giorgi Chikorashvili,

Shota Japarashvili, Tinatin Chkhartishvili, Giorgi Ian-

koshvili, Elisabeth Karalashvili, Revaz Kvaratskhelia,

Natia Rtskhiladze, Nutsa Rtskhiladze, Alisa Sanakoeva,

Anano Shubashishvili, Irina Tsereteli and Mari Tsulaia.

The traps were operating from April to November 2018.

From the collected samples several taxa were then sorted

out at ISU and prepared for further taxonomic research

and DNA barcoding, using equipment purchased by the

GIZ (Deutsche Gesellschaft fiir Internationale Zusam-

menarbeit; see below). Since the amount of collected

material was very large, this work is still in progress and

we are currently focusing on some selected groups of an-

imals (several families of dipterans, hymenopterans, and

beetles; spiders, pseudoscorpions and myriapods). The

proximate goal is to evaluate multi-taxon species diver-

sity and community structure over an elevation gradient.

Furthermore, microclimate data logger stations register-

ing air temperature, soil temperature, relative humidity

and solar radiation were installed along the transect.

©ZFMK

A prelude to the CaBOL Platform: Biodiversity Days in Georgia in 2018 and 2019 247

Photo: Giorgi lankoshvili

Fig. 9 Photo: Katharina Kurzrock

Figs 1-9. The Kintrishi Project. 1. Bjorn Rulik (ZFMK) and Levan Mumladze (ISU) with ISU students. 2. Malaise trap above the

tree line (2500 m). 3. Bjorn Rulik and Ameli Kirse (ZFMK) teaching students at Grigoleti Research Field Station. 4. Grigoleti field

station. 5. Traditional Georgian dinner prepared by local contractors from Grigoleti. 6. On the way to Kintrishi Protected Areas. 7.

Just the “normal” way to work. 8. Bridge in Kintrishi Protected Areas. 9. Electrofishing in Kintrishi River.

278 Jana Thormann et al.

Photemana Thormann

nie

Photo: Jana Thormann

Photo: Jana Thormann bie Photo: Jana Thormann

Figs 10-17. Biodiversity Days 2018. 10. Curious pigs examining fishing gear. 11. Changing a flat tire on the way to work. 12.

Bakhmaro. 13. Sceptical glance in Borjomi NP. 14. Collecting calves?! (Borjomi NP). 15. Hill top near Lentekhi. 16. Traditional

dinner in a private accommodation in Lentekhi. 17. Between Lentekhi and Tsana.

A prelude to the CaBOL Platform: Biodiversity Days in Georgia in 2018 and 2019

Besides the Malaise trap transect two additional Mal-

aise traps were set up: one was located next to the field

station in Grigoleti for one week in April 2018. The other

trap was set up close to the rangers’ station in Kintrishi,

who emptied the trap every other week, alternating with

the ISU team. This trap was contributed by the Global

Malaise Trap Program (GMP; https://biodiversitygenom-

ics.net/projects/gmp/), which is run by the Centre for Bio-

diversity Genomics, University of Guelph, Canada. After

the collecting season the material was sent to Guelph to

obtain morphospecies-based barcode sequences.

In addition to Malaise trap sampling, soil samples have

been collected along the elevation gradient to run a me-

tabarcoding study of Nematoda diversity and pitfall traps

were also set-up to sample ground beetle communities in

the Kintrishi area.

In July 2018, a group of Georgian and German scien-

tists, students and external taxon specialists (including

citizen scientists) visited the Kintrishi Protected Areas to

collect and identify specimens for DNA barcoding with-

in so-called Biodiversity Days (1.e., a concerted, intense

multi-taxon field sampling campaign, also referred to as a

Bioblitz), the first so far to take place in Georgia. During

the core dates (20—22 July) all participants were accom-

modated at the ISU field station in Grigoleti (Figs 4-5).

During the days before and after these core days several

groups of scientists also visited other areas of the country

(e.g., the Greater Caucasus) to collect specimens (Appen-

dix I: Table 1; Map (Fig. 56), Figs 12—21).

Apart from the pilot project funding by the BMBF

(German Federal Ministry of Education and Research),

the GIZ Georgia kindly supported this event under the

Integrated Biodiversity Management South Caucasus

(IBiS) initiative, funded by the BMZ (Federal Ministry

for Economic Cooperation and Development). Collecting

permits were provided by the Agency of Protected Areas

of Georgia and the Ministry of Environment Protection

and Agriculture of Georgia.

In addition to the students and scientists who were al-

ready involved into the Malaise trap project (listed above)

the following researchers participated in the Biodiversity

Days 2018: Cort Anderson (ISU), Jonas Astrin (ZFMK —

Arthropoda), Ani Bikashvili (ISU — Mollusca), Tim Boh-

nert (Uni Bonn — Botany), Albia Consul (ZFMK — her-

petology, Arthropoda), Giorgi Epitashvili (ISU — Fishes),

Marianne Espeland (ZFMK — Trichoptera, Lepidoptera),

Matthias Geiger (ZFMK — Fishes), Nils Hein (Geogra-

phisches Institut der Universitat Bonn — Arachnida), Hajo

Krammer (ZFMK — Arachnida), Katharina Kurzrock

(ZFMK — Mollusca), Ximo Mengual (ZFMK — Diptera),

Stefan Otto (independent researcher — Arachnida), André

Reimann (GBOL team; Senckenberg Naturhistorische

Sammlungen Dresden — Diptera, Scorpiones), Anke

Schafer (GBOL team; independent researcher — Dip-

tera, Botany), David Tarkhnishvili (ISU — herpetology),

Birthe Thormann (ZFMK — Coleoptera), Jana Thormann

Bonn zoological Bulletin 68 (2): 275-296

219

(ZFMK — Arthropoda), Sénke Twietmeyer (GBOL team;

Nationalpark Eifel — Aves, Mammalia), Thomas Wesener

(ZFMK — Myriapoda), Benedict Wipfler (ZFMK — Co-

leoptera).

Combining morphological and molecular data for spe-

cies investigations, the collecting efforts in 2018 result-

ed in the submission of a first publication, which gives

an updated list of the hoverflies (Diptera, Syrphidae)

of Georgia (Mengual et al., submitted). This collabora-

tion of Georgian students and external taxon specialists

contributes to the species inventory and DNA barcoding

reference library and is an example of the successful co-

operation between ISU, ZMFK and external researchers.

BIODIVERSITY DAYS 2019: THE GREATER CAU-

CASUS

After the successful trip to the Kintrishi area, the Biodi-

versity Days 2019 took place in the Stepantsminda area

(Kazbegi) in the Greater Caucasus. During the core dates

(4-7 July 2019) the participants were hosted at the ISU-

field station in Stepantsminda and, as in the previous year,

small groups of scientists also collected in other regions

of the country (e.g., Vashlovani National Park) before and

after the core dates (Appendix II: Table 2, Map (Fig. 56),

Figs 22-40).

Three Malaise traps were run during a few days around

the core dates. In addition, yellow pan, light, pitfall and

banana traps were used, as well as Winkler extractors for

sifted leaf litter.

In addition to sampling described above, collecting of

Heteroptera was done in a variety of habitats, with a focus

on brachypterous species of subalpine meadows. So far,

two Caucasian endemics have already been identified,

one of them just recently described as new to science:

Scirtetellus gudali Kiritshenko, 1951 and Myrmecophyes

(Plumiger) tomi Konstantinov & Simov, 2018 (Fig. 54)

(both Heteroptera: Miridae).

A quick scan through malaise material collected in

Stepantsminda revealed specimens from at least 13 fam-

ilies of lower Diptera. Within the family Mycetophilidae

(fungus gnats) the following discoveries are worth men-

tioning: a big series (14 33, 11 2) of Coelosia flava

(Staeger, 1840), a typical boreo-montane faunal element

(Fig. 55) and two females of the widespread but rarely

collected species Grzegorzekia collaris (Meigen, 1818).

Both species records represent the first proof for the Cau-

casus region.

At least 48 different mollusk species belonging to 24

families were recorded during the core event in Stepants-

minda and a short trip from Tbilisi to Kutaisi 1-3 July

2019. Aside from more widespread Western Palearctic

Species and species occurring also in Anatolia, 16 species

endemic to the Caucasus region were recorded. Especial-

ly noteworthy are three slug species: the first, Svanetia

©ZFMK

280 Jana Thormann et al.

Espeland

Photo: Jana Thorménn

“¢ « Wis a

Photo:Jana Thormann ey { Photo: Matthias Geiger

Figs 18-26. The Greater Caucasus. 18. Road to Tsana. 19. Kiosk in Tsana. 20. Searching for spiders. 21. Collecting in Tsana,

Greater Caucasus (1760 maz.s.1.). 22. Malaise trap in the garden of Stepantsminda field station. 23. View from field station to Mount

Kazbek (5054 m) and Gergeti Trinity Church. 24. Gergeti Trinity Church. 25. View to Stepantsminda from Gergeti Trinity Church.

26. After field work.

A prelude to the CaBOL Platform: Biodiversity Days in Georgia in 2018 and 2019 281

caucasica (Simroth, 1898), is a short-range endemic that

is only known from the Kazbegi region and was record-

ed at the Jvaris Pass. The second is an unidentified spe-

cies of the genus Gigantomilax that was collected north

of Gergeti. The genus has previously not been recorded

from the Kazbegi region. The third species, Boettgeril-

la pallens Simroth, 1912, is only known from western

Georgia, where it was only rarely recorded in the past.

The following Georgian and German researchers at-

tended the Biodiversity Days in Stepantsminda: Cort

Anderson (ISU), Ulrich Burkhardt (Senckenberg Mu-

seum ftir Naturkunde Gorlitz — Collembola), Marianne

Espeland (ZFMK — Trichoptera, Lepidoptera), Matthi-

as Geiger (ZFMK — Fishes, Macrozoobenthos), Nils

Hein (Geographisches Institut der Universitat Bonn

— Arachnida), Kai Heller (GBOL team, independent

researcher — Diptera), Bella Japoshvili (ISU — Fishes),

Elisabeth Karalashvili (ISU — Arachnida), Thorsten

Klug (ZFMK — Myriapoda), Hajo Krammer (ZFMK —

Arachnida), Christiane Lange (independent researcher

— Diptera), Cornelia Lohne (Uni Bonn — Botany), Peter

Manko (University of Presov — aquatic Insects), Dirk

Mattern (GBOL team, independent researcher — Trichop-

tera), Ximo Mengual (ZFMK — Diptera), Carsten Morkel

(GBOL team, independent researcher — Heteroptera), Le-

van Mumladze (ISU), Marco T. Neiber (CeNak Hamburg

— Mollusca), André Reimann (GBOL team; Senckenberg

Naturhistorische Sammlungen Dresden — Diptera, Scor-

piones), Dirk Rohwedder (ZFMK — Hymenoptera), Anke

Schafer (GBOL team; independent researcher — Diptera,

Botany), Axel Ssymank (Bundesamt fir Naturschutz

Bonn — Diptera, Botany), Eckart Stolle (ZFMK — Hy-

menoptera), Jens-—Hermann Stuke (GBOL team, inde-

pendent researcher — Diptera), David Tarkhnishvili (ISU

— herpetology), Jana Thormann (ZFMK — Arthropoda),

Karin Voigtlander (Senckenberg Museum ftir Naturkunde

Gorlitz — Myriapoda), Benedict Wipfler (ZFMK — Cole-

optera), Joachim Ziegler (Museum ftir Naturkunde Ber-

lin — Diptera).

During the core dates several ISU students joined the

researchers: Eka Arsenashvili, Giorgi Bananashvili, Ani

Bikashvili, Tinatin Chkhartishvili, Sandro Chubinidze,

Giorgi Epitashvili, Giorgi Iankoshvili, Shota Japarashvi-

li, Giorgi Khubashvili, Giorgi Kirtkitadze, Revaz Kvar-

atskhelia, Natia Rtskhiladze, Nutsa Rtskhiladze, Alisa

Sanakoeva, Anano Shubashishvili, Mariam Todua, Ana

Tsertsvadze, Mariam Tsulaia and Mariam Zazadze.

According to their field of interest, the students accom-

panied the scientist during their work in the field and got

training in different collecting methods and taxonomic

identification.

An additional team of international scientists made an

additional expedition one month earlier: Ximo Mengual

(ZFMK), Sander Bot (Haren, Netherlands), Jeffrey H.

Skevington (Canadian National Collection of Insects,

Arachnids and Nematodes, Agriculture and Agri-Food

Bonn zoological Bulletin 68 (2): 275-296

Canada, Ottawa, Canada), Frank Van de Meutter (Engs-

bergen, Belgium), Jeroen van Steenis (Naturalis Biodi-

versity Center, Leiden, Netherlands) and Lenze Hofstee

(Haren, Netherlands) collected Diptera in different parts

of Georgia 8—23 June 2019.

Furthermore, a joint excursion of students from the Ge-

ography department of the University of Bonn (GIUB),

the ZFMK and the ISU was held in July/August 2019.

This event aimed at teaching knowledge on the under-

lying mechanisms of the high biodiversity in Georgia to

students with different backgrounds.

OUTLOOK

As a continuation of the work in GGBC, we currently

plan a Georgian-Armenian-German initiative to estab-

lish a joint Caucasus Biodiversity Research Initiative,

the Caucasus Barcode of Life Platform (CaBOL). The

planned improvements of research and training infra-

structure in a long-term international collaboration will

provide the latest technology in molecular biodiversi-

ty research to Georgian and Armenian scientists and

students. A strong initial focus of the project will lie

in expanding the DNA barcoding reference databases

for Georgia, as initiated within GGBC, and Armenia.

CaBOL will collect DNA barcodes, and archive DNA,

tissue samples and voucher specimens of Caucasian

biodiversity, to the benefit of collections in Georgia,

Armenia and Germany. The DNA barcodes and their

metadata will be publicly available in a centrally man-

aged database, and will be used for cooperative research

projects by Georgian, Armenian and German scientists

and students. Simultaneously, CaBOL will equip labora-

tories in Tbilisi (Georgia) and Yerevan (Armenia), with

a strong hub at the Ilia State University, to train students

from the South Caucasus region and beyond. Currently,

there are no advanced biotechnology facilities avail-

able for non-medical research in Georgia and Armenia.

Beyond DNA barcoding, infrastructural improvements

and its training mission, CaBOL will expand its method-

ological spectrum by remote sensing approaches and will

include experienced forestry partners from the Universi-

ty of Gottingen (Chair of Forest Inventory and Remote

Sensing).

Acknowledgements. The GGBC-project was funded by the

German Federal Ministry of Education and Research under

grant number 01DK17048. The two field expeditions were

supported by Deutsche Gesellschaft fiir Internationale Zusam-

menarbeit Georgia under the Integrated Biodiversity Manage-

ment South Caucasus initiative, funded by the Federal Ministry

for Economic Cooperation and Development (contract number

2018: 83291308, contract number 2019: 83322223). Collecting

permits were kindly provided by the Agency of Protected Ar-

eas of Georgia and the Ministry of Environment Protection and

Agriculture of Georgia. ZFMK kindly supported its employees

©ZFMK

282 Jana Thormann et al.

Cy 7

Photo: Jana Thormann

»Photo: Marianne Espeland

: —— =a et

—- /

Photo: Jana Thormann

Figs 27-34. Biodiversity Days 2019. 27. Analyzing dung samples. 28. Winkler extractor in repurposed wardrobe. 29. Daily way

to work. 30. Clearing the road. 31. ISU students accompany scientists to the field. 32. Analyzing soil samples in the field. 33. Sno

valley on the way to Juta. 34. Juta and view to Chaukhi massif.

A prelude to the CaBOL Platform: Biodiversity Days in Georgia in 2018 and 2019 283

Photo! Marianne Espeland

4 Photo; Jana Thormann

Photo: Hajo Krammer

Figs 35—42. Various collecting sites. 35. The fishing team. 36. Truso gorge. 37. View to Mna valley. 38. Sighnaghi and Greater Cau-

casus in the background. 39. Collecting on the way to Vashlovani Protected Areas. 40. Vashlovani Protected Areas. 41. Lesson in

the field during joint excursion with Georgian and German students. 42. Looking out for birds at Eagle Gorge near Dedopliszqaro.

284 Jana Thormann et al.

Fig. 44 Photo: Matthias Geiger

Fig. 46 Photo: M. Espeland Fig.47 Photo: Benedict Wipfler

Photo: Matthias Geiger ) Fig.51 ., Photo: M. Espeland

pidae, Panorpa communis Linnaeus, 1758. 45. Gastropoda: Hygromiidae, Caucasigena eichwaldi (Pfeiffer, 1846). 46. Lepi-

do-ptera: Lycaenidae, Polyommatus amandus (Schneider, 1792). 47. Coleoptera: Cerambycidae, Rosalia alpina (Linnaeus, 1758).

48. Gastropoda: Limacidae, Eumilax brandti (Martens, 1880). 49. Teleostei: Salmonidae, Salmo ciscaucasicus Dorofeyeva, 1967.

50. Araneae: Atypidae, Atypus muralis Bertkau, 1890. 51. Lepidoptera: Nymphalidae, Melanargia russiae (Esper, 1783). 52. Di-

ptera: Syrphidae, Epistrophe leiophthalma (Schiner & Egger, 1853). 53. Diptera: Rhinophoridae, Stevenia fausti (Portshinsky,

1875). 54. Heteroptera: Miridae, Myrmecophyes tomi Konstantinov & Simov 2018. 55. Diptera: Mycetophilidae, Coelosia flava

(Staeger, 1840).

A prelude to the CaBOL Platform: Biodiversity Days in Georgia in 2018 and 2019 285

4 42 43

Fig. 56. Map of collecting sites.

with logistics, equipment and travel expenses. We are thankful

to all participants of the Biodiversity Days for collecting, iden-

tifying and providing specimens for further molecular analy-

sis and thus helping to establish a reference database. We also

thank the Georgian students who supervised the Malaise traps

in Kintrishi and who are sorting the material.

REFERENCES

APA (2019) Agency of Protected Areas of Georgia. https://apa.

gov.ge/en/protected-areas/cattestone/kintrishis-daculi-terito-

riebis-administracia. Accessed 03/11/2019

Astrin JJ, Hofer H, Spelda J, Holstein J, Bayer S, Hendrich L,

Huber BA, Kielhorn KH, Krammer HJ, Lemke M, Monje

JC, Moriniere J, Rulik B, Petersen M, Janssen H, Muster

C (2016) Towards a DNA Barcode Reference Database for

Spiders and Harvestmen of Germany. PLoS ONE 11(9):

e0162624. https://do.org/10.1371/journal.pone.0162624

Denk T, Frotzler N, Davitashvili N (2001) Vegetational patterns

and distribution of relict taxa in humid temperate forests and

wetlands of Georgia Transcaucasia. Biological Journal of the

Linnean Society 72: 287—332

Garstecki T (2017) Feasibility assessment for a World Heritage

nomination of the Colchic Forests and Wetlands under the

natural criteria. Greifswald, Germany: pp. 132

Geiger MEF, Astrin JJ, Borsch T, Burkhardt U, Grobe P, Hand R,

Hausmann A, Hohberg K, Krogmann L, Lutz M, Monje C,

Bonn zoological Bulletin 68 (2): 275-296

@ collecting sites 2018

a collecting sites 2019

44 45 46 47

Misof B, Moriniere J, Muller KF, Pietsch S, Quandt D, Rulik

B, Scholler M, Traunspurger W, Haszprunar G, Wagele W

(2016) How to tackle the molecular species inventory for an

industrialized nation — lessons from the first phase of the Ger-

man Barcode of Life initiative GBOL (2012—2015). Genome

59(9): 661-670. https://doi.org/10.1139/gen-2015—-0185

Hebert PDN, Cywinska A, Ball SL, Dewaard J (2003a). Biolog-

ical identification through DNA barcodes. Proceedings of the

Royal Society of London B 270: 313-321

Hebert PDN, Ratnasingham S, DeWaard JR (2003b) Barcod-

ing animal life: Cytochrome c oxidase subunit 1 divergences

among closely related species. Proceedings of the Royal So-

ciety of London B 270: S596—S599

Kikvidze Z, Ohsawa M (1999) Adjara, East Mediterranean

refuge of Tertiary vegetation. Pp. 297-315 in: Ohsawa M,

Wildpret W, Arco MD (eds) Anaga Cloud Forest, a compar-

ative study on evergreen broad-leaved forests and trees of

the Canary Islands and Japan. Chiba University Publications

Milne RI, Abbott RJ (2002) The origin and evolution of Tertia-

ry relict floras. Advances in Botanical Research 38: 281-314

Mittermeier RA, Gil PG, Hoffmann M, Pilgrim J, Brooks T,

Mittermaier CG, Lamoreux J, da Fonseca GAB (2004)

Hotspots Revisited: Earth’s Biologically Richest and Most

Endangered Terrestrial Ecoregions. CEMEX/Agrupacion,

Sierra Madre

Moriniére J, Hendrich L, Balke M, Beermann AJ, Konig T,

Hess M, Koch S, Miller R, Leese F, Hebert PDN, Haus-

©ZFMK

286 Jana Thormann et al.

mann A, Schubart CD, Haszprunar G (2017) A DNA

barcode library for Germany's mayflies, stoneflies and

caddisflies (Ephemeroptera, Plecoptera and _ Trichop-

tera). Molecular Ecology Resources 17: 1293-1307.

https://doi.org/10.1111/1755-0998 12683

Mumladze L, Japoshvili B, Anderson EP (2019) Fau-

nal Biodiversity Research in the Republic of Georgia: A

Short Review of Trends, Gaps, and Needs in the Cauca-

sus Biodiversity Hotspot. Preprints 2019, 2019080249.

https://doi.org/10.20944/preprints201908.0249.v1

Myers N, Mittermeier RA, Mittermeier CG, da Fonseca GAB,

Kent J (2000) Biodiversity hotspots for conservation priori-

ties. Nature 403: 853-858

Neidze V (2003) Sakartvelos sotsialur-ekonomikuri geogra-

fia (Social-economic geography of Georgia). Metsniereba,

Tbilisi, Georgia

Raupach MJ, Hendrich L, Kichler SM, Deister F, Moriniere J,

Gossner MM (2014) Building-Up of a DNA Barcode Library

for True Bugs (Insecta: Hemiptera: Heteroptera) of Germany

Reveals Taxonomic Uncertainties and Surprises. PLoS ONE

9(9): 106940. https://doi.org/10.1371/journal.pone.0106940

Rulik B, Eberle J, von der Mark L, Thormann J, Jung M,

Kohler F, Apfel W, Weigel A, Kopetz A, Kohler J, Fritzlar

F, Hartmann M, Hadulla K, Schmidt J, Hérren T, Krebs

D, Theves F, Eulitz U, Skale A, Rohwedder D, Kleeberg

APPENDIX II.

Table 1. Collecting sites of 2018, Georgia.

Locality description

1 km N of Bakhmaro

Barkhmaro

Black Sea coast near Grigoleti in front of Ilia University field station

Borjomi, pipeline road between Sakire and Tsikhisjvari, 0.65 km V of Kodiani, meadow and

open pine tree forest with large amounts of perennial herbs

Borjomi, pipeline road between Sakire and Tsikhisjvari, 1.75 km W of Kodiani, drier meadow

with flowers

Borjomi, pipeline road between Sakire and Tsikhisjvari, 2.3 km E of Sakire, dryish meadow

with some wetter areas

Borjomi, pipeline road between Sakire and Tsikhisjvari, 2.9 km E of Kodiani, very diverse

meadows

Bugdasheni river, near small Arakali

Chkhakoura — Bakhmaro road

Chkhakoura — Bakhmaro road, 2.9 km ENE of Bakhmaro, overgrazed meadows

Ghebi

Grassy peak 1 km N of Bakhmaro

Grigoleti, Ilia State Universtity Marine Biological Station

Iori River, down from the Paldo Dam

Khino , entrance

Kintrishi

Kintrishi drainage: Kintrishi River downstream of old bridge

Kintrishi drainage: Kintrishi River in Kobuleti Village, upstream of railway bridge

Kintrishi drainage: Kintrishi River near village Kveda Sameba

Bonn zoological Bulletin 68 (2): 275-296

A, Astrin JJ, Geiger MF, Wagele JW, Grobe P, Ahrens D

(2017) Using taxonomic consistency with semi-automat-

ed data pre-processing for high quality DNA barcodes.

Methods in Ecology and Evolution 8(12): 1878-1887.

https://doi.org/10.1111/2041-210X.12824

Shatilova I, Mchedlishvili N, Rukhadze L, Kvavadze E (2011)

The History of the Flora and Vegetation of Georgia (South

Caucasus). Georgian National Museum, Institute of Paleo-

biology, Tbilisi

Tarkhnishvili D, Gavashelishvili A, Mumladze L (2011) Palae-

oclimatic models help to understand current distribution of

Caucasian forest species. Biological Journal of the Linnean

Society 105(1): 231 —248

Wesener T, Voigtlander K, Decker P, Oeyen JP, Spel-

da J, Lindner N (2015) First results of the German Bar-

code of Life (GBOL) — Myriapoda project: Cryptic lin-

eages in German Stenotaenia linearis (Koch, 1835)

(Chilopoda, Geophilomorpha). ZooKeys 510: 15-29.

https://doi.org/10.3897/zookeys.510.10176

Zazanashvili N, Sanadiradze G, Bukhnikashvili A, Kandaurov

A, Tarkhnishvili D (2004) Caucasus. Pp. 148—153 in: Mitter-

meier RA, Gil PG, Hoffmann M, Pilgrim J, Brooks T, Mit-

termaier CG, Lamoreux J, da Fonseca GAB (eds) Hotspots

revisited, Earth’s biologically richest and most endangered

terrestrial ecoregions. CEMEX/Agrupacion, Sierra Madre.

Latitude Longitude Altitude Collection date

41.8576 42.3237 2055 m 19.07.18

41.8576 42.3240 2050 m 19.07.18

42.0535 41.7260 19.07.18

41.7271 43.3773 1863 m 22.07.18

41.7323 43.3221 1772 m 23.07.18

41.7325 43.3068 1552 m 22.07.18

41.7272 43.377] 2190 m 23.07.18

41.2356 43.6720 01.11.18

41.8607 42.3574 1935 m 19.07.18

41.8610 42.3569 1900 m 19.07.18

42.7774 43.4956 1410 m 17.07.18

41.8611 42.3237 2192 m 19.07.18

42.0529 41.7272 Om 18.—20.07.18

41.8465 45.1357 23.06.18

41.7637 41.9788 385m 27.07.18

41.7369 41.9839 21.07.18

41.7520 41.9760 451m 21.07.18

41.7880 41.9630 21.07.18

41.7370 41.9810 21.07.18

41.7889 41.9603 21.07.18

41.8033 41.7778 21.07.18

41.7991 41.8480 21.07.18

©ZFMK

A prelude to the CaBOL Platform: Biodiversity Days in Georgia in 2018 and 2019

Table 1. Continued.

Locality description

Kintrishi drainage: Kintrishi River SE of Khutsubani Village

Kintrishi drainage: Kintrishi River, upstream of gravel quarry E oft Kobuleti

Kintrishi National Park, at bisection behind ranger station; under bark and dead wood rela-

tively close to the river; humid leaf litter

Kintrishi Nature Reserve

Kintrishi Nature Reserve, Khino

Kintrishi NR

Kintrishi NR, Khino monestry

Kintrishi NR, picnic spot

Kintrishi Protected Areas

Kintrishi ProtectedAareas, Didvake

Kintrishi Protected Areas, Didvake Village

Kintrishi Protected Areas, monastery at Khino, meadows

Kintrishi Protected Areas, monastery at Khino, meadows, meadow — pine forest mosaic

Kintrishi River, side valley

Kintrishi, Khino

Kintrishi, National Park

Kintrishi-National Park, woods and meadows around Khino Monastery

Kintrishi-National Park, wayside between Didvake and Khino along Kintrishi River

Near Lentekhi, antennae

North from Atsana, along the road

Above Khino Church

Above Khino Church, second meadow

Paliastomi-Rioni drainage: Oragvisghele Stream SW of Lanchkhuti Village

Paliastomi-Rioni drainage: Swlana Stream S of Jurukveti Village, upstream of 02-18

Paliastomi-Rioni drainage: Swlana Stream W of Jurukveti Village

Paliastomi-Rioni drainage: Swlana Stream W of Jurukveti Village, upstream of 02-18

Rioni drainage: Gubistskali River E of Samtredia Town, above confluence with Rioni

River and Rioni Channel

Road from Sakire to Tsikhisjvari

Road to Barkhmaro

Road to Chkhakoura - Bakhmaro road, 5 km NE of Bakhmaro, meadows on ridge

along road

Road to Khino, Kintrishi Protected Areas, 6 km WNW of monastery at Khino, sunny,

exposed rock face along road

Road to Tsana

Samtskhe-Javakheti, road from Sakire to Tsikhisjvari

Tsana

Tskhmori, Chalistskali Waterfall

Below Khino, meadow

Western Meskheti Mountains

Bonn zoological Bulletin 68 (2): 275-296

Latitude

41.8029

41.8016

41.7303

41.7345

41.7380

41.7345

41.7369

41.7286

41.7319

41.7621

41.7366

41.7349

41.7463

41.7172

41.7287

42.7752

41.7346

41.7287

41.7440

41.7169

41.7173

42.7774

42.0559

41.7283

41.7344

42.0850

42.0654

42.0783

42.0730

42.1625

41.7303

41.7271

41.7326

41.8609

41.8863

41.7376

42.8739

41.7303

42.8889

42.5313

41.7195

41.7553

Longitude Altitude

41.8384

41.8095

41.9817

41.9888

42.0076

41.9888

41.9839

42.0781

41.9753

41.9785

41.9787

41.9710

42.0167

42.0467

42.0782

42.7608

41.9758

42.0782

42.0832

42.0552

42.0469

42.7502

42.0594

42.0841

42.0845

42.0246

41.9209

41.9164

41.9168

42.4107

43.3348

43.3773

43.3081

42.3572

42.3614

42.3612

42.0074

43.1500

43.3348

43.1430

43.4709

42.0408

42.1112

287

Collection date

20.07.18

22.07.18

20.07.18

18.07.18

20.07.18

29.07.18

21.07.18

19.07.18

18.07.18

17.07.18

20.07.18

25.07.18

19.07.18

21.07.18

23.07.18

18.07.18

25.07.18

19.07.18

23.07.18

20.07.18

19.07.18

24.07.18

19.07.18

24.07.18

22.07.18

19.07.18

18.07.18

24.07.18

23.07.18

24.07.18

18.07.18

09.08.18

©ZFMK

288 Jana Thormann et al.

Table 1. Continued.

Locality description Latitude 41.9782 Altitude Collection date

Malaise Trap, ranger station Kintrishi 41.7621 41.9785 318m Apr—Nov. 2018

Malaise Trap Grigoleti field station 42.0528 41.7269 7m 1725.04.18

, i 20.04.-

Malaise Trap Transect Kintrishi #1 41.7294 42.0776 1035 m 05.11.2018

Malaise Trap Transect Kintrishi #2 41.7294 42.0775 1020m_— Oh

7 3 05.11.2018

Malaise Trap Transect Kintrishi #3 ange 420773; 103m a pP re

: : 05.11.2018

Malaise Trap Transect Kintrishi #4 41.7441 42.0834 1264m_ 20.08

7 | 05.11.2018

Malaise Trap Transect Kintrishi #5 41.7434 42.0818 »«1252m_— 0h

“ : 3 05.11.2018

Malaise Trap Transect Kintrishi #6 41.7433 42.0841 1235 m eee

: : 05.11.2018

Malaise Trap Transect Kintrishi #7 41.7371 41.9792 404m oO Ute

: : 05.11.2018

Malaise Trap Transect Kintrishi #8 41.7378 41.9786 403m eta

: : 05.11.2018

Malaise Trap Transect Kintrishi #9 41.7378 41.9782 401m gaat

7 : 05.11.2018

Malaise Trap Transect Kintrishi #10 41.7479 42.0959 1697m Oh

: : 05.10.2018

Malaise Trap Transect Kintrishi #11 41.7477 42.0951 1637m Oh

7 : 05.10.2018

Malaise Trap Transect Kintrishi #12 41.7482 42.0940 1634m

: : 05.10.2018

Malaise Trap Transect Kintrishi #13 41.7552 42.1125 268m Oh

, | 05.10.2018

Malaise Trap Transect Kintrishi #14 41.7553 42.1128 2280 m Agee

: : 05.10.2018

Malaise Trap Transect Kintrishi #15 41.7554 42.1129 280m Oh

7 ; 05.10.2018

Malaise Trap Transect Kintrishi #16 41.7625 42.1157 465m OH

: : 05.10.2018

Malaise Trap Transect Kintrishi #17 41.7619 42.1162 2462m 9 24 0b

: : 7 05.10.2018

Malaise Trap Transect Kintrishi #18 41.7616 42.1158 2450m 4h

, : : 05.10.2018

A prelude to the CaBOL Platform: Biodiversity Days in Georgia in 2018 and 2019 289

APPENDIX ILI.

Table 2. Collecting sites of 2019, Georgia.

Locality description Latitude Longitude Altitude Collection date

1.4 km WSW IImazlo 41.4240 45.0080 - 30.06.2019

10 km S. of Kasristskali, Vashlovani NR 41.2235 46.5353 390 m 10.07.2019

14 km E of Mestia 43.0254 42.8907 2550 m 13.06.2019

14.5 km E of Mestia, forest with Rhododendron 43.0284 42.8788 2345 m 13.06.2019

15 km E of Mestia 43.0266 42.9101 2860 m 13.06.2019

2 lakes between Amali River and Terek 42.7219 44.6225 1538 m 09.07.2019

2.8 km nnw Jikurebi Lake 41.5980 45.3260 - 01.07.2019

20 km S of Stepandsminda 42.5105 44.4949 2550 m 1.-7.7.2019

5 km W of Naduknari 42.0640 45.0636 1045 m 08.07.2019

7 km W of Ushguli, along the road 42 91:79 42.9366 2290 m 16.06.2019

7 km W of Ushguli, hilltop 42.9062 42.9370 2615 m 16.06.2019

7.5 km N of Mestia, path to glacier 43.1017 42.7276 1850 m 14.06.2019

7.5 km N of Mestia, path to glacier 43.1133 42.7379 1800 m 14.06.2019

7.7 km NE Mele, meadow 42.8211 43.1607 1450 m 20.06.2019

Abastumani area, along path 41.8234 42.8400 2025 m 11.06.2019

Akavreta river, left tributary of Adjaristskali River 41.5821 41.9679 - 01.06.2019

Akhaldaba northeast of Borjomi, Nedzvi Sanctuary 41.9028 43.5167 890 m 15.07.2019

Akhaltsikhe, Snotskali River valley 42.5881 44.6667 1802 m 10.07.2019

Akhaltsikhe-Sno, Snotskali River valley 42.5997 44.6528 1780 m 10.07.2019

Aktas Golu 41.2360 43.2490 — 11.07.2019

Alazani Chanal, near Tibaani 41.6016 46.0139 — 22.05.2019

Alazani River, down from the Shilda 41.9351 45.6891 — 25.07.2019

Alazani River, tributary of Kabali — Alazani, Tsnori bridge 41.6807 46.0756 = 22.05.2019

Algeti River 0.8 km WNW Tskhrakudaani 41.6750 44 3790 - 10.07.2019

Algeti River 3.4 km NW Abrameti 41.6340 44.4690 — 13.07.2019

Algeti River N Partskhisi 41.5790 44.5670 — 13.07.2019

Almasiani, southern end of village 42.5544 44.4969 1990 m 07.07.2019

ce ia aa Dariali Monastery complex, 10 km N of Stepanzminda; 42.7360 44 634] 1350 m Le a

et a = Gveleti big waterfalls, rich, variable meadows, scree; Kazbegi 42.7053 446149 1450-1700m ates 6

Alpana towards Kutaisi, Rioni Valley, limestone cliff along road 42.5581 42.8206 380 m 02.07.2019

Alpana towards Kutaisi, Rioni Valley, right side, north of Mekvena 42.4914 42.7839 370m 02.07.2019

Sie towards Kutaisi, Rioni Valley, southern entrance of canyon-like part of 42.5219 42.7942 380 m 02.07.2019

Alpana, limestone cliffs on left side of Rioni Valley 42.5592 42.8486 420m 02.07.2019

Ananuri towards Zhinvali Dam, ca 4.5 km along road north of the dam 42.1525 44.7531 940 m 03.07.2019

Ananuri, slope opposite monastery, beyond bridge 42.1606 44.7028 880 m 03.07.2019

Aragvi River NE Zhinvali 42.1120 44.7780 - 07.07.2019

Aragvi River W Bulachauri 42.0360 44.7460 — 07.07.2019

Aragvi River W Choporti 41.9730 44.7560 — 07.07.2019

Aragvi Valley, between junction towards Denali and Tsivtskaro 42.2175 44.6728 910m 07.07.2019

Aragvi Valley, Bibiliani, southern end of village 42.3244 44.6814 1050 m 07.07.2019

Bakhmaro 41.8600 42.3200 2000 m 03.08.2019

Batumi, Botanical Garden 41.6990 41.7220 - 19.07.2019

290 Jana Thormann et al.

Table 2. Continued.

Locality description Latitude Longitude Altitude Collection date

Mountain station at Kobi 42.5112 44.4929 2957 m 07.07.2019

Between Kanobi and Pkhelshe along river Kesia 42.5980 44.5410 - 07.07.2019

Between Kvishkheti and Khashuri 41.9625 43.5636 740 m 08.06.2019

Between Tianeti and Akhmeta 42.0640 45.0640 1000 m 08.07.2019

Between Tskneti and Kojori 41.6700 44.6690 - 15.07.2019

Blashoviskhevi River SW Norio 41.7800 44.9700 — 02.07.2019

Borjomi 41.8320 43.3820 - 28.07.2019

aa pes! = Gveleti waterfalls, fast-flowing, rocky stream with a lot of 42.7074 44.6247 1493 m Behe, eh

Bughdasheni Lake 41.1980 43.6890 ~ 12.07.2019

Bursa Stream, tributary to Alasani River about 3 km S of Sanavardo 41.8670 45.8100 - 08.07.2019

Chargali, Pschawi-Aragwi 42.3342 44.9031 1029 m 05.07.2019

Charnali, Sarpi 41.5560 41.6110 — 19.07.2019

Chili-Chili River in Beshtasheni 41.6410 44.1090 — 13.07.2019

Chkheri River, Stepanzminda 42.6712 44.6123 2000 m 02.—03.07.2019

Chkheri Valley northwest of Stepanzminda 42.6708 44.6170 1960 m 05.07.2019

Chkheri Valley, right side 42.6706 44.6097 2030 m 04.07.2019

Chrami River N Tikilisa 41.5970 43.9600 - 13.07.2019

Cow and horse pasture W of Stepandsminda 42.6668 44.6303 1812m 1-7.7.2019

Daliari Valley 42.7371 44.6318 1319m 08.07.2019

Dariali gorge 42.7030 44.6270 — 05.07.2019

eae nee ee of the monastery complex of the Holy Archangels 42.7362 44 6333 1340 m

Debeda River E Kirach-Mughanlo 41.3340 45.0680 — 30.06.2019

Debeda River N Khanji-Gazlo 41.3570 45.0050 -- 30.06.2019

Debeda River N Kirach-Mughanlo 41.3400 45.0510 — 30.06.2019

Debeda River W Didi Mughanlo 41.3890 44.9430 _ 30.06.2019

so eg tat road to Eagle Canyon, very dry meadow dominated by Eryngi- 41.4875 46.0944 770 m 10.07.2019

Enguri Dam 42.8100 42.0450 — 04.08.2019

Entrance Vashlovani Nationalpark 41.2143 46.5368 380 m 10.07.2019

Fluvial plane Tergi = Terek 42.6532 44 6356 1733 m 04.07.2019

Forest 42.0644 45.0635 1059 m 08.07.2019

Former road, tunnel 42.6900 44.6357 1633 m 08.07.2019

Gelati Monastery, walls and stones along path on the southside of the monastery 42,2939 42.7683 420m 01.07.2019

Gergeti Sameba, valley west of church 42.6628 44.6106 2150 m 04.07.2019

Gergeti, close to road to Gergeti Trinity Church 42.6669 44.6128 2175m 13.07.2019

Gergeti, Ghkheri Valley 42.6700 44.6103 2103 m 13.07.2019

Gergeti, Ghkheri Valley 42.6708 44.6119 1900 m 18.07.2019

Gergeti, north of village 42.6669 44.6303 1830 m 04.07.2019

Gergeti, road to Gergeti Trinity Church 42.6720 44.6100 - 04.07.2019

Gergeti, valley S of Gergeti Trinity Church 42.6617 44.6156 2080 m 13.07.2019

Gergeti Trinity Church parking spot 42.6650 44.6143 2257-m 1.-—7.7.2019

Goderdzi Pass 41.6330 42.5010 ~ 20.07.2019

Goderdzi Pass 41.6610 42.6070 — 19.07.2019

A prelude to the CaBOL Platform: Biodiversity Days in Georgia in 2018 and 2019 291

Table 2. Continued.

Locality description Latitude Longitude Altitude Collection date

Gori Castle, southern side of castle hill 41.9856 44.1086 620 m 03.07.2019

Gori, Stalin Park 41.9867 44.1136 600 m 03.07.2019

Grigoleti 42.0530 41.7280 i Lata 9

Group of trees near Vashlovani NP 41.2524 46.5188 465 m 10.07.2019

Gudauri, lift station 42.5105 44.4954 3000 m 07.07.2019

Gudauri Pass, at Tufa formations 42.5336 44.4750 2210-2230m 11.07.2019

Gveleti 42.7071 44.6254 1500 m 04.07.2019

Gveleti 42.7215 44.6225 1900 m 09.07.2019

Gveleti Lakes; Kazbegi Municipality 42.7227 44.6238 -

Gveleti north of Stepanzminda slopes near the Great waterfall 42.7047 44.6184 1620 m pies ae

Gveleti north of Stepanzminda valley below the waterfalls 42.7061 44.6224 1520 m noe oie

Gveleti Waterfalls 42.7044 44.6205 — 1. -7.7.2019

Gveleti Waterfalls 42.7070 44.6252 1483 m 04.07.2019

Gveleti Waterfalls 42.7078 44.6253 1478 m 12.07.2019

Gveleti, near small waterfall 42.7025 44.6194 1645 m 17.07.2019

Gveleti, small waterfall 42.7050 44.6150 1600 m 06.07.2019

Gveleti, way to waterfall 42.7060 44.6170 1700 m 06.07.2019

Gveleti Valley 42.7074 44.6247 1485 m 02.07.2019

Heretiskari, way to Sighnaghi 41.7110 46.0870 200 m 09.07.2019

Tori River 8.5 km SE Sagaredscho 41.6680 45.3880 - 01.07.2019

Tori River N Qaracop 41.6130 45.5390 - 01.07.2019

Tori River NE Sartichala 41.7230 45.1810 - 01.07.2019

Jandara Reservoir 1.6 km SE Jandari 41.4350 45.1870 - 02.07.2019

Jandara Reservoir 2.8 km SE Mzianeti 41.4510 45.2120 - 02.07.2019

Jandara Reservoir 4.7 km SE Mzianeti 41.4400 45.2280 - 02.07.2019

ate ce a valley and Juta Valley, valley bottom, rich in succulents; 42.5627 44.7070 1830 m 05.07.2019

Juta 42.5720 44.7310 ~ 05.07.2019

Juta 42.5797 44.7431 2160 m 05.07.2019

Juta at Sno-Valley, village 42.5795 44.7459 - 05.07.2019

Juta Valley, Juta mountainside with rich meadows; Kazbegi Municipality 42.5800 44.7423 2100-2200 m 05.07.2019

Juta, brook above the village 42.5839 44.7486 2185 m 10.07.2019

Juta, slope 42.5796 44.7436 2148 m 09.07.2019

Juta, village border 42.5794 44.7433 2150 m 10.07.2019

Jutistskali River 42.5838 44.7484 2182 m 10.07.2019

Jutistskali Valley, right side, near confluence with Kora River 42.5622 44.7061 1860 m 05.07.2019

Jvari Monastery, along road east of the monastery 41.8403 44.7367 600 m 01.07.2019

Jvari Pass 42.5189 44.4669 2360 m 07.07.2019

Kapatadze Lake 41.5730 45.3220 ~ 01.07.2019

Kasbek Mountains 5.6 km W Stepanzminda 42.6590 44.5760 - 05.07.2019

Keda, Kveda Agara 41.6020 41.9020 - 19.07.2019

Khde-riverbed above Dariali-Monastery-complex 42.7360 44.6342 1360 m 02.07.2019

Khino, Kintrishi 41.7290 42.0780 = 30.—31.07.2019

292 Jana Thormann et al.

Table 2. Continued.

Locality description Latitude Longitude Altitude Collection date

Khomisdziri, Pschawi-Aragwi 42.3880 44.9185 1070 m 05.07.2019

Khornabuji 41.4724 46.0847 780 m 10.07.2019

Pieced Dacictel cog tall-herbs and mountainside with shorter vegetation; 42.5994 44.5444 1980-2200m 07.07.2019

Kintrishi 41.7490 42.1000 ~ 31.07.2019

Kirkhbulaki River e Qulalisi 41.3270 43.4840 - 11.07.2019

Kobi, near Kobi Pass 42.4983 44.493] 2760 m 14.07.2019

Kobi, near Kobi Pass 42.5047 44.493] 2860 m 14.07.2019

Kobi, near ski lift base station 42.5572 44 4975 1825m 14.07.2019

Kobi Valley/ river Tergi= Terek 42.5809 44.4642 2009 m 07.07.2019

Kochki River S Epremovka 41.1890 43.7480 - 12.07.2019

Korolistavi, Mtirala 41.6420 41.7430 - 18.07.2019

Ksani river, near Mukhrani Village 41.9628 44.5265 - 30.06.2019

Kumisi Lake, S of Tbilisi 41.5770 44 8240 ~ 16.07.2019

Kura drainage: Aragvi River near Naoza 41.9720 44.7550 - 07.07.2019

ae Seger Pshavis Aragvi River near Tsiprani Village, upstream of Zhinvali 42.2340 44 8450 - 05.07.2019

ao Gees ae tributary stream to Pshavis Aragvi River about 42.3870 44.9180 - 05.07.2019

Kura River 1.2 kmNn Metekhi 41.9340 44 3420 ~ 09.07.2019

Kura River 1.3 km NNW Tedotsminda 42.0350 44 0620 - 08.07.2019

Kura River 1.5 km W Khtsisi 41.9800 43.6550 - 08.07.2019

Kura River 1.6 km E Ilmazo 41.4280 45.0430 - 30.06.2019

Kura River 1.9 km ESE Ilmazo 41.4180 45.0420 — 30.06.2019

Kura River 2 km S Karayalari 41.5990 44.9600 - 29.06.2019

Kura River 2.0 km ESE Ilmazo 41.4200 45.0440 - 30.06.2019

Kura River 2.3 km NNE Teliani 41.9480 44.2820 - 09.07.2019

Kura River 2.3 km W Mtskheta 41.8390 44 6800 - 07.07.2019

Kura River 6.6 km E Khidistavi 41.9600 44.2100 — 09.07.2019

Kura River 7.5 km E Khidistavi 41.9650 44.2200 — 09.07.2019

Kura River E Variani 42.0730 44 0400 ~ 08.07.2019

Kura River N Akhalsheni 42.0050 43.7230 ~ 08.07.2019

Kura River NW Akhalsopeli 42.0130 43.7650 - 08.07.2019

Kura River NW Dzegvi 41.8500 44 5990 - 07.07.2019

Kura River SE Gachiani 41.5780 44.9990 - 29.06.2019

Kura River SE Gori 41.9710 44.1210 — 09.07.2019

Kura River, Rustawi 41.5510 45.0100 — 29.06.2019

Kura valley 2.3 km W Akhalsheni 41.4870 45.0380 ~ 29.06.2019

Kura valley 2.7 km W Akhalsheni 41.4840 45.0340 - 29.06.2019

Kura valley SE Rustawi 41.5200 45.0230 — 29.06.2019

Kutaisi, Bagrati Cathedral 42.2775 42.7050 210m 02.07.2019

Kutaisi, Botanical Garden 42.2792 42.7097 160 m 01.07.2019

Kveda Chkhutuneti 41.5020 41.8510 ~ 19.07.2019

lakes 3.4 km N Tsdo 42.7160 44 6240 _ 03.07.2019

Martini Canal 3.4 km N Jandari 41.4730 45.1670 - 02.07.2019

A prelude to the CaBOL Platform: Biodiversity Days in Georgia in 2018 and 2019 293

Table 2. Continued.

Locality description Latitude Longitude Altitude Collection date

Meadows with basaltic rocks; Kazbegi Municipality 42.5797 44.4700 2000-2300 m 07.07.2019

Mineral Springs, Baidara Valley north of Jvari Pass 42.5319 44.4722 2250 m 05.07.2019

Middle station, Kobi 42.5317 44.4934 2513 m 07.07.2019

Mna-Valley, W of Shevardeni 42.5800 44.4700 - 07.07.2019

Monastery Church 42.6648 44.6148 2169 m 03.07.2019

Monestry N of Stepandsminda 42.7362 44 6330 1899 m 1.-7.7.2019

Mountain pasture at road between Achmeta and Tianeti 42.0635 45.0641 1030 m 08.07.2019

Nadarbazevis Tba 41.9990 44.2870 - 10.07.2019

Nakerala Pass 42.3758 43.0372 1230 m 02.07.2019

Nakhshirgele towards Kutaisi, ca 200 m east of bridge over Tchishura River 42.2150 42.7983 130m 01.07.2019

Nakhshirgele towards Kutaisi, ca 500 m along road to Broloskedi 42.2300 42.7908 150m 01.07.2019

Narvani River, Kobi 42.5616 44.5102 1959 m 04.07.2019

Navenakhevi, forest near Navenakhevi Cave 42.2461 42.8469 300 m 02.07.2019

Near to Lagodechi NP 41.8335 46.2825 550 m 09.07.2019

Near Tsana, along the road 42.8887 43.1429 1757 m 18.06.2019

Near Tsana, along the road 42.9012 43.1422 1835 m 18.06.2019

Near Tsana, along the road 42.9160 43.1428 1975 m 19.06.2019

Near Tsana, meadow 42.8889 43.1430 1760 m 18.06.2019

Near Ushguli, close to river Inguri 42.9499 43.0719 2270 m 15.06.2019

Near Ushguli, path to glacier 42.9437 43.0539 2220 m 15.-17.06.2019

Near Ushguli, up to the ruins 42.9101 43.0070 2295 m 17.06.2019

Near Vashlovani NP 41.2636 46.5261 ~ 10.07.2019

NW of Stepanzminda 42.6710 44.6100 2050 m 04.07.2019

NW of Stepanzminda 42.6740 44.6330 2200 m 01.—07.07.2019

On the way to Sighnaghi, village Heretiskari, oak grove 41.7106 46.0870 215m -

on the way to Telawi 42.0605 45.0617 1555 m =

E of Sno, alluvial forest 42.5996 44.6531 1781 m 10.07.2019

Ozero Zres 41.3880 43.4230 — 11.07.2019

Pass from Ushguli to Tsana 42.9140 43.0911 2575 m 18.06.2019

Pastures | km E Stepanzminda 42.6580 44.6600 — 06.07.2019

Pastures around Gergeti Trinity Church 42.6650 44.6150 - 06.07.2019

Path from Chkheri riverbed to road to Gergeti Trinity Church 42.6694 44.6119 steel 04.07.2019

Plain at Akhaltsikhe 42.5889 44.6657 1802 m 09.07.2019

Poti, Paliastomi Lake 42.1150 41.7030 - 29.07.2019

Rioni Valley, ca 1 km along road towards Ambrolauri 42.5489 42.8558 440m 02.07.2019

Riverbed at Terek between Achkhoti and Stepanzminda 42.6359 44.6285 1755 m 05.07.2019

Riverside W of Stepandsminda 42.6707 44.6097 2044 m 1.-7.7.2019

Road from Abastumani to Sairme 41.7963 42.8444 1700 m HSS

Road from Abastumani to Sairme 41.7969 42.8439 1725 m 10.-11.06.2019

Road from Abastumani to Sairme 41.8385 42.8194 2260 m 10.06.2019

Road from Abastumani to Sairme 41.8643 42.7784 1830 m a

294 Jana Thormann et al.

Table 2. Continued.

Locality description Latitude Longitude Altitude Collection date

Road from Abastumani to Sairme, near river 41.7773 42.8372 1386 m 10.-11.06.2019

Road from Abastumani to Sairme, small creek 41.8638 42.7890 1800 m 11.06.2019

Road from Sakire to Tsikhisjvari 41.7305 43.3343 1900 m 09.06.2019

Road from Sakire to Tsikhisjvari, hilltop and surroundings 41.7250 43.3606 2519 m _

Road to Gergeti Trinity Chruch 42.6670 44.6140 2100 m 01.—07.07.2019

Road to Gergeti Trinity Chruch 42.6730 44.6160 2250 m 01.-07.07.2019

Road to Vashlovani National Park, 7km NW of gate, dry steppe 41.2742 46.5146 450m 10.07.2019

Rustawi, lake close to Kura River 41.5490 45.0020 _ 29.06.2019

S of Lakhushdi, meadow 42.9988 42.6501 1270 m 13.-14.06.2019

S of Lakhushdi, meadow 42.9990 42.6502 1270 m 13.-14.06.2019

S of Sioni 42.5960 44.5700 1900 m 07.07.2019

S of Stepanzminda 42.6370 44.6310 - 05.07.2019

Saguramo 41.8944 44.7521 551m 21.06.2019

Satsire, southern outskirts of village 42.3350 42.9300 560 m 02.07.2019

Scrub and undergrowth near Sighnaghi 41.6128 45.9329 660 m —

Sheep and horse pasture 42.6504 44.6514 1861 m 1-7.7.2019

Shevardini > Truso Gorge 42.5914 44.4378 2100 m 15.07.2019

Shevardini > Truso Gorge, near Tufa formations 42.5828 44 4303 2132m 15.07.2019

Side valley (Kora River) of Sno Valley 42.5637 44.7074 1845 m 05.07.2019

Sighnaghi 41.6210 45.9180 — 09.07.2019

Sighnaghi, surroundings of “Best Host” 41.6213 45.9180 727 m 10.-11.07.2019

Sioni 42.5963 44.5702 2000 m 07.07.2019

Slope at Chkheri riverbed near Gergeti Trinity Church 42.6722 44.6114 2070 m 04.07.2019

Small lake 2.0 km ENE Patara Gondra 41.3070 43.4750 - 11.07.2019

Small lake 2.2 km ENE Imera 41.6500 44.2150 ~ 10.07.2019

Small lake N Sulda 41.2820 43.3720 _ 11.07.2019

Small oakwood 41.7102 46.0876 229 m 10.07.2019

Small river 1.6 km WSW Tokhliauri 41.7210 45.4030 ~ 01.07.2019

Small river valley SW Manglisi 41.6940 44.3790 - 10.07.2019

Small valley 1.1 km NE Abrameti 41.6200 44.5080 — 13.07.2019

Small valley 4.8 km SE Giorgitsminda 41.6960 45.3830 — 01.07.2019

Small valley near Kanobi at Kesia River 42.5975 44.5406 1925 m 07.07.2019

Sno, Snotskali River valley 42.5989 44.6500 1780 m 11.07.2019

Snostskali River 0.6 km NW Sno 42.6090 44.6330 - 04.07.2019

Snostskali River 0.6 km SE Achkhoti 42.6180 44.6240 ~ 03.07.2019

Snostskali River 0.8 km SE Sno 42.6000 44.6450 - 04.07.2019

Snostskali River SE Akhaltsikhe 42.5880 44.6670 — 04.07.2019

Soramula River 1.7 km ENE Agara 42.0470 43.8410 - 08.07.2019

Stepantsminda, station of Ilia State University 42.6550 44.6492 1830 m 04.07.2019

Stepanzminda 42.6530 44.6550 1800 m 02.07.2019

Stepanzminda 42.6535 44 6496 1823 m 03.—06.07.2019

Stepanzminda 42.6549 44.6495 1900 m 1 —7.7.2019

Stepanzminda 42.6581 44 6566 1900 m 06.07.2019

Stepanzminda 42.6641 44.6144 2163 m 07.07.2019

Stepanzminda 42.6656 44.6365 1797 m 03.07.2019

A prelude to the CaBOL Platform: Biodiversity Days in Georgia in 2018 and 2019 295

Table 2. Continued.

Locality description Latitude Longitude Altitude Collection date

Stepanzminda 42.6704 44.6096 2034 m 03.07.2019

Stepanzminda, along road to loane Natlismcemeli Orthodox Church, overgrazed

meadows with Hippophae rhamnoides, Kazbegi Municipality secees ee ere ee

Stepanzminda, cow and horse pasture W of Stepandsminda 42.6678 44.6316 1810m —

Stepanzminda, Elm Monastery 42.6575 44.6567 1950 m 17.07.2019

Stepanzminda, Gveleti 42.7161 44.6239 1400 m 09.07.2019

Stepanzminda, Gveleti, Tibistskali River 42.7047 44.6208 1540 m 12.07.2019

Stepanzminda, Gveleti, Tibistskali River 42.7078 44.6253 1480 m 12.07.2019

Stepanzminda, hilltop north of Gergeti Trinity Church 42.6753 44.6158 2250 m _

Stepanzminda, Ilia State University field station 42.6548 44.6490 1820 m 12.07.2019

Stepanzminda, Kasbek mountain 42.6667 44.6008 2470 m 18.07.2019

Stepanzminda, near Snotskali River mouth 42.6367 44.6308 1750 m 16.07.2019

Stepanzminda, near village 42.6628 44.6394 1745 m 16.07.2019

Stepanzminda, parking bay on the way to the Gergeti Trinity Church 42.6669 44.6136 2120 m —

Stepanzminda, pass to Gergeti Glacier 42.6588 44.5736 2900 m 05.07.2019

Stepanzminda, road to Gergeti Trinity Church 42.6678 44.6106 2160 m 18.07.2019

Stepanzminda, slope opposite pastures 42.6672 44.6292 1825 m -

Stepanzminda, Snotskali River mouth 42.6369 44.6325 1760 m 16.07.2019

Pace Meee with some drier areas, flower-rich 42.6755 44 6130 awe 04.07.2019

Stepanzminda, Tergi River 42.6519 44.6392 1740 m 15.07.2019

Stepanzminda, way to hilltop north of Gergeti Trinity Church 42.6725 44.6161 1900-2250m -

Streamcourse 42.6711 44.6118 1996 m 09.07.2019

Streamcourse to monastery 42.6706 44 6096 2023 m 09.07.2019

Street to Baisubani 41.8399 46.1121 300 m 09.07.2019

Sulfur springs at Dschari Pass 42.5346 44.4756 2216m 11.07.2019

Surami Pass, eastern side 42.0472 43.4981 910m 03.07.2019

Tabatskuri 41.6600 43.6130 - 23.07.2019

Coombe, river Tegi= Terek 42.6843 44 6362 1594 m 08.07.2019

Valley station, Kobi 42.5573 44.4976 1988 m 07.07.2019

Tba Bazalet’i 42.0440 44.6810 — 07.07.2019

Tba Khanjali 41.2390 43.5610 — 10.07.2019

Tba Mada 41.1710 43.7730 - 12.07.2019

Tba Paravani 41.3980 43.7870 13.07.2019

Tba Saghamo 41.2970 43.7540 — 12.07.2019

Tbilisi National Park 41.8808 45.0204 1270 m 22.06.2019

Tbilisi, Botanical Garden 41.6872 44 8056 470 m 07.07.2019

Tbilisi, Chilitba Lake 41.8150 44.6800 — 25.05.2019

Tbilisi, Dighomi 41.7780 44.7010 — 17.07.2019

Tbilisi, Dighomi, next to cemetery 41.7710 44.7670 —- 17.07.2019

Tbilisi, Kakheti Highway/Aleksandre Tvalchrelidze I Turn, hotel garden 41.6892 44.9078 480m 07.07.2019

Tedzami River 1.4 km NE Zemo Khandaki 41.9130 44.3160 - 09.07.2019

Tedzami River, tributary to Kura river, near Ertatsminda Village 41.8760 44.3144 - 30.06.2019

Telawi, Guest House LeNi 41.9152 45.4719 760 m 08.07.2019

Telawi, surroundings of hostel 41.9241 45.4701 685 m 08.07.2019

296 Jana Thormann et al.

Table 2. Continued.

Locality description Latitude Longitude Altitude Collection date

Teliankhevi River W Paldo 41.6110 45.4940 - 01.07.2019

ne drainage: Narvani River at Kobi Village close to confluence with Terek 42.5612 44 5093 = 04.07.2019

Terek drainage: Snostskali River near Akhaltsikhe Village 42.5940 44.6600 - 06.07.2019

nobis a ae River SE of Sno Village, small side channel of main 42.6003 44 6469 - 02.07.2019

Terek River 0.4 km W Pansheti 42.6350 44.6240 — 03.07.2019

Terek River 0.5 km S Kanobi 42.5850 44.5240 ~ 04.07.2019

Terek River 1.0 km WSW Stepanzminda 42.6530 44.6350 - 03.07.2019

Terek River 1.3 km SW Stepanzminda 42.6490 44.6340 - 03.07.2019

Terek River 1.6 km W Ukhati 42.5580 44.5020 - 04.07.2019

Terek Valley near confluence with Khde River, north of Dariali Monastery 42.7369 44.6314 1300 m 06.07.2019

Terek Valley, Darial Gorge, east of Tsdo 42.6936 44.6414 1560 m 05.07.2019

Terek Valley, Darial Gorge, near Gveleti 42.7036 44.6269 1480 m 05.07.2019

Terek Valley, Darial Gorge, south of Gveleti 42.7014 44 6283 1480 m 05.07.2019

Terek Valley, Gveleti Small Waterfall 42.7050 44.6147 1720 m 06.07.2019

Tkhilistskaro 41.8727 46.0050 337m 09.07.2019

Tkibuli towards Ambrolauri, below Nakerala Pass 42.3869 43.0119 1110m 02.07.2019

Truso Gorge at Terek 42.5797 44.4699 2023 m 07.07.2019

Truso Gorge, slope 42.5777 44.4647 2140-2212 m 07.07.2019

Nine. flowery meadows and scree fields, 400 m E of Abano; Kazbegi 42.6039 44.3034 2180-2300 m 3.072019.

Truso Valley, steep mountain side with flowers and scree; Kazbegi Municipality 42.5869 44.4305 2130-2200 m 3.07.2019.

Tsdo Village 42.6928 44.6353 1750m 17.07.2019

Tsdo, old road around the road tunnel 42.6900 44 6361 1620-1630 m 09.07.2019

Tsdo, small river valley close to the road tunnel 42.6842 44.6347 ees 09.07.2019

Tsiprani, Pschawi-Aragwi 42.2345 44.8450 852 m 05.07.2019

Ughviri Lake 43.0319 42.8273 1905 m 13.06.2019

Ureki, Shekvetili 41.9410 41.7660 ~ 02.08.2019

Ushguli 42.9143 43.0077 2082 m 15.06.2019

Vachiani Lake 41.3600 43.4390 — 11.07.2019

Vashlovani NP, bungalows near border to Azerbaijan 41.1110 46.6470 - 26.—27.07.2019

Walk to Gveleti waterfall 42.7047 44.6204 1555m —

Way from Bakuriani to Borjomi 41.7940 43.4690 - 23.07.2019

Wetland N Kvakhvreli 41.9640 44.1700 _ 09.07.2019