Dtsch. Entomol. Z. 70 (2) 2023, 311-335 | DOI 10.3897/dez.70.107425 Ate BERLIN Explosive radiation versus old relicts: The complex history of Ethiopian Trechina, with description of a new genus and a new subgenus (Coleoptera, Carabidae, ‘Trechin1) Arnaud Faille!, Sylvia Hofmann?, Yeshitla Merene**°, David Hauth?, Lars Opgenoorth®, Yitbarek Woldehawariat*, Joachim Schmidt® Department of Entomology, Stuttgart State Museum of Natural History, Stuttgart, Germany Leibniz Institute for the Analysis of Biodiversity Change, Museum Koenig, 53113 Bonn, Germany Faculty of Biology, Philipps University Marburg, Marburg, Germany Department of Zoological Sciences, Addis Ababa University, Addis Ababa, Ethiopia Amhara Agricultural Research Institute, Bahir Dar, Ethiopia Dn OF W DY HF General and Systematic Zoology, University of Rostock, Rostock, Germany https://zoobank. org/8D3E277C-424C-440B-8FES8-78085239C2A2 Corresponding authors: Amaud Faille (arnaud.faille@smns-bw.de); Joachim Schmidt (schmidt@agonum.de) Academic editor: Harald Letsch @ Received 3 June 2023 # Accepted 10 August 2023 @ Published 27 September 2023 Abstract The trechine beetle fauna (Coleoptera, Carabidae) of the Ethiopian Highlands is known to be highly diverse in species, and many species groups were recognized to be characterized by unusual character states of external and genital morphology. Earlier authors described several genera and subgenera of Ethiopian Trechina endemic to certain high mountains of the country. However, the relationships of these species groups and their evolutionary history are unknown so far. Here, we present the first molecular phylo- genetic analysis of Ethiopian Trechina, detect several synonymic names under 7rechus sensu lato, and introduce two new species groups to the country’s fauna: the monotypic genus Baehria Schmidt & Faille, gen. nov., with the type species B. separata sp. nov. from Mt. Choke in northern Ethiopia, and the 7rechus subgenus Abunetrechus Schmidt & Faille, subgen. nov., with the type species T. bipartitus Raffray, 1885; this subgenus includes three species of northern Ethiopia. We show that the composition of the Ethiopian fauna is based on multiple events of immigration, which started simultaneously with or some million years after the Oligocene-Early Miocene orogenic events north and south of the Rift Valley. Our results support the habitat island hypothesis for the evolution of the Ethiopian highland fauna. We found no evidence for an alternative hypothesis assuming a close connection of the Trechina im- migration to Ethiopia and Pleistocene cooling. We, thus, conclude that the geomorphological development rather than the climatic changes are the main drivers of the diversification of the high-altitude Trechina fauna in Ethiopia. Key Words Abunetrechus, Baehria, biogeography, checklist, mountains of East Africa, new species, new synonymy, phylogeny, 7rechus Introduction the southern African fauna, and two species were found in Ethiopia so far (Basilewsky 1974; Merene et al. 2023). The biogeographic and evolutionary history of the Tre- chini fauna of Ethiopia is complex and poorly under- stood. Members of two trechine subtribes are known to occur in the country. Within the subtribe Trechodina, the genus Pachydesus Motschulsky is a typical element of Similarly, only two species of Trechodes Blackburn oc- cur in Ethiopia; the genus belongs to Trechodina and is widely distributed in tropical-subtropical regions of Afri- ca, Madagascar, Indochina, the Philippines and Australia (Casale and Laneyrie 1982; Magrini et al. 2005; Faille et Copyright Arnaud Faille et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. si2 al. 2021). Another Trechodina genus, Perileptus Schaum, has a distribution similar to Trechodes but also occurs in the warm temperate zone of the Palearctic region. Four species of Perileptus are noted for the Ethiopian fauna (Casale and Laneyrie 1982; Deuve 2004; Merene et al. 2023). Based on our field investigations in Ethiopia, spe- cies of Pachydesus, Perileptus, and Trechodes have their habitats along brooks and rivers at lower and medium el- evations, with the highest occurrences of Pachydesus at altitudes up to 3200 m. Most of the Trechini species of the Ethiopian fauna belong to the megadiverse Holarctic genus Trechus Clair- ville (in the widest sense) of the subtribe Trechina and have their occurrences restricted to afromontane and afroalpine environments (Fig. 1). Molecular phylogenetic analyses have shown that the taxon Trechus (sensu lato) is polyphyletic (Faille et al. 2010, 2013). However, so far, the systematic position of the Ethiopian and East African representatives has not been addressed. In the mountains of East Africa, several Trechus (sen- su lato) lineages occur in widely separated high-altitude habitats (“mountains islands”): 21 species are placed in the subgenera E/gonophyes Jeannel, Elgonotrechus Jean- nel, and Trechus s. str. on Mt. Elgon, two species of the subgenus Meruitrechus Jeannel on Mt. Meru, and 64 7 aR. “ale al ‘g4/ Z “ if ¥ z aH Ze ‘ Gughe Highlands | L, MARS» KENYA NERV OK Guassa Plate \ A a | Aire 7 Guassa Plateau | Sh 4 rea. . Arnaud Faille et al.: The complex history of Ethiopian Trechina beetles species are placed in the subgenera Abyssinotus Quéin- nec & Ollivier, Archeotrechus Magrini, Quéinnec & Vi- gna Taglianti, Minitrechus Vigna Taglianti & Magrini, and Trechus s. str.) in the Ethiopian Highlands (Jeannel 1954a; Casale and Laneyrie 1982; Geginat 1995, 2008; Pawlowski 2003; Vigna Taglianti and Magrini 2010; Ortufio and Novoa 2011; Magrini et al. 2012; Schmidt and Faille 2018; Quéinnec et al. 2021). The eyeless Trechini species Nunbergites aethiopicus Pawlowski & Stachowiak, 1991, is special to the coun- try’s fauna. This species was found in the Simien Moun- tains (Fig. 1) and is considered closely related to Neotre- chus J. Muller of the subtribe Trechina from the eastern Mediterranean region (Pawlowski and Stachowiak 1991). Four additional Trechini genera and a total of 13 species were described from the mountains of northern Ethiopia just recently (Quéinnec et al. 2021): Aethiopsis Quéinnec & Ollivier, from Mt. Abune Yosef, and Afrotrechus Quéin- nec & Ollivier, Deuveopsis Quéinnec & Ollivier, and Nilotrechus Quéinnec & Ollivier from Mt. Choke (Fig. 1). Based on morphological character analyses, the authors conclude that the hypothesis of phylogenetic relationships between these genera and Trechodina is debatable. Overall, these data indicate that the Ethiopian High- lands might be considered a hotspot of Trechini diversity, “Ss, ny, * SOMALILAND _ | Arsi Volcanoes Bale Mountains ~~_+%. SOMALIA Figure 1. Map of Ethiopia showing the main topographic features of the country. Seven prominent volcanic massifs which are dis- cussed in the text are highlighted. The base map was downloaded from www.freeworldmaps.net (01-07-2023). dez.pensoft.net Dtsch. Entomol. Z. 70 (2) 2023, 311-335 not only in terms of species numbers but also lineage di- versity. Remarkably, in some Trechini genera, several species seem to be highly endemic to single Ethiopian volcanos (Quéinnec et al. 2021). However, phylogenetic relationships among Ethiopian Trechini species and spe- cies groups are largely unknown, and taxonomic concepts are in flux (Ortufio and Novoa 2011; Schmidt and Faille 2018). Also, the biogeographical history and evolution of the East African high-altitude Trechini fauna are rarely known. Related questions are part of a persisting contro- versial debate (Jeannel 1954b; Mani 1968; Ortufio and No- voa 2011; Schmidt and Faille 2018; Quéinnec et al. 2021). In recent years, a more intensive field investigation of the diversity, ecology and distribution of the high-altitude ground beetle fauna of Ethiopia was performed by authors of the present study. As a result, several new synonymies could be detected, and two additional, previously unrec- ognized trechine species groups were identified and are described in this paper. Most importantly, representatives of all Ethiopian 7rechus (s. |.) species groups and the gen- era recently described by Quéinnec et al. (2021) became available for molecular analysis. Based on this materi- al, we present the first dated phylogeny of the Ethiopian species. Using sequence fragments of two mitochondrial and two nuclear ribosomal genes, we aimed at answering the following questions: Are species groups recently de- scribed by Quéinnec et al. (2021) representatives of Tre- china or Trechini? How many independent immigration events caused the Ethiopian high-altitude trechine fauna to emerge? When did immigration of Palearctic trechine taxa occur — in the course of the Late Cenozoic cooling or, alternatively, in response to the major orogenic events in East Africa, starting in the Oligocene? Materials and methods Materials Specimens used for the morphological studies are listed in the respective Material sections of the relevant taxa, below. Specimens included in the molecular phylogenetic analyses are listed in Suppl. material 1. Taxonomy and identification of Ethiopian 7rechus species and species groups follow Jeannel (1927), Pawlowski (2001, 2003), Ortufio and Novoa (2011), Schmidt and Faille (2018) and Quéinnec et al. (2021). Institutional codes used in the tax- onomic treatment are as follows: CAF Arnaud Faille working collection, Stuttgart, Germany; CSCHM_ Joachim Schmidt working collection, later to be deposited in the Zoologische Staatssam- mlung, Munich; MNHN - Muséum National d’ Histoire Naturelle, Paris; UARK — University of Arkansas Arthropod Collection; NHMAA Natural History Museum, Addis Ababa University. 313 Morphological studies Specimens were examined by stereomicroscope Lei- ca M205-C. The photographs were taken with a Lel- ca DFC450 digital camera using a motorised focussing drive, light base Leica TL5000 Ergo, diffused light with Leica hood LED5000 HDI, subsequently processed with Leica LAS application software, and enhanced with CorelDRAW Graphics Suite X5. Body size was measured from the tip of mandibles in opened position to the apex of the longer elytron. The width of the head was measured across the widest portion including compound eyes. The width of pronotum and the width of elytra were measured at their widest points. The length of pronotum was measured along the median line. The widths of pronotal, apical and basal margins were measured between the tips of the apical and basal angles, respectively. The length of elytra was measured from the tip of the scutellum to the apex of the longer elytron. The length of the hind tibia was measured along its maximum length including its basal joint. The length of aedeagal median lobe was measured across the longest distance without consideration of the sagittal aileron. The follow- ing abbreviations were used in the species descriptions: AL Length of aedeagal median lobe; EL Length of elytra; EW Width of elytra; HW Width of head; PL Length of pronotum; PAW Width of pronotal apical margin; PBW Width of pronotal basal margin; PW Width of pronotum; TL Length of hind tibia. Molecular data acquisition Specimens used for the molecular study were collected alive by hand in the field and preserved in absolute etha- nol. Genomic DNA was isolated from whole specimens using a non-destructive extraction protocol (Rowley et al. 2007) and the DNeasy Tissue Kit (Qiagen GmbH, Hilden, Germany). The specimens were then dry mounted on a card, male genitalia extracted and included in a drop of water-soluble dimethyl hydantoin formaldehyde res- in (DMHF) on a transparent card beneath the specimen. Vouchers and DNA samples are deposited in the collec- tions of the State Museum of Natural History, Stuttgart (CAF, SMNS). In the phylogenetic analyses, we included 55 specimens belonging to 49 species of Ethiopian Tre- chini, including representatives of all genera and subgen- era described so far (Suppl. material 1). We complement- ed this data set with representatives of Trechodina and Trechus subgenera from Tanzania (Meruitrechus Jeannel) and Yemen (Arabotrechus Mateu), as well as representa- tives of all the lineages and clades of Jrechus Clairville and allied genera identified in Faille et al. (2013). dez.pensoft.net 314 Arnaud Faille et al.: The complex history of Ethiopian Trechina beetles Table 1. Node ages (My) and 95% HPD (height posterior density) of high-altitude trechine groups endemic to Ethiopia obtained from BEAST2 based on the concatenated data set. For calibration, two age constraints (ac) were implemented or combined with available substitution rates (sr) (see text, for details). Baehria gen. nov. Trechus (s. 1.) Abunetrechus subg. nov. Trechus (s. 1.) subgen. Abyssinotus Trechus (s. 1.) subgen. Minitrechus Stem Crown The sampling includes most of the type species of the genera treated in this paper, including 7rechus. Oth- er genera of uncertain affinities were also included: Anchotrechus Jeannel from the Canary Islands, Paratre- chus Jeannel and Oxytrechus Jeannel from Ecuador, Du- valiomimus Jeannel from New Zealand, Tasmanorites Jeannel from Tasmania, Bhutanotrechus Uéno from Bhu- tan, Agonotrechus Jeannel from Nepal, Trechisibus Mot- schulsky from Chile. We used Anillini and Bembidiini as outgroups. Two species of Patrobini were selected to root the tree since they are known to belong to the subfamily Trechinae but are outside the group revised in the present study (Maddison et al. 2019). We amplified three DNA fragments, two mitochondrial (the 5’ end of cytochrome c oxidase subunit 1, cox] and a fragment containing the 5’ end of large ribosomal unit plus the Leucine transfer plus the 3’ end of NADH dehydroge- nase subunit 1, rrnL+trnL+nad1; ca 740 bp) and two nucle- ar (large and small ribosomal unit, LSU and SSU rRNA). For the primers used, see Suppl. material 2, and for the general PCR conditions, see Faille et al. (2010). Sequences were assembled and edited with Bioedit v. 7.0 (Hall 1999) and Geneious Prime 2019.2.3 (Kearse et al. 2012). New sequences have been deposited in GenBank database, with accession numbers indicated in Suppl. material 1. A few sequences were taken from Faille et al. (2010, 2011, 2013, 2014) and Fresneda et al. (2019) (Suppl. material 1). We aligned the sequences using the online version of MAFFT v.7 (Katoh et al, 2019) using the L-INS-i algo- rithm and default parameters. Maximum likelihood anal- yses were conducted on the concatenated alignment using RAXxML v.7.2 (Stamatakis 2006), with thorough boot- straps, 20 runs, 500 reps, and a GTR+I+G evolutionary model (Stamatakis 2006, 2014). We used the default val- ues for other parameters of the search (Stamatakis 2014). Divergence time estimations were performed based on the concatenated data set, partitioned by genes and co- dons using BEAST2 v.2.6.7 (Bouckaert et al. 2019). We specified HK Y substitution models, a birth-death prior, and a relaxed log-normal clock. To calibrate the tree, we implemented two age constraints: cl, upper age range of 24 Mya for the 7rechus clade of southern Ethiopia based on the maximum age of the Bale Mountains (Abbate and Bruni 2015) (uniform distribution); c2, a minimum age of 98 Mya for the Trechini stem group based on the ear- liest fossil of that group (log-normal distribution M=2, S=1.105, offset=99). The oldest trechini fossils are known from Burmese amber (coll. D.R. Maddison and coll. A. Faille; data will be published by A. Faille and co-workers dez.pensoft.net 15.19 (9.86-21.13)| 2.13 (1.03-3.27) | 10.67 (6.83-14.62) | 3.62 (1.52-5.96) | na. | 17.5 (13.01-22.23) | na. actsr [22.87 (15.9-30.08)| 4.19 (2.12-6.85) | 17.74 (12.93-22.96)| 6.01 (3.02-9.36) | na. | 26.22 (21.51-30.96) | na. Crown 16.57 (12.44-21.04) 22.87 (20.89-24.00) Crown Stem elsewhere). The amber is dated as about 98.79 + 0.62 Ma (earliest Cenomanian; Shi et al. 2012). We also used the rates estimated for the same gene fragments in a previ- ous work on Carabidae, all with a normal distribution and a standard deviation of 0.3 (i.e. 0.0016 changes/branch/ Ma for 16S; 0.0013 for 28S; and 0.0145 for COI; Andt- jar et al. 2012). Two dating approaches were performed: i) using the age constraints only, and 11) including age constraints and available rates. Six (1) and three (11) runs were performed, each with 100 million generations, and a thinning range of 10,000. Replicate runs were then com- bined with BEAST2 LogCombiner v.2.6.7 by re-sam- pling logs and trees from the posterior distributions at a lower frequency (11) and using a burn-in of 10% for each data-set, resulting in a final set of ~18,000 (1) and ~27,000 trees (11). Convergence and stationary levels were verified with Tracer v1.7.1 (Rambaut et al. 2018). We annotated the tree information with TreeAnnotator v.2.6.7 and visu- alized it with FigTree v.1.4.2. Results In accordance with previous molecular phylogenetic stud- ies on Trechinae (Faille et al. 2013, 2021; Maddison et al. 2019), we recovered a monophyletic Trechini clade with Trechodina forming the sister clade of Trechina. The genus Oxytrechus is sister to the remaining Trechina, and the Tas- manian 7asmanorites together with the Chilean representa- tive of Trechisibus form a well-supported clade. These two taxa are sister clade to the remaining Trechina. Inside this large clade of Trechina, the main clades found in Faille et al. (2013) are largely recovered: the Isotopic clade (Clade 2 in Faille et al. 2013), the Pyrenean hypogean clade (Clade 1.1 in Faille et al. 2013), the Epaphius clade, the Dina- ro-Alpine clade (Clade 1.3.2.1 in Faille et al. 2013) and the Trechus clade (Clade 1.3.2.2 in Faille et al. 2013). Phylogeny of Ethiopian Trechina The Ethiopian high-altitude trechine fauna consists of at least three isolated clades, all of which cluster within Tre- china (Figs 2, 3). Because Nungbergites was not available to our study, the systematic position of this taxon remains to be tested. The monotypic genus Baehria gen. nov. from Mt. Choke in northern Ethiopia (see below, for description) forms a well-supported clade together with the Trechina SH lis} Dtsch. Entomol. Z. 70 (2) 2023, 311-335 ‘aIYDO poYSITYSTy st ‘Aou Uds vi/yapg otwMOpus ueIdoIy)Y oy} WIM OPR]O SNYIadjDAD J-SNULWUOL]OANG] S1OYdstuUsY UIDYINOS SY], ‘(CT OZ) ‘Te 39 [wy JO S}Nsor snorasid 0} JZOI Spe] SUTYDON PosOfOd ‘posdeyjoD ‘ase spou sANsadser 9y} JO GqH %S6 94} Apioods sieq Adin ‘AjoATOOdsol ‘¢/< sanjea de.ns}oog pue g6'Q< SOTIGeqoid 10119}SOd 0} JOJOI SOpOU YURI JB SIVJS PUL SOTDIID YORT_ ‘(1XO} 99S STIRJOP OF) SJUTRIISUOS O8ev OM} SUISN pue eyep sdUONbos poyeusyeou0s oy) UO poseg ZTLSVA_ Woy pourejqgo sem 904 oY], “(¢ ‘SITY Ul sonuTUOD ‘9027 9Y} Jo Wed [eseq) sdno1sjno pue sopeoq tuTydory, Jo AuosoyAyd pojesqiyes-owiy SLOWeI °Z WANS eAWO Sz os- Sl- ool SZL- ZOL1 siyesodwa} snjajjauag Z9ZT Sryuane snqojed — ti. SeuIgoljed Lp winjejnuny uoipiquieg ZO “ds sueyoojydry LPZIW Snoeeo snyjiuy ry TUN y = GOLLT weueon Ju “oysary 1ui|! O9PLT Jpnenye sapoysa, ay * L4 snjejoge snjdajiad 1LZ7 WAewyes snsapAysed * eu | poyosl | 6EZLL7 snuevojenba snysaiAxQ Pez] sueinbue ‘su snqisiyoass 8P9L7 ‘ds sawouewse, jUIYOOd | * euUIudeL | Z90/-7 Is@maipue snysejqouoby UeAYOg snydajo1aq] ZS7 sisugjossua snyoaujosapoq O0Z7 Snuvew snday tv luiqos sisdoday 6ZS1L7 sisueenquuis sisdoiydedz 6SZ1 SousIWW snwajqoyaeL, LLOT Snosip snwelg (eL0z se 38 ales WZ ape|y) apels s1doj03) % 99¢%7 sdeaiqo/6 sajasimobbayg fe | P£9L-7 sioojxeyas snyseqoueNYg —S—— (€Loz 2 79 ayes ul |", apelD) apeya ueaBodAy ueauashg ZBL Suaqns "Fy PezT sneegheys J OE9T Ssmero "FZ PSZ7 syeoes sniydedz SLET Haug snyoajojydA Ore7 99810 W La yssnyneyos SNUWS/GOUBLSpPapy zr snuejuowesdns sdoueeydepies pj snueyinaig sdouseydeseg fl * ¢ Big ul panuruos dez.pensoft.net 316 Arnaud Faille et al.: The complex history of Ethiopian Trechina beetles Minitrechus Aethiopsis guassaensis_L2238 Aethiopsis nr. guassaensis_L2247 . Nilotrechus sp. nr. reebae_L1821 Abyssinotus ) 4 3 Deuveopsis (Abayopsis) sp. nr. basilewskianus_L1810 Deuveopsis (Abayopsis) sp. nr. basitewskianus_L1811 * Kl Trechus sp. Mt. Choke_L1806 ij | Afrotrechus abyssinicus_L1817 Trechus (s. I.) = Deuveopsis lobeliae_L1815 Afrotrechus (Abyssiniopsis) amharicus_L1820 Ny * Abyssinotus sp. nr. salomon_L1814 Abyssinotus sp. nr. gigas_L1823 fej 7 a * & Abyssinotus sp. nr. gigas_L1819 Mountains of EB Trechus sp. Guassa_L2239 N-Ethiopia Trechus sp. Abune Yosef_L1838 Tr. sublaevis_L1839 Speotrechus mayeti_L900_A4 i Pc | / Trechus clade (Clade 1.3.2.2 in Faille et a/ 2013) -30 -25 -20 -15 -10 -5 0 Mya Figure 3. Subtree of the ultrametric time-calibrated phylogeny of Trechini beetles as shown in Fig. 2 (see there for details). Collapsed, colored clades refer to previous results of Faille et al. (2013). Clades which include Ethiopian 7rechus (sensu lato) spe- cies are highlighted by different colors (see text for details). continued in Fig. 2 Dinaro-Alpine hypogean clade (Clade 1.3.2.1 in Faille ef a/ 2013) Jeannelius birsteini L515 dez.pensoft.net Dtsch. Entomol. Z. 70 (2) 2023, 311-335 genera Duvaliomimus from New Zealand and Paratrechus from South and Central America (in the following the BDP clade). Based on our dataset, the sister group of this clade is formed by the Palearctic “isotopic clade” of Trechina (Faille et al. 2013), although with low support (Fig. 2). All other Ethiopian high-altitude trechine species clus- ter within the megadiverse genus 7rechus (sensu lato). Within the genus 7rechus, Abunetrechus subgen. nov. (see below, for description) is a member of a well-sup- ported clade which includes the monotypic Anchotrechus Jeannel from Tenerife, the monotypic Arabotrechus Ma- teu from Yemen, as well as the subgenus Meruitrechus Jeannel with two species from Mt. Meru, Tanzania (in the following the “AAMA clade’). The tree shows low supported basal branching of Abunetrechus subgen. nov. within the AAMA clade (Fig. 3). The remaining Ethiopian T7rechus (s. |.) form two clades that strictly separate the northern and southern Ethiopian fau- nas (Fig. 3). A sister relationship of the northern and south- ern Ethiopian clades is not supported by our tree. Moreover, the results do not support a close relationship of these two clades with any of the Holarctic Trechus (s. |.) lineages. The southern Ethiopian Jrechus clade includes all species known to occur in the Bale and Arsi Mountains and the Gughe Highlands, south and west of the Rift Valley. The two species from the Gughe Highlands form a separate lineage within one of the two main clades of South Ethiopian 7rechus (Fig. 3). This clade comprises only species with markedly small body size (Schmidt and Faille 2018). The taxa Archeotrechus and Minitrechus are closely related and representatives of a terminal clade within southern Ethiopian 7rechus (s. |.) (Fig. 3). Our phylogenetic analyses show a similar picture for the trechine fauna of the highlands in northern Ethiopia. Samples of this clade originated from the Abune Yosef Massif, Guassa Plateau, and Mt. Choke. Based on the dat- ed tree, all of the Trechini genera and subgenera recent- ly described by Quéinnec et al. (2021) cluster within the northern Ethiopian 7rechus (s. |.) clade (Fig. 3); these are Abyssinotus, Abayopsis, Abyssiniopsis, Aethiopsis, Afro- trechus, Deuveopsis, and Nilotrechus. The systematic positions of the 7rechus (s. |.) species from the Simien Mountains in northern Ethiopia and of T. aethiopicus Alluaud, 1918 from the mountains near Addis Abeba remain unknown due to the lack of molec- ular material. Noteworthy, among the Palearctic species groups, both the monotypic genera Anchotrechus Jeannel from Tener- ife, and Speotrechus Jeannel from France, are nested within 7rechus (s. 1.) in our phylogeny. Molecular dating of the Trechina species groups endemic to Ethiopia Stem and crown group ages of the trechines endemic to Ethiopia, as calculated with BEAST2 using two different dating approaches, are summarized in Table 1. 317 Taxonomic results Baehria Schmidt & Faille, gen. nov. https://zoobank.org/7 1FD62CE-3922-4927-9A70-E94D7DD7A7A0 Figs 4-15 Type species. Baehria separata sp. nov., herewith designated. Diagnosis. Representative of subtribe Trechina due to presence of bidentate mandibles (absence of retina- cle) and dorsally closed aedeagal median lobe (Jeannel 1926). Comparatively large trechines characterized by ro- bust head, large mandibles, small but markedly protrud- ed eyes, smooth, markedly convex tempora, cordiform pronotum, straight pronotal basal margin with large, rect- angular to acute laterobasal angles, slender elytra, short metepisternum, reduced hindwings, moderately slender antenna and legs, protibia with a complete longitudinal groove on external surface, presence of a row of long ad- hesive hairs on apical margins of 4" tarsomeres which are as long as the 5" tarsomeres, and by a markedly slender median lobe of aedeagus, which possesses a small, slight- ly sclerotized endophallic copulatory piece. The new ge- nus is particularly distinguished from other representa- tives of Trechina by unusual chaetotaxy of head capsule and elytra as follows: clypeus plurisetose, each side with 3—5 setae; submentum with three setae each side; anterior elytral discal seta located in the 4" interval, adjoined to the 4" stria. Etymology. The new genus name is given in memo- riam of our dear friend and colleague, the distinguished entomologist Martin Baehr, Munich (10.03.1943- 17.04.2019). Description. Head: Large and robust, without pilos- ity. Mandibles large, moderately slender, with bidentate dentition pattern as shown in Fig. 8. Labrum with apical margin moderately emarginated, with six setae near api- cal margin. Clypeus each side with three or four setae (Figs 5, 7; seldom only two setae at one of the sides). Eyes moderately small, as long as or slightly shorter than tempora, markedly convexly protruded (Figs 5, 7). Two supraorbital setae each side in normal position for Tre- china. Supraorbital furrows unevenly bent in posterior half, markedly deep in front and middle portions, slightly flatter near insertion of posterior supraorbital seta. Tem- pora markedly convex, markedly wrinkled to the neck, smooth. Mid of head convexly elevated, with a distinct transverse depression between supraorbital area and neck (Figs 5, 7). Antennae slender, with third antennomere longest, 1/9-1/10 longer than first respectively fourth antennomere, and with second antennomere about 2/3 of length of third. Suborbital seta present. Apical tooth of mentum bifid, sensory pits of mentum present; submen- tum with three setae each side (Fig. 9). Prothorax: Pronotum rather small, without pilosity, moderately transverse, cordate, broadest distinctly before middle, with lateral margin markedly concave before base, and with basal margin slightly smaller than apical dez.pensoft.net 318 margin (Figs 4, 6). Disc moderately convex. Anterior mar- gin straight or slightly concave in middle with anterior angles small but distinct, rounded, moderately protruded. Basal margin straight along internal 3/4, with laterobasal angles slightly shifted posteriad. Lateral margin convexly rounded in anterior 2/3 and concave towards laterobasal angles, latter large, rectangular or sharp at tip, sometimes slightly protruded laterally. Marginal gutter moderately broad throughout. Median longitudinal impression sharp- ly incised, disappearing near apex, somewhat deepened before base. Anterior and posterior transverse impres- sions shallow and smooth. Laterobasal foveae large, in- ternally and externally (towards lateral gutter) distinctly sloped, without punctures but with fine transversal wrin- kles. Lateral and laterobasal setae present, with the for- mer situated at or slightly anterad of maximum width of pronotum. Proepisternum glabrous and smooth. Pterothorax: Elytra without pilosity, long and very slender ovate, very slightly convex or flattened in middle of disc, in dorsal view broadest distinctly posterad mid- dle, shoulders flatly rounded (Fig. 13), apical sinuation distinct, apex rounded with the indication of a very ob- tuse apical angle. Striae 1-8 complete, moderately deep impressed, impunctate, intervals moderately convex, par- ascutellar stria free, 1/6—1/9 of length of elytra. Recurrent preapical stria deep, long, in most specimens reaching the apex of the fifth stria. Parascutellar seta present. Anteri- or discal seta located in the 4" interval, adjoined to the 4" stria, located near the end of the anterior elytral 5" (Fig. 13); second discal seta located at the 3™ stria about at elytral middle (in most specimens, the posterior setif- erous pore together with the 3" stria is switched into the 4" interval); posterior discal seta (= subapical seta near the end of 3" stria) present, located about 1/9 of elytral length from elytral apex; subapical seta of the recurrent stria isolated, distinctly removed from this stria by dis- tance of 2—3 diameters of the setiferous pore. Number and positions of the setae of the marginal umbilicate se- ries as in Trechus s. str. Metepisternum very short, gla- brous and smooth, with outer margin about as long as anterior margin. Legs: Moderately long and robust. Protibia distinctly dilated towards apex, straight, with longitudinal groove on dorsal surface complete, and with several fine setae on anterior surface near apex (Fig. 12). Two basal protar- someres of males dilated and dentoid at the inner apical border (Fig. 10). Fourth pro-, meso- and metatarsomeres each with a row of long adhesive hairs on apical margins which are as long as the 5" tarsomeres (Figs 10, 11). Male genitalia (Figs 14, 15): Aedeagal median lobe markedly elongated, slender tube-like, in lateral view slightly sinusoidal, with apex distinctly bent upwardly, simple, with apical lamella insignificant; basal bulb rath- er small with large sagittal aileron. Endophallus with a small, very slightly sclerotized copulatory piece. Param- eres with 3-4 apical setae. Distribution. So far only known from Mt. Choke in northern Ethiopia (Fig. 1). dez.pensoft.net Arnaud Faille et al.: The complex history of Ethiopian Trechina beetles Relationships and identification. Based on the mo- lecular data, Baehria gen. nov. is representative of a well-supported clade comprising Duvaliomimus Jean- nel from New Zealand and Paratrechus Jeannel from South and Central America (Fig. 2). Apart from a gen- eral “Duvalius-like” appearance, the three genera share some common features like the large size, cordate pro- notum, rounded head with salient temples, and pubes- cent protibiae. Baehria gen. nov. differs from both these genera by the presence of three or four setae instead of two on each side of the clypeus, and by the position of the anterior discal seta on elytra: in Baehria gen. nov., this seta inserts in the 4" interval instead on the 3” stria in Duvaliomimus and on the 5" stria in Paratrechus. Baehria gen. nov. differs additionally from Paratrechus by the simple apex of the aedeagal median lobe, which is button-like shaped in Paratrechus (Jeannel 1928; Barr 1982: Townsend 2010). Baehria separata Schmidt & Faille, sp. nov. https://zoobank.org/430BB485-1056-462B-8088-FF992DE5F947 Figs 4-15 Type material. Holotype male, with label data: Ethio- pia, Amhara, Mt. Choke, crater valley, alt. 3780-3900 m, 10°42'12"N, 37°50'58"E, 27.11.2019, leg. D. Hauth, J. Schmidt, Yeshitla M., Yitbarek W. (CSCHM). Paratypes: 39 males, 54 females, same data as holo- type (CAF, CSCHM, NHMAA); 2 males, Mt. Choke, crater valley, alt. 3700-3800 m, 10°41'14"N, 37°50'07"E, 24.11.2019, leg. D. Hauth, J. Schmidt, Yeshitla M., Yitba- rek W. (CSCHM); 6 males, 7 females, Mt. Choke, western crater valley, alt. 3500-3600 m, 10°41'00"N, 37°50'35"E, 01.V.2022, leg. J. Schmidt, Yeshitla M., (CSCHM). Additional material. 3 males, 3 females, W-slope Mt. Choke, alt. 3370 m, 10°38'07"N, 37°45'51"E, 23.11.2019, leg. D. Hauth, J. Schmidt, Yeshitla M., Yitbarek W. (CAF, CSCHM); 6 males, 7 females, W-slope Mt. Choke, alt. 3700-3900 m, 10°42'17"N, 37°50'29"E, 25.11.2019, leg. D. Hauth, J. Schmidt, Yeshitla M., Yitbarek W. (CAF, CSCHM); 43 males, 31 females, W-slope Mt. Choke, “Shoa Kidaneberet” valley, alt. 3700-3800 m, 10°39'08"N, 37°49'45"E, 8.V.2022, leg. J. Schmidt, Ye- shitla M. (CSCHM); 10 males, 10 females, N-slope Mt. Choke, alt. 3800-3950 m, 10°43'16"N, 37°51'15"E, 26.11.2019, leg. D. Hauth, J. Schmidt, Yeshitla M., Yit- barek W. (CSCHM); 18 males, 20 females, N-slope Mt. Choke, alt. 3750-3850 m, 10°43'51"N, 37°52'15"E, 09.V.2022, leg. J. Schmidt, Yeshitla M. (CSCHM); 3 males, 2 females, N-slope Mt. Choke, above Gumadur, alt. 3750-3850 m, 10°44'10"N, 37°53'48"E, 05.V.2022, leg. J. Schmidt, Yeshitla M. (CSCHM); 1 male, N-slope Mt. Choke, N of Waber, alt. 3450-3600 m, 10°44'48"N, 37°46'22"E, 07.V.2022, leg. J. Schmidt, Yeshitla M. (CSCHM); 13 males, 5 females, Mt. Choke, eastern cra- ter valley, alt. 3700-3800 m, 10°42'59"N, 37°54'13"E, 06.V.2022, leg. J. Schmidt, Yeshitla M. (CSCHM). Dtsch. Entomol. Z. 70 (2) 2023, 311-335 319 3 7 Figures 4—7. Baehria separata Schmidt & Faille, gen. nov., sp. nov., dorsal aspect of body (4, 6) and head (5, 7) of paratypes; 4, 5. Male; 6, 7. Female. The small white circles in Figs 5 and 7 mark the insertion points of the clypeal setae. Etymology. The specific epithet refers to the markedly Description. See description of genus. separated distributional area of the taxon, which is, based Body length: 6.9-7.5 mm (@ = 7.19 mm, n= 20). on current knowledge, far away from its next relatives. It Proportions (n = 20): PW/HW = 1.18—1.25 (0 = 1.21); is built by the past participle of the Latin verb separare. PW/PL=1.30—-1.38 (@ = 1.34); PW/PBW = 1.48-1.54(0= dez.pensoft.net 320 Arnaud Faille et al.: The complex history of Ethiopian Trechina beetles Figures 8-15. Baehria separata Schmidt & Faille, gen. nov., sp. nov. 8. Left and nght mandible, dorsal aspect; 9. Ventral aspect of head; the small white circles mark the insertion points of the setae on submentum; 10. Right male protarsomeres, left latero-ventral as- pect; 11. Left male metatarsomeres, right lateral aspect; 12. Right male protibia, dorsal aspect; 13. Anterior part of elytra and pronotal base; the arrows point to the insertions of the anterior elytral discal setae; 14. Aedeagus, left lateral aspect; 15. Aedeagus, dorsal aspect. 1.50); PBW/PAW = 0.92-1.00 (0 = 0.96); EW/PW =1.42— _ brown, basal 3/4 of femora light brown; antennal base in 1.48 (O = 1.45); EL/EW = 1.46-1.56 (O = 1.51). some specimens more widely brightened. Colour: Dark brown to blackish, moderately shiny in Microsculpture: Same in males and females. Head both sexes; palpi light brown, labrum and scapus reddish with deeply engraved, rather large, almost isodiametric dez.pensoft.net Dtsch. Entomol. Z. 70 (2) 2023, 311-335 sculpticells on disc and supraorbital area, slightly small- er sculpticells on clypeus. Pronotum with moderately deep engraved, slightly transverse sculpticells on disc and markedly deep engraved sculpticells near base; the sculpticells are somewhat smaller than on head disc. Elytral intervals with more finely engraved sculpticells which are more transverse than on pronotum. Aedeagus. Proportion EL/AL (n = 10): 2.40-2.64 (QO = 2.52). Median lobe in lateral view unevenly bent, dorsally with a distinct concavity before middle (Fig. 14), in dorsal view not or very slightly broadened before apex (Fig. 15). Differential diagnosis. See Diagnosis and Identifica- tion sections of the genus, above. Distribution and geographical variability. The type series was collected on the western side of the crater valley of Mt. Choke. Additional populations were collected on the north eastern side of the crater valley and on northern and western slopes of Mt. Choke. Specimens of these pop- ulations differ +/- distinctly from those of the type series and from each other by the curvature of the aedeagal medi- an lobe, the number of lightened basal antennomeres, and the depth of the engraving of the elytral microsculpture. Slight differences were also found in the DNA sequence segments of the three investigated specimens represent- ing three different populations (Fig. 2; Suppl. material 1). Further morphological and molecular genetic studies are needed to answer the question of whether certain popula- tions represent separate species or subspecies. Habitat. Specimens of Baehria separata gen. nov., sp. nov. have been found in stone packs traversed by run- ning water in small steep streams in the afroalpine zone, together with Dytiscidae beetles (Fig. 16). Based on this finding, Baehria separata gen. nov., sp. nov. seems to be adapted to a rheophilic way of life. Trechus Clairville, 1806 Abunetrechus Schmidt & Faille, subgen. nov. https://zoobank. org/608E 14D6-39EC-4CEC-97C9-68870B659966 Type species. 7rechus bipartitus Raffray, 1885, herewith designated. Diagnosis. Representative of Trechina and TJrechus s. |. sensu Jeannel (1926, 1927, 1928) due to presence of bidentate mandibles (absence of retinacle), dorsally closed aedeagal median lobe, well-developed compound eyes, protibia glabrous on anterior surface, presence of two elytral dorsal setae situated in third interval, elytral intervals glabrous, 4+2+2 pattern of umbilicate setae, and two basal tarsomeres of male dilated. Externally, Abunetrechus subgen. nov. reminders a non-specialised high-altitude Trechus of moderate body size, short mandibles, antenna and legs, moderately large eyes, rounded humeri, short metepisternae and hindwings reduced to short stubs. 7rechus sensu lato, Abunetrechus subgen. nov. is characterized by the combination of following character states: bisetose clypeus (Figs 18, fs all 20, 22), smooth tempora, pronotum with fully rounded laterobasal angles and with laterobasal setae markedly protruded anteriorly (Figs 18, 20, 21), elytral striae 3 and 4 merging at level of the anterior discal seta (Fig. 23), elytral preapical seta of the third interval present and situated about at level of the elytral apical tenth, protibia with longitudinal groove on external surface; aedeagus with two moderately sclerotized portions of the endophallus arranged one behind the other in apical half of the median lobe (Figs 24—29). Etymology. The subgenus name combines the name of the Abune Yosef Massif in northern Ethiopia, where the species of this subgenus occur, with the name of the genus Trechus. Description. Head: Size averaged for Trechus sensu lato, without pilosity. Mandibles short, with dentition pat- tern as in 7rechus sensu stricto. Labrum with apical margin moderately emarginated, with six setae near apical margin. Clypeus each side with one long seta (Figs 18, 20, 22; very seldom with an additional very fine seta situated interior of one of the primary setae). Eyes moderately large, convexly protruded, more than two times as long as tempora, latter moderately convex (Figs 18, 20, 22). Two supraorbital se- tae each side in normal position for 7rechus. Supraorbital furrows moderately deep and almost evenly bent through- out. Tempora moderately convex, markedly wrinkled to the neck, smooth. Mid of head convexly elevated. Anten- nae short, with third antennomere slightly longer than ped- icellus. Suborbital seta present. Apical tooth of mentum truncate or slightly bifid, sensory pits of mentum present but very small; submentum with 4—7 setae. Prothorax: Pronotum with size averaged for Trechus sensu lato, without pilosity, slightly transverse, broadest distinctly before middle, with lateral margin completely rounded towards base, and with laterobasal angles indis- tinct. Basal margin (between insertion points of laterobasal setae) distinctly broader than apical margin. Disc marked- ly convex. Anterior margin slightly or moderately concave with anterior angles shortly rounded, moderately protrud- ed. Basal margin straight or slightly convex in middle and with outer quarters markedly shifted anteriorly towards lateral margin (Figs 18, 20, 21). Lateral margin convex- ly rounded throughout; laterobasal angle fully rounded or marked as a very small blunt tooth. Marginal gutter very narrow throughout. Median longitudinal impression slightly incised, disappearing near apex and base, not deepened within area of posterior transverse impression. Anterior and posterior transverse impressions very shal- low, smooth or (posterior transverse impression) some- times finely wrinkled. Laterobasal foveae rather small, moderately impressed, smooth. Lateral and laterobasal se- tae present, with the former situated near maximum width of pronotum. Proepisternum glabrous and smooth. Pterothorax: Elytra without pilosity, slender ovate, markedly domed towards disc, not flattened in middle of disc, in dorsal view broadest slightly posterad middle, shoulders flatly rounded, apical sinuation very slightly developed or indistinct, apex rounded or marked as an ob- tuse apical angle. Parascutellary stria short to moderately dez.pensoft.net 322 Figure 16. North-exposed slope on Mt. Choke with Erica forest and a steep small brook at an altitude of 3600 m during dry sea- son (May, 2022). The stone pack in the brook 1s habitat of Baeh- ria separata Schmidt & Faille, gen. nov., sp. nov.: the beetles were collected between the stones along which the water flows (in order to find the beetles, the creek bed was partially dug up). long, free; striae 1-8 almost complete, moderately deep impressed in middle of disc, less deeply towards sides, disappearing near base, crenulated, striae 3 and 4 merg- ing at level of the anterior discal seta; intervals slightly convex. Recurrent preapical stria deep, long, connected with the apex of the fifth stria. Parascutellar seta present. Anterior discal seta situated at merging point of the 3 and 4" stria, near the end of the anterior elytral 5" (Fig. 23); second discal seta located at the 3 stria somewhat be- hind elytral middle; posterior discal seta (= subapical seta near end of 3™ stria) present, located about 1/10 of elytral length from elytral apex; subapical seta of the recurrent stria isolated, removed from this stria by distance of 1—2 diameters of the setiferous pore. Number and positions of the setae of the marginal umbilicate series as in Trechus s. str. Metepisternum very short, glabrous and smooth, with outer margin about as long as anterior margin. Legs: Short and moderately robust. Protibia distinctly dilated towards apex, straight, glabrous, with longitudi- nal groove on dorsal surface complete. Two basal protar- someres of males dilated and dentoid at the inner apical border. Chaetotaxy as in 7rechus sensu stricto. Male genitalia (Figs 24-29): Aedeagal median lobe moderately large, in lateral view markedly curved, with apical lamella short, latter with distinct terminal capitu- dez.pensoft.net Arnaud Faille et al.: The complex history of Ethiopian Trechina beetles lum; basal bulb and saggital aileron averaged. Endophal- lus with a moderately large, moderately sclerotized folding structure (copulatory piece) in the shape of a half- open cylinder or cone which is located in apical half of the median lobe and directed to its longitudinal axis, and with the open part of the copulatory piece facing ventrad. Apicad of this piece, an additional slightly more strongly sclerotized folding structure is developed which is shaped as a small plate (best visible in lateral view); the basal part of this piece overlaps with the apex of the more dor- sal copulatory piece. Parameres with 2-4 apical setae. Remarks. In his redescription of Trechus bipartitus, Jeannel (1927) noted the presence of a single copulatory piece which is characterized by a long sinusoidal appen- dix. However, in his figure of the left lateral view of the copulatory piece (Jeannel 1927: 195) he merged the more strongly sclerotized folding structure near median lobe apex with the more basad located larger copulatory piece which leads to the impression of a single, very long piece. Distribution. Northern Ethiopia Plateau (Fig. 1): Three species are known so far, two from Mt. Abune Yo- sef (1. bipartitus Raffray, T! lalibelae Quéinnec & Olli- vier) and one from the Guassa Plateau (7. habeshaicus Quéinnec & Ollivier). Relationships and identification. Based on the mo- lecular data, Abunetrechus subgen. nov. is representa- tive of a clade comprising Anchotrechus Jeannel from Tenerife, the Trechus subgenus Arabotrechus Mateu from Yemen, and the 7rechus subgenus Meruitrechus Jeannel from Mt. Meru, Tanzania (Fig. 3; in the follow- ing called the AAMA clade). Abunetrechus subgen. nov. differs from all species groups of the AAMA clade by bisetose clypeus. A quadrisetose clypeus was hypothe- sized plesiomorphic character state in Trechini (Schmidt et al. 2021). Within this tribe, a bisetose clypeus is also developed in the genus Omalodera Blanchard from Chil1, the Caucasian genus A/anorites Belousov of the Neotre- chus Phyletic Series, and two Epaphiopsis Uéno species occurring in the central Himalaya (Belousov 1998; Naito 2023). However, this character state has to be considered homoplasic because none of these taxa cluster within Trechus sensu lato (Faille et al. 2013, 2021; Maddison et al. 2019, see Fig. 2 in this paper). Abunetrechus sub- gen. nov. additionally differs from all other species of the AAMA clade by rounded pronotal laterobasal angles, from Meruitrechus by presence of the elytral preapical seta of the third interval, the isodiametric sculticells on elytra less deeply engraved, and eight striae well marked, from Anchotrechus by smaller and stouter body, glabrous elytra and much shorter aedeagal median lobe, and from Arabotrechus by the smaller body size, presence of a sec- ond discal setae (missing in Arabotrechus, as well as in T: aethiopicus and some species of E/gonotrechus Jean- nel. Abunetrechus subgen. nov. shares the elytral striae 3 and 4 merging at level of the anterior discal seta with Arabotrechus (based on a single investigated specimen; larger series would be necessary to confirm the stability of this character). Dtsch. Entomol. Z. 70 (2) 2023, 311-335 Trechus (Abunetrechus) bipartitus Raffray, 1885 Figs 17, 18, 23-25 Trechus bipartitus Raffray, 1885: 318; locus typicus: “col du mont Abouna- Yousef (4024 my)”. Trechus bipartitus: Jeannel 1927: 194. Trechus bipartitus: Pawtowski 2003: 157. Trechus bipartitus: Ortufiio and Novoa 2011: 137. Trechus (s. str.) bipartitus: Quéinnec et al. 2021: 20. Type material. Not studied. The lectotype was designated by Quéinnec et al. (2021). Identification 1s based on the redescriptions of the species, including habitus and male genital figures of the type specimens, presented by Jeannel (1927) and Quéinnec et al. (2021), as well as on compre- hensive material collected at the type locality (see below). Additional material. 6 males, 9 females, Ethio- pia, Amhara, Mt. Abuna Yosef, N-slope, 3800-3950 m, 12°07'52"N, 39°11'39"E, 4.111.2019, leg. D. Hauth, J. Schmidt, Yeshitla M., Yitbarek W. (CAF, CSCHM); 6 males, 12 females, ditto, S-slope, 3850-3900 m, 12°07'29"N, 39°11'21"E, 5.11.2019, leg. D. Hauth, J. Schmidt, Yeshitla M., Yitbarek W. (CSCHM, NHMAA); 5 males, 2 females, ditto, S-slope, 3700-3850 m, 12°09'10"N, 39°09'35"E, 5.1II.2019, leg. D. Hauth, J. Schmidt, Yeshitla M., Yitbarek W. (CSCHM). Additions to the species description. Mature species with elytra including suture blackish brown (Quéinnec et al. 2021: “suture distinctly brown shiny”; this informa- tion 1s probably based on confusion with 7? habeshani- cus). Body length 4.14.7 mm (Quéinnec et al. 2021: “> 4.5 mm’). Length of aedeagus 0.98-1.05 mm. PW/ PL=1.29-1.37;n=20 (Quéinnec et al. 2021: PW/PL= 1.6; this value is very probably based on a measurement error). PW/TL = 1.06—1.15; n= 20. EL/AL = 2.15-2.46; n= 10. Differential diagnosis. See Key to species of the sub- genus Abunetrechus, below. Distribution. Endemic to the Abune Yosef Mountains of the northern Ethiopian Highlands. Habitat. As in 7) (Abunetrechus) lalibelae Quéinnec & Ollivier (see below). Trechus (Abunetrechus) lalibelae Quéinnec & Ollivier Figs 19, 20, 26, 27 Trechus (Abunetrechus) lalibelae: Quéinnec et al. 2021: 21; locus typi- cus: Mt. Abuna Yosef, 12°08'32"N, 39°10'59"E. Type material. Not studied. Identification is based on the original description, including habitus and male genital figures of the type specimens (Quéinnec et al. 2021), as well as on comprehensive material collected at the type locality (see below). Additional material. 13 males, 11 females, Ethio- pia, Amhara, Mt. Abuna Yosef, N-slope, 3800-3950 m, 12°07'52"N, 39°11'39"E, 4.111.2019, leg. D. Hauth, J. Schmidt, Yeshitla M., Yitbarek W. (SCHM, NHMAA); 1 323 male, 1 female, ditto, S-slope, 3850-3900 m, 12°07'29"N, 39°11'21"E, 5.01.2019, leg. D. Hauth, J. Schmidt, Ye- shitla M., Yitbarek W. (CSCHM); 1 male, ditto, S-slope, 3700-3850 m, 12°09'10"N, 39°09'35"E, 5.III.2019, leg. D. Hauth, J. Schmidt, Yeshitla M., Yitbarek W. (CSCHM). Additions to the species description. Body length 4.2—5.1 mm (Quéinnec et al. 2021: 4.8—5.1 mm). Length of aedeagus 0.98—1.02 mm. PW/PL = 1.24—1.30; n = 20. PW/TL = 0.99-1.04; n = 20. EL/AL = 2.39-2.65; n= 10. Differential diagnosis. See Key to species of the sub- genus Abunetrechus, below. Distribution. Endemic to the Abune Yosef mountains of the northern Ethiopian Highlands. Habitat. Specimens of 7: /alibelae were found syn- topic with 7: bipartitus and T. sublaevis Raffray under stones and in humus and rotten plant material near brooks in the afroalpine zone. Trechus (Abunetrechus) habeshanicus Quéinnec & Ollivier Figs 21, 22, 28, 29 Trechus (Abunetrechus) habeshanicus: Quéinnec et al. 2021: 23; locus typicus: Guassa Plateau 10°17'19"N, 39°48'13"E. Type material. Not studied. Identification is based on the original description, including habitus and male genital figures of the type specimens (Quéinnec et al. 2021), as well as on comprehensive material collected at the type locality (see below). Additional material. 98 exx. (males, females), Ethi- opia, Amhara, northern Guassa Plateau, near Guas- sa Comm. Lodge 3330 m, 10°17'17"N, 39°47'54"E, 18.V.2022, leg. J. Schmidt, Yeshitla M. (CAF, CSCHM, NHMAA); 360 exx. (males, females), ditto, “Aste wuha” 3400 m, 10°24'N, 39°48'E, 19.V.2022, leg. J. Schmidt, Yeshitla M. (CAF, CSCHM, NHMAA); 11 exx. (males, females), ditto, “Yegana Natural Forest’, river valley, 3125 m1 0°26'03"N,_ 39547'16"E, *20/V. 2022 leg= J. Schmidt, Yeshitla M. (CSCHM). Additions to the species description. Mature species with elytra blackish brown, and with suture and first inter- val reddish brown lightened in most specimens (Quéinnec et al. 2021: “suture distinctly darkened”; this information is probably based on confusion with 7! bipartitus). Body length 4.1-5.0 mm (Quéinnec et al. 2021: 4.1-4.3 mm). Length of aedeagus 0.70—0.80 mm. PW/PL = 1.28—1.38; n=20. PW/TL = 1.08-1.13; n= 20. EL/AL = 3.00-3.33; n= 10. Differential diagnosis. See Key to species of the sub- genus Abunetrechus, below. Distribution. Endemic to the Guassa Plateau of the northern Ethiopian Highlands. Habitat. Specimens of 7? habeshanicus were found syntopic with 7? guassaensis (Quéinnec & Ollivier) and two hitherto undescribed Trechus species in humus and rotten plant material along brooks in the afromontane zone. dez.pensoft.net 324 Arnaud Faille et al.: The complex history of Ethiopian Trechina beetles Figures 17-20. 7rechus subgenus Abunetrechus nov., dorsal aspect of body (17, 19), head and pronotum (18, 20). 17, 18. 7) bipartitus Raffray; 19, 20. 7: /alibelae Quéinnec & Ollivier. The small white circles in Figs 18 and 20 mark the insertion points of the clypeal setae. dez.pensoft.net Dtsch. Entomol. Z. 70 (2) 2023, 311-335 o20 21 WW Gd 1mm Figures 21-23. Trechus subgenus Abunetrechus nov., dorsal aspect of body (21) and head (22), and anterior part of left elytron (23). 21, 22. 7. habeshaicus Quéinnec & Ollivier; 23. 7) bipartitus Raffray. The small white circles in Fig. 22 mark the insertion points of the clypeal setae; the arrow in Fig. 23 points to the insertion of the anterior elytral discal seta. Revised key to species of Abunetrechus subgen. nov. Remarks. Quéinnec et al. (2021) proposed a key to the coloration patterns of the elytra. However, we found that species of their 7rechus (s. str.) bipartitus group, which in- — these character states are unsuitable due to high intraspe- cludes the three species we here assigned to the subgenus cific variability. In addition, the pronotal proportion value Abunetrechus nov. For species’ differentiation, the authors — was probably erroneously presented by Quéinnec et al. used total body length, proportions of the pronotum and (2021; see Additions to the species descriptions, above). 1 Body more slender, with shoulders more gently rounded (Fig. 19) and with appendages slightly longer: tibiae about as long as width of pronotum (PW/TL < 1.05). Endemic to Mt. Abune Yosef ........cccccceeeesseeeeeeeeeee Trechus (Abunetrechus) lalibelae Quéinnec & Ollivier Body more robust, with shoulders broader (Figs 17, 21) and with appendages shorter: tibiae distinctly shorter than width of pronotum (EA oe IA opt eee na nee ne teat ana ee he eee Aer he ornate ConA nee he tent Urea re coh Pee rehe erence Pree eee ee eee ner tr 2 Aedeagal median lobe much smaller (length: 0.70-0.80 mm; EL/AL > 2.9), its ventral margin almost evenly curved from base to apex (lateral view, Fig. 28). Endemic to the Guassa Plateau ..........cccccsecseeeeeeees Trechus (Abunetrechus) habeshaicus Quéinnec & Ollivier Aedeagal median lobe larger (length: 0.98-1.05 mm; EL/AL < 2.5), its ventral margin almost straight near apex (lateral view, Fig. 24). ERICe Me TOs Vite A DUM OSC) canes nssracord ee se.cinme em anre eoraciare a Eek sense FEevars esac anaes Trechus (Abunetrechus) bipartitus Jeannel dez.pensoft.net 326 0.5mm Arnaud Faille et al.: The complex history of Ethiopian Trechina beetles Figures 24-29. 7rechus subgenus Abunetrechus nov., aedeagus in left lateral aspect (24, 26, 28) and dorsal aspect (25, 27, 29). 24, 25. 7: bipartitus Raffray; 26, 27. 7: lalibelae Quéinnec & Ollivier; 28, 29. 7) habeshaicus Quéinnec & Ollivier. Subgenus Minitrechus Vigna Taglianti & Magrini, 2010 Type species. 7: gypaeti Vigna Taglianti & Magrini, 2010. New synonymy. Archeotrechus Magrini, Quéinnec & Vigna Taglianti, 2012 (type species: 7: relictus Magrini, Quéinnec & Vigna Taglianti, 2012), syn. nov. Remarks. Based on the molecular data, all 7rechus species known to occur in the mountains of southern Ethiopia (Bale and Arsi Mountains, Gughe Highlands), form a monophyletic clade (Fig. 3). This clade includes species characterized by widely differing body sizes, shapes, and proportions, and by many other morpholog- ical characters, including elytral chaetotaxy, the number of dilated male protarsomeres, and the extent of the dor- sal opening of the aedeagal median lobe. Similar char- acter states can likewise be found in 7rechus sensu lato Species occurring in northern Ethiopia which, however, dez.pensoft.net do not cluster within the south Ethiopian clade. At the current state of knowledge, a morphological definition of this clade together with a differential diagnosis with respect to other species groups of 7rechus sensu lato can- not be presented here and require more comprehensive morphological investigations. For the monophyletic southern Ethiopian Trechus clade, the oldest valid species group name is Minitrechus Vigna Taglianti & Magrini, which was given for a very tiny, depigmented species from Mt. Enkuolo (Vigna Ta- glianti and Magrini 2010). The subgenus Archeotrechus Magrini, Quéinnec & Vigna Taglianti was described two years later for a likewise tiny and depigmented species from the Bale Mountains, which is additionally char- acterized by a very wide dorsal opening of aedeagus (Magrini et al. 2012). In our phylogeny, the type spe- cies of both of these subgenera cluster together within Dtsch. Entomol. Z. 70 (2) 2023, 311-335 one of the two main clades of South Ethiopian Trechus, both of which are highly supported by the molecular data (Fig. 3). Consequently, the status of Archeotrechus as a separate subgenus within 7rechus sensu lato can no lon- ger be maintained. A complete list of species we propose to summa- rize within the subgenus Minitrechus, is shown in the checklist of the Ethiopian Trechini species, see Discus- sion, below. Trechus (Minitrechus) patrizii Jeannel Figs 30-35 Trechus Patrizzi [sic!] Jeannel, 1960: 265; locus typicus: “mont Chillalo”. Trechus Patrizii Jeannel (1960): 266. Trechus patrizii Jeannel: Casale and Laneyrie (1982): 125. Trechus (s. str.) patrizzii [sic!] Jeannel: Lorenz (2005): 186. Trechus patrizii Jeannel: Ortufio and Novoa (2011): 135. Trechus (s. str.) oromiensis Magrini, Quéinnec & Vigna Taglianti, 2012: 26; locus typicus: Oromia Province, Bale massif, South of Goba, alt. about 3200 m. Trechus patrizii Jeannel: Schmidt and Faille (2018): 37. Trechus oromiensis Magrini et al.: Schmidt and Faille (2018): 37. Trechus (s. str.) patrizii Jeannel: Quéinnec et al. 2021: 65. Trechus (s. str.) oromiensis Magrini et al.: Quéinnec et al. 2021: 65. New synonymy. 7rechus patrizii Jeannel, 1960 = Trechus oromiensis Magrini, Quéinnec & Vigna Taglianti, 2012, syn. nov. Type material examined. 7rechus patrizii Jeannel: Holotype female, with label data “TYPE” (printed on red card), “A.O.I. Arussi occ. / Reg. Aselle m. 2600 ca / pend. M.te Cillalo / S. Patrizi 20:27.4.38", “Trechus / patrizii nov. /R. Jeannel det., 19” in UARK (Fig. 30). Paratype male, with label data “A.O.I. Arussi occ. / Torr. Asciabaca / S. Patrizi 28 .IV.38 /m 2500”, “Trechus / patrizii n.”, “Lectotype / E. Quéinnec dés. 1994” (printed on red card), “Trechus / patrizi1 / MNHN Paris” (printed and handwritten on red card) in MNHN (Fig. 32). Remarks. Jeannel (1960: 266) stated that the type specimen is deposited in the S.L. Straneo collection. Sig- nificant parts of the Straneo collection together with the 7. patrizii specimen cited by Jeannel (1960) are now pre- served in the UARK (M. Pavesi, pers. comm. 2018). The above cited 7’ patrizii specimen from the UARK collec- tion has thus to be considered the holotype of 7! patrizii, while the (unpublished) lectotype designation made by E. Quéinnec for the specimen preserved in the MNHN has to be considered unjustified. Trechus oromiensis Magrini et al.: Type material not studied. Identification is based on the detailed description of this distinctive taxon and comprehensive material from the type locality (see Schmidt and Faille 2018). Additional material. For comprehensive material studied see our previous paper (Schmidt and Faille 2018). Note that in this study, 77 oromiensis Magrini, Quéinnec & Vigna Taglianti was erroneously treated as a distinct yea species. In the meantime, the following additional ma- terial was available for us: Ethiopia, Oromia, SE-slope of Mt. Chillalo, Dhaba village, alt. 3200 m, 19.II.2020, 7.861644°N, 39.27711°E, leg. J. Schmidt, C. Wirkner, Yeshitla M. (1 female: CSCHM); ditto, Bale Mts., Web river N Dinsho, alt. 3000 m, 5.1.2019, 07°07'18'"N, 39°46'03"E, leg. R. Emmerich, J. Schmidt, Yeshitla M. (12 specimens: CSCHM); ditto, Bale Mts., forest re- main W Dinsho, alt. 3100 m, 8.II.2019, 07°06'16"N, 39°44'46"E, leg. R. Emmerich, J. Schmidt, Yeshitla M. (12 specimens: CSCHM); ditto, Bale Mts., Sebsebe Was- hia Forest, Salgen Valley, alt. 2720-2800 m, 3.11.2019, 07°02'08"N, 39°36'06"E, leg. R. Emmerich, J. Schmidt, Yeshitla M. (7 specimens: CSCHM); ditto, Bale Mts., Sebsebe Washia Forest, Salgen Valley, alt. 3130 m, 4 11.2019, 07°02'08"N, 39°36'06"E, leg. R. Emmerich, J. Schmidt, Yeshitla M. (30 specimens: CSCHM); ditto, Bale Mts., Angeso Valley S Goba, alt. 3050 m, 5.I.2020, 6.932923°N, 39.951341°E, leg. J. Schmidt, C. Wirkner, Yeshitla M., Yitbarek W. (53 specimens: CSCHM); dit- to, Bale Mts., Shaya Valley SW Goba, alt. 3100-3150 m, 6.11.2020, 6.991843°N, 39.884397°E, leg. J. Schmidt, C. Wirkner, Yeshitla M., Yitbarek W. (15 specimens: CSCHM). Justification of the new synonymy. Jeannel (1960) noted for his 7. patrizii the absence of pronotal basolat- eral setae as diagnostic character. Up to today, T° patrizii is considered the only Ethiopian species bearing this par- ticular character (for development of this character state in 7? amharicus Ortufio & Novoa, for which absence of pronotal basolateral setae was likewise determined, see Quéinnec et al. 2021). However, based on re-investiga- tion of the type material of 7’ patrizii, we found that a pore is present in the normal position for the basolateral seta both sides of the pronotum (Figs 30, 31). Very prob- ably, absence of these setae is based on preservation ar- tefacts. The 7. patrizii type specimens correspond in all external and genital diagnostic characters with the many specimens we previously identified as 7’ oromiensis Magrini et al. from the Bale Mts, Mt. Enkuolo and from the type locality of 7) patrizii, Mt. Chillalo (Schmidt and Faille 2018). In far most of these specimens, the pronotal basolateral seta is present but lost on one or both sides in very few cases (Figs 32, 33; Schmidt and Faille 2018: 38, figs 58-60). Consequently, we conclude junior synonymy for the taxon 7’ oromiensis Magrini, Quéinnec & Vigna Taglianti under 7! patrizii Jeannel. Diagnosis. Within the 7rechus fauna of the Bale and Arsi Mountains, 7 patrizii 1s easily recognized by ab- sence of the posterior elytral discal seta (Schmidt and Faille 2018). Beside 7: patrizii, absence of the posterior elytral discal seta is also characteristic for 7’ amharicus Ortufio & Novoa and 7? aethiopicus Alluaud. Trechus pa- trizii differs from 7: amharicus by two male protarsomeres dilated, by presence of an apical disc on aedeagal median lobe, and by very differently sclerotized endophallus (for comparison see Ortufio and Novoa 2011: 134, fig. 3b, d, and Schmidt and Faille 2018: 37, figs 64-66). Trechus patrizii differs from T- aethiopicus by the pronotum with dez.pensoft.net 328 30 ee BOs 1. Arussi OGG+ © Re Gecite ut 2000 cal & 4000 m a.s.l. are particularly species-rich areas (see summary in Quéinnec et al. 2021). Trechus patrizii is the only species known to occur on the Arsi Volcanoes as well as in the Bale Mts. We, therefore, assume that the Cenozoic uplift of the volcanic massifs on both sides of the Rift Valley led to massive, if not ex- plosive, radiation of the regional carabid beetle fauna as it was likewise described from other volcanic mountains of lower latitudes, e.g. the Canary Islands (Contreras-Diaz et al. 2007) and Hawaii (Liebherr 2015). It remains unknown whether the phylogenetic position of the enigmatic new genus Baehria contrasts with this scenario. Future studies using a larger taxon sampling are required to show wheth- er Baehria gen. nov. is a relic of a moderately old Trechina lineage with its ancestor having immigrated to the Ethio- pian Highlands at about the same time as the ancestors of the highly species diverse Abyssinotus and Minitrechus. Given the limited taxon and gene sampling our evolu- tionary scenarios have to be considered preliminary. Also, the results of our study do not allow for conclusions con- cerning the relationship between the species diverse North Ethiopian Trechus clade (Abyssinotus) and the likewise di- verse South Ethiopian clade (Minitrechus). It remains open whether the evolution of these groups results from a single dez.pensoft.net Arnaud Faille et al.: The complex history of Ethiopian Trechina beetles Oligocene-Early Miocene immigration event from the Ce- nozoic Boreal. Alternatively, independent immigrations of two ancestral trechines into two separated volcanic areas of the country, which were uplifted at different times, are pos- sible. Especially species of the Simien Mountains in Ethio- pia, including the enigmatic genus Nunbergites, representa- tives of the diverse Trechina fauna from the Elgon Massif, and additional Holarctic Trechina lineages need to be in- cluded in future molecular analyses to better understand the relationships of the Ethiopian lineages. Moreover, using a larger number of molecular markers may improve the res- olution of the more basal branches in the phylogenetic tree. Evidence emerged from our study regarding the taxo- nomic positions of most of the trechine groups known to occur in Ethiopia. We could reject an earlier hypothesis of potential relationships of some of the highland trechines with Trechodina (Quéinnec et al. 2021). Further, we could show that the species group diversity is much lower than suspected by earlier authors. Consequently, morphological character states which were used to define species groups on subtribal, genus or subgenus levels, for the Ethiopian high-altitude fauna are often unsuitable to define natural groups, e.g. bidentate or tridentate mandibles, patterns of chaetotaxy and pilosity, the number of dilated male protar- someres and the extent of the dorsal closure of the aedeagal median lobe (e.g., Pawlowski 2003; Vigna Taglianti and Magrini 2010; Magrini et al. 2012; Quéinnec et al. 2021). In fact, the molecular data demonstrate that the high mor- phological diversity in both, the northern and the southern Ethiopian TJrechus clades, Abyssinotus and Minitrechus, are the result of independent morphological radiations within these clades. These radiations might have been fos- tered by 1) the absence or very limited presence of addition- al hygrophilic ground beetles in the high-altitude forests of Ethiopia, such as Nebriini, Patrobini, Platynini and Pteros- tichini, and 11) interspecific concurrence. The latter argu- ment is supported by the fact that up to eight Abyssinotus species can be found syntopically, e.g., along slopes of Mt. Choke (own unpubl. data). Within trechine beetles, this is a unique ecologic-faunistic phenomenon, supporting Jean- nel’s observation on the abundance of 7rechus on Mount Elgon (Jeannel 1950). Detailed morphological studies, in- tensive fieldwork and a significant extension of the molec- ular databases are required to test these hypotheses and to understand the evolution of the high diversity of Trechina species and their morpho-types in the Ethiopian Highlands. Acknowledgements We thank the Ethiopian Wildlife Conservation Authority, the Ethiopian Biodiversity Institute, the College of Natural and Computational Sciences and the Department of Plant Biology and Biodiversity Management, Addis Ababa Uni- versity, the Bale Mountains National Park, the Arsi Moun- tain National Park, and the Abune Yosef, Choke Mountain and Menz Guassa Community Conservation offices, for their cooperation and kind permission to conduct field work. Many thanks are due to Katinka Thielsen (Mar- Dtsch. Entomol. Z. 70 (2) 2023, 311-335 burg), Christian Wirkner and Ralph Emmerich (Rostock), Awol Assefa, Mekbib Fekadu and Woge Abebe (Addis Ababa), for supporting or organising fieldwork in Ethio- pia. We thank the following colleagues for providing spec- imens for the molecular studies: Carmelo Andujar, Paula Arribas, Hervé Bouyon, Hervé Brustel, Teo Deli¢, Cyrille D’Haese, Vasily Grebennikov, Jifi Hajek, Pierre Moret, Michel Perreau, Alfred Newton & Margaret Thayer, Pe- dro Oromi, and Ian Townsend. We are particularly grateful to Maurizio Pavesi (Milano), for his indispensable help in finding the repository of the holotype of Trechus patrizii, and Ray Fisher (UARK) for providing photographs of this specimen. Martin Fikacek (Pragae) and Pier Mauro Gia- chino (Torino) made valuable comments to an earlier ver- sion of our manuscript which helped to improve the paper. Y.M. kindly thanks the Catholic Academic Exchange Ser- vice (KAAD) for the Ph.D. scholarship and the Amhara Agricultural Research Institute (ARARI) for maintaining his job position for the period of his study. This research was funded by the German Research Council (DFG) in the framework of the joint Ethio-European DFG Research Unit 2358 ’The Mountain Exile Hypothesis. How humans benefited from and re-shaped African high-altitude eco- systems during Quaternary climate changes” with grants OP 219/5-1, 2 to L.O. and SCHM 3005/2-1 to J.S. References Abbate E, Bruni P (2015) Geology of Ethiopia: A review and geomor- phological perspectives. In: Billi P (Ed.) Landscapes and Landforms of Ethiopia. Springer International Publishing, 33-63. https://doi. org/10.1007/978-94-017-8026-1 2 Andujar C, Serrano J, GOmez-Zurita J (2012) Winding up the molecular clock in the genus Carabus (Coleoptera: Carabidae): Assessment of methodological decisions on rate and node age estimation. 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The Open Database License (ODbL) is a license agreement intended to allow us- ers to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited. Link: https://do1.org/10.3897/dez.70.107425.suppl1 3o0 Supplementary material 2 Primers used for DNA amplification and sequencing Authors: Arnaud Faille, Sylvia Hofmann, Yeshitla Merene, David Hauth, Lars Opgenoorth, Yitbarek Woldehawariat, Joachim Schmidt Data type: docx Copyright notice: This dataset is made available under the Open Database License (http://opendatacom- mons.org/licenses/odbl/1.0). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for oth- ers, provided that the original source and author(s) are credited. Link: https://do1.org/10.3897/dez.70.107425.suppl2 dez.pensoft.net