taxonID	type	format	identifier	references	title	description	created	creator	contributor	publisher	audience	source	license	rightsHolder	datasetID
052D87A15861131CFE8CAA7EFE2FFA19.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/3666542/files/figure.png	https://doi.org/10.5281/zenodo.3666542	Figures 2–5. Habitus of the Grouvellinus species collected from the Tawan Hills Park. 2: G. andrekuipersi; 3: G. nigerquadratus, sp. nov. (paratype male from Tawau River); 4: G. nigerquadratus, sp. nov. (male from Lucia River); 5: G.cruxniger, sp. nov. (paratype male from Lucia River). Scale: 1 mm.	Figures 2–5. Habitus of the Grouvellinus species collected from the Tawan Hills Park. 2: G. andrekuipersi; 3: G. nigerquadratus, sp. nov. (paratype male from Tawau River); 4: G. nigerquadratus, sp. nov. (male from Lucia River); 5: G.cruxniger, sp. nov. (paratype male from Lucia River). Scale: 1 mm.	2020-01-24	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo		Zenodo	biologists	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo			
052D87A15861131CFE8CAA7EFE2FFA19.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/3666562/files/figure.png	https://doi.org/10.5281/zenodo.3666562	Figure 18. Map of Sabah and Brunei (Borneo) with collecting site of Grouvellinus species with new species record in red fonts.	Figure 18. Map of Sabah and Brunei (Borneo) with collecting site of Grouvellinus species with new species record in red fonts.	2020-01-24	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo		Zenodo	biologists	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo			
052D87A1586C1318FE9AAC99FEEDFD1D.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/3666542/files/figure.png	https://doi.org/10.5281/zenodo.3666542	Figures 2–5. Habitus of the Grouvellinus species collected from the Tawan Hills Park. 2: G. andrekuipersi; 3: G. nigerquadratus, sp. nov. (paratype male from Tawau River); 4: G. nigerquadratus, sp. nov. (male from Lucia River); 5: G.cruxniger, sp. nov. (paratype male from Lucia River). Scale: 1 mm.	Figures 2–5. Habitus of the Grouvellinus species collected from the Tawan Hills Park. 2: G. andrekuipersi; 3: G. nigerquadratus, sp. nov. (paratype male from Tawau River); 4: G. nigerquadratus, sp. nov. (male from Lucia River); 5: G.cruxniger, sp. nov. (paratype male from Lucia River). Scale: 1 mm.	2020-01-24	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo		Zenodo	biologists	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo			
052D87A1586C1318FE9AAC99FEEDFD1D.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/3666546/files/figure.png	https://doi.org/10.5281/zenodo.3666546	Figure 10. Grouvellinus nigerquadratus, sp. nov. (paratypes): (a) anteriodorsal aspect with pronotum; (b) anterioventral aspect with prosternum; (c) posterioventral aspect with meso-, metaventrite, and ventrites 1–5.	Figure 10. Grouvellinus nigerquadratus, sp. nov. (paratypes): (a) anteriodorsal aspect with pronotum; (b) anterioventral aspect with prosternum; (c) posterioventral aspect with meso-, metaventrite, and ventrites 1–5.	2020-01-24	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo		Zenodo	biologists	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo			
052D87A1586C1318FE9AAC99FEEDFD1D.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/3666540/files/figure.png	https://doi.org/10.5281/zenodo.3666540	Figure 1. (a) Co-authors and assistants of the Taxon Expedition to Tawau Hills Park, Sabah, Malaysian Borneo, performing microhabitat-specific sampling; (b) Co-authors during beetle collection at a blacklight trap installed in the vicinity of the Tawau River; (c) The first author (second from the right) instructing Taxon Expedition participants on the identification of riffle beetles; (d) the transportable molecular field laboratory at the Tawau Park Station; (e) Type locality of Grouvellinus borneensis, sp. nov., G. cruxniger, sp. nov., G. nigerquadratus, sp. nov., and collection site of G. sp. 1; (f) Type locality of Grouvellinus luciaensis, sp. nov., and collection site of G. andrekuipersi, G. nigerquadratus, sp. nov. and G. sp. 2.	Figure 1. (a) Co-authors and assistants of the Taxon Expedition to Tawau Hills Park, Sabah, Malaysian Borneo, performing microhabitat-specific sampling; (b) Co-authors during beetle collection at a blacklight trap installed in the vicinity of the Tawau River; (c) The first author (second from the right) instructing Taxon Expedition participants on the identification of riffle beetles; (d) the transportable molecular field laboratory at the Tawau Park Station; (e) Type locality of Grouvellinus borneensis, sp. nov., G. cruxniger, sp. nov., G. nigerquadratus, sp. nov., and collection site of G. sp. 1; (f) Type locality of Grouvellinus luciaensis, sp. nov., and collection site of G. andrekuipersi, G. nigerquadratus, sp. nov. and G. sp. 2.	2020-01-24	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo		Zenodo	biologists	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo			
052D87A1586C1318FE9AAC99FEEDFD1D.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/3666548/files/figure.png	https://doi.org/10.5281/zenodo.3666548	Figure 11. Grouvellinus nigerquadratus, sp. nov. (paratypes): (a) aedeagus, lateral; (b) aedeagus, ventral; (c) ovipositor (half), ventral. Scale bar = 0.1 mm.	Figure 11. Grouvellinus nigerquadratus, sp. nov. (paratypes): (a) aedeagus, lateral; (b) aedeagus, ventral; (c) ovipositor (half), ventral. Scale bar = 0.1 mm.	2020-01-24	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo		Zenodo	biologists	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo			
052D87A1586C1318FE9AAC99FEEDFD1D.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/3666562/files/figure.png	https://doi.org/10.5281/zenodo.3666562	Figure 18. Map of Sabah and Brunei (Borneo) with collecting site of Grouvellinus species with new species record in red fonts.	Figure 18. Map of Sabah and Brunei (Borneo) with collecting site of Grouvellinus species with new species record in red fonts.	2020-01-24	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo		Zenodo	biologists	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo			
052D87A158681304FEB6AB8CFE9BFD65.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/3666542/files/figure.png	https://doi.org/10.5281/zenodo.3666542	Figures 2–5. Habitus of the Grouvellinus species collected from the Tawan Hills Park. 2: G. andrekuipersi; 3: G. nigerquadratus, sp. nov. (paratype male from Tawau River); 4: G. nigerquadratus, sp. nov. (male from Lucia River); 5: G.cruxniger, sp. nov. (paratype male from Lucia River). Scale: 1 mm.	Figures 2–5. Habitus of the Grouvellinus species collected from the Tawan Hills Park. 2: G. andrekuipersi; 3: G. nigerquadratus, sp. nov. (paratype male from Tawau River); 4: G. nigerquadratus, sp. nov. (male from Lucia River); 5: G.cruxniger, sp. nov. (paratype male from Lucia River). Scale: 1 mm.	2020-01-24	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo		Zenodo	biologists	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo			
052D87A158681304FEB6AB8CFE9BFD65.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/3666550/files/figure.png	https://doi.org/10.5281/zenodo.3666550	Figure 12. Grouvellinus cruxniger, sp. nov. (paratypes): (a) anteriodorsal aspect with pronotum and elytra; (b) ventral aspect with prosternum, meso-, metaventrite, and ventrites 1–5.	Figure 12. Grouvellinus cruxniger, sp. nov. (paratypes): (a) anteriodorsal aspect with pronotum and elytra; (b) ventral aspect with prosternum, meso-, metaventrite, and ventrites 1–5.	2020-01-24	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo		Zenodo	biologists	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo			
052D87A158681304FEB6AB8CFE9BFD65.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/3666540/files/figure.png	https://doi.org/10.5281/zenodo.3666540	Figure 1. (a) Co-authors and assistants of the Taxon Expedition to Tawau Hills Park, Sabah, Malaysian Borneo, performing microhabitat-specific sampling; (b) Co-authors during beetle collection at a blacklight trap installed in the vicinity of the Tawau River; (c) The first author (second from the right) instructing Taxon Expedition participants on the identification of riffle beetles; (d) the transportable molecular field laboratory at the Tawau Park Station; (e) Type locality of Grouvellinus borneensis, sp. nov., G. cruxniger, sp. nov., G. nigerquadratus, sp. nov., and collection site of G. sp. 1; (f) Type locality of Grouvellinus luciaensis, sp. nov., and collection site of G. andrekuipersi, G. nigerquadratus, sp. nov. and G. sp. 2.	Figure 1. (a) Co-authors and assistants of the Taxon Expedition to Tawau Hills Park, Sabah, Malaysian Borneo, performing microhabitat-specific sampling; (b) Co-authors during beetle collection at a blacklight trap installed in the vicinity of the Tawau River; (c) The first author (second from the right) instructing Taxon Expedition participants on the identification of riffle beetles; (d) the transportable molecular field laboratory at the Tawau Park Station; (e) Type locality of Grouvellinus borneensis, sp. nov., G. cruxniger, sp. nov., G. nigerquadratus, sp. nov., and collection site of G. sp. 1; (f) Type locality of Grouvellinus luciaensis, sp. nov., and collection site of G. andrekuipersi, G. nigerquadratus, sp. nov. and G. sp. 2.	2020-01-24	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo		Zenodo	biologists	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo			
052D87A158681304FEB6AB8CFE9BFD65.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/3666552/files/figure.png	https://doi.org/10.5281/zenodo.3666552	Figure 13. Grouvellinus cruxniger, sp. nov. (paratypes): (a) aedeagus, lateral; (b) aedeagus, ventral; (c) ovipositor (half), ventral. Scale bar = 0.1 mm.	Figure 13. Grouvellinus cruxniger, sp. nov. (paratypes): (a) aedeagus, lateral; (b) aedeagus, ventral; (c) ovipositor (half), ventral. Scale bar = 0.1 mm.	2020-01-24	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo		Zenodo	biologists	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo			
052D87A158681304FEB6AB8CFE9BFD65.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/3666562/files/figure.png	https://doi.org/10.5281/zenodo.3666562	Figure 18. Map of Sabah and Brunei (Borneo) with collecting site of Grouvellinus species with new species record in red fonts.	Figure 18. Map of Sabah and Brunei (Borneo) with collecting site of Grouvellinus species with new species record in red fonts.	2020-01-24	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo		Zenodo	biologists	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo			
052D87A158741300FEB9AA7AFDE2FD60.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/3666544/files/figure.png	https://doi.org/10.5281/zenodo.3666544	Figures 6–9. Habitus of the Grouvellinus species collected from the Tawan Hills Park. 6: G. luciaensis, sp. nov. (paratype male from Lucia River); 7: G. borneensis, sp. nov. (paratype male from Tawau River environment); 8: G. sp.1 (female from Tawau River environment); 9: G. sp.2 (female from Lucia River). Scale: 1 mm.	Figures 6–9. Habitus of the Grouvellinus species collected from the Tawan Hills Park. 6: G. luciaensis, sp. nov. (paratype male from Lucia River); 7: G. borneensis, sp. nov. (paratype male from Tawau River environment); 8: G. sp.1 (female from Tawau River environment); 9: G. sp.2 (female from Lucia River). Scale: 1 mm.	2020-01-24	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo		Zenodo	biologists	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo			
052D87A158741300FEB9AA7AFDE2FD60.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/3666554/files/figure.png	https://doi.org/10.5281/zenodo.3666554	Figure 14. Grouvellinus luciaensis, sp. nov. (paratypes): (a) anteriodorsal aspect with pronotum; (b) anterioventral aspect with head and prosternum; (c) ventral aspect with prosternum and mesoventrite, (d) posterioventral aspect with metaventrite and ventrites 1–5.	Figure 14. Grouvellinus luciaensis, sp. nov. (paratypes): (a) anteriodorsal aspect with pronotum; (b) anterioventral aspect with head and prosternum; (c) ventral aspect with prosternum and mesoventrite, (d) posterioventral aspect with metaventrite and ventrites 1–5.	2020-01-24	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo		Zenodo	biologists	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo			
052D87A158741300FEB9AA7AFDE2FD60.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/3666540/files/figure.png	https://doi.org/10.5281/zenodo.3666540	Figure 1. (a) Co-authors and assistants of the Taxon Expedition to Tawau Hills Park, Sabah, Malaysian Borneo, performing microhabitat-specific sampling; (b) Co-authors during beetle collection at a blacklight trap installed in the vicinity of the Tawau River; (c) The first author (second from the right) instructing Taxon Expedition participants on the identification of riffle beetles; (d) the transportable molecular field laboratory at the Tawau Park Station; (e) Type locality of Grouvellinus borneensis, sp. nov., G. cruxniger, sp. nov., G. nigerquadratus, sp. nov., and collection site of G. sp. 1; (f) Type locality of Grouvellinus luciaensis, sp. nov., and collection site of G. andrekuipersi, G. nigerquadratus, sp. nov. and G. sp. 2.	Figure 1. (a) Co-authors and assistants of the Taxon Expedition to Tawau Hills Park, Sabah, Malaysian Borneo, performing microhabitat-specific sampling; (b) Co-authors during beetle collection at a blacklight trap installed in the vicinity of the Tawau River; (c) The first author (second from the right) instructing Taxon Expedition participants on the identification of riffle beetles; (d) the transportable molecular field laboratory at the Tawau Park Station; (e) Type locality of Grouvellinus borneensis, sp. nov., G. cruxniger, sp. nov., G. nigerquadratus, sp. nov., and collection site of G. sp. 1; (f) Type locality of Grouvellinus luciaensis, sp. nov., and collection site of G. andrekuipersi, G. nigerquadratus, sp. nov. and G. sp. 2.	2020-01-24	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo		Zenodo	biologists	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo			
052D87A158741300FEB9AA7AFDE2FD60.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/3666556/files/figure.png	https://doi.org/10.5281/zenodo.3666556	Figure 15. Grouvellinus luciaensis, sp. nov. (paratypes): (a) aedeagus, lateral; (b) aedeagus, ventral; (c) ovipositor (half), ventral. Scale bar = 0.1 mm.	Figure 15. Grouvellinus luciaensis, sp. nov. (paratypes): (a) aedeagus, lateral; (b) aedeagus, ventral; (c) ovipositor (half), ventral. Scale bar = 0.1 mm.	2020-01-24	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo		Zenodo	biologists	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo			
052D87A158741300FEB9AA7AFDE2FD60.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/3666562/files/figure.png	https://doi.org/10.5281/zenodo.3666562	Figure 18. Map of Sabah and Brunei (Borneo) with collecting site of Grouvellinus species with new species record in red fonts.	Figure 18. Map of Sabah and Brunei (Borneo) with collecting site of Grouvellinus species with new species record in red fonts.	2020-01-24	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo		Zenodo	biologists	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo			
052D87A15870130CFEBEAA71FB83FD6A.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/3666544/files/figure.png	https://doi.org/10.5281/zenodo.3666544	Figures 6–9. Habitus of the Grouvellinus species collected from the Tawan Hills Park. 6: G. luciaensis, sp. nov. (paratype male from Lucia River); 7: G. borneensis, sp. nov. (paratype male from Tawau River environment); 8: G. sp.1 (female from Tawau River environment); 9: G. sp.2 (female from Lucia River). Scale: 1 mm.	Figures 6–9. Habitus of the Grouvellinus species collected from the Tawan Hills Park. 6: G. luciaensis, sp. nov. (paratype male from Lucia River); 7: G. borneensis, sp. nov. (paratype male from Tawau River environment); 8: G. sp.1 (female from Tawau River environment); 9: G. sp.2 (female from Lucia River). Scale: 1 mm.	2020-01-24	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo		Zenodo	biologists	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo			
052D87A15870130CFEBEAA71FB83FD6A.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/3666558/files/figure.png	https://doi.org/10.5281/zenodo.3666558	Figure 16. Grouvellinus borneensis, sp. nov. (paratypes): (a) anteriodorsal aspect with pronotum and elytra; (b) ventral aspect with prosternum and mesoventrite; (c) posteriodorsal aspect with metaventrite and ventrites 1–5.	Figure 16. Grouvellinus borneensis, sp. nov. (paratypes): (a) anteriodorsal aspect with pronotum and elytra; (b) ventral aspect with prosternum and mesoventrite; (c) posteriodorsal aspect with metaventrite and ventrites 1–5.	2020-01-24	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo		Zenodo	biologists	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo			
052D87A15870130CFEBEAA71FB83FD6A.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/3666540/files/figure.png	https://doi.org/10.5281/zenodo.3666540	Figure 1. (a) Co-authors and assistants of the Taxon Expedition to Tawau Hills Park, Sabah, Malaysian Borneo, performing microhabitat-specific sampling; (b) Co-authors during beetle collection at a blacklight trap installed in the vicinity of the Tawau River; (c) The first author (second from the right) instructing Taxon Expedition participants on the identification of riffle beetles; (d) the transportable molecular field laboratory at the Tawau Park Station; (e) Type locality of Grouvellinus borneensis, sp. nov., G. cruxniger, sp. nov., G. nigerquadratus, sp. nov., and collection site of G. sp. 1; (f) Type locality of Grouvellinus luciaensis, sp. nov., and collection site of G. andrekuipersi, G. nigerquadratus, sp. nov. and G. sp. 2.	Figure 1. (a) Co-authors and assistants of the Taxon Expedition to Tawau Hills Park, Sabah, Malaysian Borneo, performing microhabitat-specific sampling; (b) Co-authors during beetle collection at a blacklight trap installed in the vicinity of the Tawau River; (c) The first author (second from the right) instructing Taxon Expedition participants on the identification of riffle beetles; (d) the transportable molecular field laboratory at the Tawau Park Station; (e) Type locality of Grouvellinus borneensis, sp. nov., G. cruxniger, sp. nov., G. nigerquadratus, sp. nov., and collection site of G. sp. 1; (f) Type locality of Grouvellinus luciaensis, sp. nov., and collection site of G. andrekuipersi, G. nigerquadratus, sp. nov. and G. sp. 2.	2020-01-24	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo		Zenodo	biologists	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo			
052D87A15870130CFEBEAA71FB83FD6A.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/3666560/files/figure.png	https://doi.org/10.5281/zenodo.3666560	Figure 17. Grouvellinus borneensis, sp. nov. (paratypes): (a) aedeagus, lateral; (b) aedeagus, ventral; (c) ovipositor (half), ventral. Scale bar = 0.1 mm.	Figure 17. Grouvellinus borneensis, sp. nov. (paratypes): (a) aedeagus, lateral; (b) aedeagus, ventral; (c) ovipositor (half), ventral. Scale bar = 0.1 mm.	2020-01-24	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo		Zenodo	biologists	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo			
052D87A15870130CFEBEAA71FB83FD6A.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/3666562/files/figure.png	https://doi.org/10.5281/zenodo.3666562	Figure 18. Map of Sabah and Brunei (Borneo) with collecting site of Grouvellinus species with new species record in red fonts.	Figure 18. Map of Sabah and Brunei (Borneo) with collecting site of Grouvellinus species with new species record in red fonts.	2020-01-24	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo		Zenodo	biologists	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo			
052D87A1587C130CFDADAA7AFE97FA2C.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/3666544/files/figure.png	https://doi.org/10.5281/zenodo.3666544	Figures 6–9. Habitus of the Grouvellinus species collected from the Tawan Hills Park. 6: G. luciaensis, sp. nov. (paratype male from Lucia River); 7: G. borneensis, sp. nov. (paratype male from Tawau River environment); 8: G. sp.1 (female from Tawau River environment); 9: G. sp.2 (female from Lucia River). Scale: 1 mm.	Figures 6–9. Habitus of the Grouvellinus species collected from the Tawan Hills Park. 6: G. luciaensis, sp. nov. (paratype male from Lucia River); 7: G. borneensis, sp. nov. (paratype male from Tawau River environment); 8: G. sp.1 (female from Tawau River environment); 9: G. sp.2 (female from Lucia River). Scale: 1 mm.	2020-01-24	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo		Zenodo	biologists	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo			
052D87A1587C130CFDADAA7AFE97FA2C.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/3666558/files/figure.png	https://doi.org/10.5281/zenodo.3666558	Figure 16. Grouvellinus borneensis, sp. nov. (paratypes): (a) anteriodorsal aspect with pronotum and elytra; (b) ventral aspect with prosternum and mesoventrite; (c) posteriodorsal aspect with metaventrite and ventrites 1–5.	Figure 16. Grouvellinus borneensis, sp. nov. (paratypes): (a) anteriodorsal aspect with pronotum and elytra; (b) ventral aspect with prosternum and mesoventrite; (c) posteriodorsal aspect with metaventrite and ventrites 1–5.	2020-01-24	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo		Zenodo	biologists	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo			
052D87A1587C130DFDADAC83FE08FE0E.taxon	http://purl.org/dc/dcmitype/StillImage	image/png	https://zenodo.org/record/3666544/files/figure.png	https://doi.org/10.5281/zenodo.3666544	Figures 6–9. Habitus of the Grouvellinus species collected from the Tawan Hills Park. 6: G. luciaensis, sp. nov. (paratype male from Lucia River); 7: G. borneensis, sp. nov. (paratype male from Tawau River environment); 8: G. sp.1 (female from Tawau River environment); 9: G. sp.2 (female from Lucia River). Scale: 1 mm.	Figures 6–9. Habitus of the Grouvellinus species collected from the Tawan Hills Park. 6: G. luciaensis, sp. nov. (paratype male from Lucia River); 7: G. borneensis, sp. nov. (paratype male from Tawau River environment); 8: G. sp.1 (female from Tawau River environment); 9: G. sp.2 (female from Lucia River). Scale: 1 mm.	2020-01-24	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo		Zenodo	biologists	Freitag, Hendrik;Molls, Christian;Bouma, Aglaia M.;Garces, Jhoana M.;Rossato, Marzia;Cosentino, Emanuela;Delledonne, Massimo			
