Resolving the taxonomic conundrum in Graphoderus of the east Palearctic with a key to all species (Coleoptera, Dytiscidae) Author Holmgren, Sandra Author Angus, Robert Author Jia, Fenglong Author Chen, Zhen-ning Author Bergsten, Johannes text ZooKeys 2016 574 113 142 http://dx.doi.org/10.3897/zookeys.574.7002 journal article http://dx.doi.org/10.3897/zookeys.574.7002 1313-2970-574-113 BA21C9C6CD8849F186D3A1AC01331E9A Taxon classification Animalia Coleoptera Dytiscidae Graphoderus elatus Sharp, 1882 new status Graphoderus elatus Sharp, 1882: 695 (original description); Graphoderus cinereus sensu Horn (1883) (in part); Graphoderus zonatus sensu Zimmermann (1920) (in part), Gschwendtner (1937) (in part), Kamiya (1940) , Balfour-Browne (1946) , Zaitsev (1972) , Lafer (1989) , Zeng (1989) , Mori and Kitayama (1993) , Nilsson (1995) , Nilsson et al. (1997) ; Graphoderus zonatus zonatus sensu Nilsson (2003a) (in part), Nilsson and Hajek (2015) (in part); Graphoderus zonatus verrucifer sensu Nilsson et al. (1997) , Nilsson (2003a) (in part), Nilsson and Hajek (2015) (in part); Graphoderus perplexus sensu Wallis (1939) (in part), Larson (1975) (in part), Nilsson et al. (1999) , Nilsson (2001) (in part), Nilsson (2003a) (in part), Nilsson (2015) (in part), Nilsson and Hajek (2015) (in part). Type locality. Russia> East Siberia> Amurland. Type material. Lectotype ♂ (BMNH), by present designation. Labeled: "Eastern Siberia 995 elatus . Sharp Coll. 1905-313. Data in NHRS JLKB 000023379. Lectotype Graphoderus elatus Sharp, 1882 Des. S. Holmgren et al., 2015". Paralectotype ♂ (BMNH). Labeled: "Red River. Am. Bor. 995 var. Paralectotype. Sharp Coll. 1905-313. Data in NHRS JLKB 000023380. Graphoderus perplexus Sharp 1882 Det. J. Bergsten, 2015". Lectotype justification. Sharp (1882) based his description of Graphoderus elatus on two male specimens, one from "Eastern Siberia (Amurland)" and the other from "North America (Red River)". He gave Graphoderus elatus the number 995. These two syntype specimens, present in Sharp's collection (BMNH), were studied and genitalia extracted. Both are pinned through cards, these cards being mounted on longer pins. The Siberian specimen is labeled on the face of the card, in Sharp's handwriting "Eastern Siberia 995 elatus ", while the North American specimen is labeled "Red River. Am. Bor. 995 var". The syntype specimen from Red River Am. Bor. we consider conspecific with Graphoderus perplexus . The Lectotype of Graphoderus perplexus (designated by Larson 1975 ) was studied at BMNH but is a female so the shape of male genitalia could not be compared. The characters Sharp (1882) used to distinguish Graphoderus elatus from Graphoderus perplexus are unconvincing as already argued by Wallis (1939) . The syntype specimen from Siberia has the concave outline of the penis' apex central lobe in lateral view. The penis is also of the longer type (Fig. 2k) and belongs to what we initially called Graphoderus perplexus -like specimens from the east Palearctic. We therefore designate the Siberian specimen as lectotype of Graphoderus elatus Sharp, 1882, and have so labeled it. What we initially called Graphoderus perplexus -like specimens from the east Palearctic belong to Graphoderus elatus which is here reinstated as a valid species (stat. n.). The American specimen although belonging to Graphoderus perplexus , is a paralectotype of Graphoderus elatus . As Sharp's description of Graphoderus elatus is very short and was based on a mix of two species we provide a redescription and documentation (Fig. 7) based on the designated lectotype followed by a discussion of the intraspecific variations as here interpreted from all examined material (Table 2). Figure 7. The designated lectotype for Graphoderus elatus Sharp, 1882 (BMNH). a head and pronotum (2.1 mm long, maximum width 6.5 mm) in dorsal view b ventral view (body length 14.0 mm) c penis in dorsal view (2.8 mm, long 0.55 mm width) d penis in lateral view e central penis lobe with concave dorsal margin f entire genitalia with parameres surrounding the penis g lappets of aedeagal ring. Table 2. Studied material of Graphoderus elatus . Sex, catalog number (ID), deposition, locality information, latitude (Lat.), longitude (Lon.), date collected and collector of the studied Graphoderus elatus specimens. ♂=male, ♀=female.

Sex	Catalog ID	Museum	Locality	Lat.	Lon.	Date	Collectors
NHRS	BMNH
NHRS	NHRS	51.5583°N	157.709°E	1999-07-27
NHRS	NHRS	51.5583°N	157.709°E	1999-07-27
NHRS	NHRS	51.5583°N	157.709°E	1999-07-27
NHRS	NHRS	51.5583°N	157.709°E	1999-07-27
NHRS	NHRS	51.5583°N	157.709°E	1999-07-27
NHRS	NHRS	51.5583°N	157.709°E	1999-07-27
NHRS	NHRS	53.0283°N	158.6454°E	1997-07-09
NHRS	NHRS	53.0283°N	158.6454°E	1997-07-09
NHRS	NHRS	53.0283°N	158.6454°E	1997-07-09
NHRS	NHRS	52.493°N	142.683°E	2002-07-29
NHRS	NHRS	52.493°N	142.683°E	2002-07-29
NHRS	NHRS	45.7965°N	149.9002°E	1995-08-29
NHRS	NHRS	45.7965°N	149.9002°E	1995-08-29
NHRS	NHRS	45.7965°N	149.9002°E	1995-08-28
NHRS	NHRS	45.7965°N	149.9002°E	1995-08-28
NHRS	NHRS	45.0172°N	141.8491°E
NHRS	NHRS	45.0172°N	141.8491°E
NHRS	NHRS	45.0172°N	141.8491°E	2009-09-13
NHRS	NHRS	52.0011°N	127.6842°E	1975-06-20
NHRS	ZMUM	52.0402°N	113.3856°E	1973-08-06
NHRS	ZMUM	52.0402°N	113.3856°E	1971-08-06
NHRS	ZMUM	52.0402°N	113.3856°E	1973-08-06
NHRS	OMNH	45.1059°N	141.6328°E	1987-08-01
NHRS	OMNH	45.1059°N	141.6328°E	1987-07-31
NHRS	OMNH	45.1059°N	141.6328°E	1987-07-31
NHRS	OMNH	45.1059°N	141.6328°E	1993-07-25
NHRS	OMNH	45.1059°N	141.6328°E	1993-07-25
NHRS	OMNH	45.1059°N	141.6328°E	1993-07-25
NHRS	OMNH	45.3103°N	141.6207°E	1992-08-22
NHRS	OMNH	45.0853°N	141.8197°E	1992-08-21
NHRS	OMNH	45.0853°N	141.8197°E	1992-08-21
NHRS	OMNH	45.0853°N	141.8197°E	1992-08-21
NHRS	BMNH	61.4372°N	131.0155°E	1970-07-21
NHRS	BMNH	61.4372°N	131.0155°E	1970-07-21
NHRS	BMNH	37.2952°N	100.1797°E	2013-06-05
NHRS	BMNH	37.2952°N	100.1797°E	2013-06-05
BMNH	37.2952°N	100.1797°E	2013-06-05
BMNH	37.2952°N	100.1797°E	2013-06-05
SYSU	37.2952°N	100.1797°E	2013-06-05
SYSU	37.2952°N	100.1797°E	2013-06-05
BMNH	37.2952°N	100.1797°E	2013-06-05
SYSU	37.2952°N	100.1797°E	2013-06-05
SYSU	37.2952° N	100.1797° E	2013-06-05
NHRS	ZIN	48.5941°N	110.8558°E	1987-08-29
NHRS	ZIN	48.5941°N	110.8558°E	1987-08-29
NHRS	ZIN	69.5267°N	146.6575°E	1891-07-16
ZIN	69.5267°N	146.6575°E	1891-07-16
ZIN	69.5267°N	146.6575°E	1891-07-16
NHRS	ZIN	67.8181°N	134.0181°E
BMNH	49.0892°N	127.5374°E
BMNH	49.0892°N	127.5374°E
SYSU	2013-07-22
SYSU	2014-07-24
SYSU	2014-07-24

Description of the Lectotype ♂. Body length 14.0 mm; maximum elytral width 8.3 mm. Head (Fig. 7a) dorsally testaceous; basal black band extending between eyes, retracted under anterior of pronotum, extending apically to eyes with no apparent separation from eyes; V-shaped black marking anteriorly. Head ventrally testaceous; clypeus testaceous; maxillary palpi yellow with apical palpomere piceous distally. Labial palpi yellow with apical palpomere darkened along its inner margin. Antenna testaceous with antennomeres more or less piceous in distal half. Pronotum (Fig. 7a) 2.1 mm long; maximum width 6.5 mm; testaceous; transverse black bands separated from anterior margin by testaceous band and from posterior margin by narrow testaceous, more piceous band; black bands do not reach sides of pronotum; anterior black band at edges with narrow posteriorly directed projections; posterior black band with narrow lateral portions projecting from basal margin of band. Elytron 10.5 mm long; smooth, yellow with black irrorations; irrorations reduced along margins of elytra; sutural midline black; scutellum piceous. Ventral side (Fig. 7b) testaceous-rufous, darkened due to age so that minor variations in its color not apparent. Forelegs testaceous; midlegs partly testaceous, mesotibia and mesotarsus rufous-testaceous with golden setae along edge; metatrochanter and metafemur testaceous; metatibia rufous with golden setae along edge; metatarsus rufous-piceous with long golden setae along edge. Elytral epipleuron testaceous; broad anteriorly, gradually tapering along edges of abdomen but relatively broad also in first part of posterior half. Protarsal claws similar in size and shape, shorter than protarsomere V; mesotarsal claws similar in size and shape. Posterior metatarsal claws almost three times as long as anterior metatarsal claws. Protarsomeres I-III enlarged with three larger adhesive discs basally and about 32 smaller discs distally. Mesotarsomeres with two more or less regular rows of seven discs, left mesotarsus with one additional smaller disc on mesotarsomere I and a second on mesotarsomere II and right one with only one extra disc, on mesotarsomere II. Penis in dorsal view (Fig. 7c) about 2.8 mm long; width 0.55 mm; apex trifid with three distinct lobes. Penis in lateral view (Fig. 7d) with lateral apical lobes fairly slender; central penis lobe sclerotized along edge, longer than side lobes, its dorsal margin concave (Fig. 7e). Parameres with external margins straighter medially, their apices convergent (Fig. 7f). Lappets of aedeagal ring sclerite short and wide, their outer apical margins rounded (Fig. 7g). Intraspecific variation. Body length between 13.9 and 16.3 mm; maximum elytral width between 8.0 and 9.6 mm. Pronotum length 1.8 to 2.5 mm long; width 5.3 to 7.2 mm; smooth in males; either deeply wrinkled (when also elytra granulated) or smooth in females; in smooth specimens either shining with anterior row of impressed punctures very distinct, or matt with puncture-row less distinct; anterior black band of pronotum mostly continuous, sometimes thin and weak or non-continuous; shape of posterior black band of pronotum varies, separated from posterior margin by testaceous band which sometimes is partly piceous. Elytron between 10.4 and 12.7 mm long; smooth in males; smooth or granulated in females. Male posterior metatarsal claws almost three times as long as anterior metatarsal claws; female posterior metatarsal claws less than twice as long as anterior metatarsal claws which are slightly curved apically. Protarsomeres I-III enlarged in males with three larger adhesive discs basally and 28-66 smaller discs distally; mesotarsomeres in males with irregular rows of 14-31 adhesive discs; in populations with granulated females, number of adhesive discs in males are in upper range. Penis in dorsal view between 2.4 and 2.8 mm long; width between 0.6 and 0.7 mm. Shape of lappets in aedeagal ring sclerite variable which also applies to outer apical margin. Distribution. (Fig. 8) The distribution covers Russia, east of the Yenisei-Angara river to the pacific coast, north to the East Siberian Sea and south to Qinghai in China and Hokkaido in Japan. Specimens from the following regions in east Palearctic were examined: Russia: Yakutia, Kamchatka, Chita Region, Amur Region, Kuril Islands, North Sakhalin, Verkhoyansk and Indigirka. Japan: Hokkaido. Mongolia: Onon river. China: Heilongjiang, Inner Mongolia and Qinghai. Figure 8. Distribution map of Graphoderus elatus based on examined specimens. The lectotype is imprecisely marked in former Amur Region.