Ecclisister bickhardti costaericae Reichensperger 1935: 204 Ecclisister costaericae E. burchellii E. burchellii E. burchellii E. burchellii E. burchellii E. burchellii E. burchellii E. burchellii E. burchellii E. burchellii E. burchellii Ecclisister species. E E. costaericae Ecclisister bickhardti costaericae Reichensperger Mazur 1984: 304 1997: 159 2011: 116 Helava et al. 1985: 324 Taxonomic Review and Natural History Notes of the Army Ant-Associated Beetle Genus Ecclisister Reichensperger (Coleoptera: Histeridae: Haeteriinae) Tishechkin, Alexey K. Kronauer, Daniel J. C. Beeren, Christoph Von The Coleopterists Bulletin 2017 2017-06-27 71 2 279 288 9LBRX Reichensperger, 1935 Reichensperger 1935 Insecta Histeridae Ecclisister Animalia Coleoptera 3 281 Arthropoda species costaericae stat. nov.      Ecclisister bickhardti costaericae Reichensperger 1935: 204.   Table 1.Information for barcoded  Ecclisister costaericaepopulations. acc. no. = accession number; AKT = Alexey K. Tishechkin; Coll. = collection; CvB = Christoph von Beeren; Dep. = depository; DK = Daniel Kronauer; GB = Griffin Burke; ID = identification number; Lat. = Latitude; Long. = Longitude; LSAM = Louisiana State Arthropod Museum; SP = Sebastian Pohl.     GenBank  Voucher   Sample ID  acc. no.  type  Dep.  Host species  Collectors  Collec. Date  Country  Region  Exact Site  Lat.  Lon.  cvb266hist002  KX420353 private CvB   E. burchellii CvB, DK, 12-Mar-2013 Costa Heredia: LSBS 10.423 -84.022  collection  foreli SP, GB Rica Canton  Sarapiqui  cvb266hist003  KX420352 private CvB   E. burchellii CvB, DK, 12-Mar-2013 Costa Heredia: LSBS 10.423 -84.022  collection  foreli SP, GB Rica Canton  Sarapiqui  cvb266hist004  KX420351 private CvB   E. burchellii CvB, DK, 12-Mar-2013 Costa Heredia: LSBS 10.423 -84.022  collection  foreli SP, GB Rica Canton  Sarapiqui  cvb285hist001  KX420350 private CvB   E. burchellii CvB, DK, 13-Mar-2013 Costa Heredia: LSBS 10.423 -84.022  collection  foreli SP, GB Rica Canton  Sarapiqui  cvb571hist005  KX420348* museum LSAM   E. burchellii CvB, DK, 08-Apr-2013 Costa Heredia: LSBS 10.423 -84.022  specimen  foreli SP, GB Rica Canton  Sarapiqui  cvb572hist001  KX420349 museum LSAM   E. burchellii CvB, DK, 08-Apr-2013 Costa Heredia: LSBS 10.423 -84.022  specimen  foreli SP, GB Rica Canton  Sarapiqui  cvb572hist003  KX420354 museum LSAM   E. burchellii CvB, DK, 08-Apr-2013 Costa Heredia: LSBS 10.423 -84.022  specimen  foreli SP, GB Rica Canton  Sarapiqui  Ecc002  KX420355 museum LSAM   E. burchellii AKT 01-Jul-2008 Ecuador Orellana Tiputini -0.638 -76.150  specimen Biodiversity  Station  Ecc005  KX420347 museum LSAM   E. burchellii AKT 28-Mar-2009 Peru Jun´ın Los Olivos -11.050 -74.259  specimen  Ecc007  KX420357 museum LSAM   E. burchellii AKT 28-Mar-2009 Peru Jun´ın Los Olivos -11.050 -74.259  specimen  Ecc008  KX420356 museum LSAM   E. burchellii AKT 28-Mar-2009 Peru Jun´ın Los Olivos -11.050 -74.259  paratype  * Record taken from von Beeren and Tishechkin (2017).   Fig. 1.  Ecclisisterspecies. E . bickhardtilectotype: a) dorsal view, b) lateral view, c) ventral view, d) frontal head, e) aedeagus, dorsal view, f) aedeagus, lateral view;  E. costaericaelectotype: g) dorsal view, h) lateral view, i) ventral view, j) frontal head, k) aedeagus, dorsal view, l) aedeagus, lateral view. Scale bars: a–c, g–i = 1 mm; d and j = 0.2 mm. Dorsal and lateral views of aedeagi are at the same scale.    Ecclisister bickhardti costaericaeReichensperger:  Mazur 1984: 304,  1997: 159,  2011: 116;  Helava et al.1985: 324  Zoobank Identifier. urn:lsid:zoobank.org:act: 920C67FF-E5B2-47A9-B2C2-31EBCCCDDEA8   Type Material Examined.   Lectotype(present designation; Fig. 1g–l). A male mounted on a point and labeled “ Costa Rica F. Nevermann  25.11.1934/  Eciton burchelli/ Type! Reichensperger /  Ecclisister bickhardti costaericae/ LECTOTYPE  Ecclisister bickhardti costaericaeReich. A. K. Tishechkin and C. von Beerendes. 2015” ( FIMAK).   Paralectotype.A specimen of undetermined sex with the same label data as lectotype, but “ Paratype! Reichensperger” instead of “ Type! Reichensperger” and our “ Paralectotype” instead of “ Lectotype” labels and an extra label “ Hamburg Farm Reventazon Ebene Limon” ( FIMAK).  Additional Material Examined.   BRAZIL(new country record): Pará: 4 specimens: Altamira – Maraba, km 18, 3°09ʹS 52°03ʹW,  14.iv.1986, in an  E. burchelliibivouac, N. Dégallier(ND); 1 specimen: Belém, Utinga( IPEAN), 1°27ʹS 48°26ʹW, 18.viii.1984, in a nest of  E. burchelliiinside a hollow tree stump, N. Dégallier (ND).   COSTA RICA: Heredia: 2 specimens: La Selva Biological Station, 10.4305°N 84.0064°W,  8.iv.2013, with  E. burchellii, C. von Beeren, G. Burke, D. Kronauer, S. Pohl (LSAM); 1 specimen: Puerto Viejo, 10°26ʹN 83°59ʹW,  2.viii.1965, with  E. burchellii, C. W. Rettenmeyer (UCS).   Limon: 29 specimens: Hamburg Farm, 10°13ʹN 83°29ʹW, multiple dates between  10.xi.1927and 1.i.1938and undated, all with  E. burchellii, F. Nevermann (FIMAK, FMNH, USNM, ZMHUB).   ECUADOR(new country record): Orellana: 1 specimen: Tiputini Biodiversity Station, 0.6376°S 76.1499°W,  31.vii.2008, with  E. burchellii, A. K. Tishechkin (LSAM).   GUYANA(new country record): Cuyuni-Mazaruni: 1 specimen: Kartabo, 6°23ʹN 58°42ʹW,  15.iv.1919(ND).   PANAMA: Colón: 3 specimens: San Lorenzo Forest, Smithsonian Research Institutioncanopy crane site, 9°17ʹN 79°58ʹW,  3.x. 2003and 21.x. 2003, with  E. burchellii, A. K. Tishechkin (LSAM).   Panamá: 1 specimen: Barro Colorado Island, 9°11ʹN 79°51ʹW, with  E. burchellii, O. Park (FMNH); 6 specimens: same locality and host data, multiple dates between  4.iii.1955and 23.iii.1966, C. W. Rettenmeyer( FMNH, UCS);  4 specimens: same locality and host data, multiple dates between  26.iii.1967and 1.v.1967, R. Akre, R. Torgerson( WSU).   PERU(new country record): Jun´ın: 5 specimens:  11 kmNW Puerto Ocopa, Los Olivos, 11°3.00ʹS 74°15.52ʹW,  1,220 m,  28. iii.2009, with  E. burchellii, riding beneath heads of major workers in emigration column, A. K. Tishechkin( LSAM).   Loreto: 1 specimen: R´ıo Sucusari, 2 kmupstream from R´ıo Napo, 3°15ʹS 73°14ʹW,  27.viii.1992, with  E. burchellii, P. E. Skelley (PK).   Host Specificity and Phoretic Transport in  Ecclisister.We found nine  E. costaericaespecimens during  Ecitoncolony emigrations at LSBS. They were collected from two of 12  E. burchellii forelicolonies, while we never found any  E. costaericaespecimens associated with other host species. However, actual prevalence of  E. costaericaein  E. burchellii forelicolonies might be higher, because the discovery rate during army ant colony emigrations is expected to be low.  Ecclisister costaericaehas been observed to be phoretic on the underside of the heads of major worker ants (AKT, personal observation), which makes it difficult for the collector to spot them. In an emigration of  E. burchellii cupiensSantschimonitored in March 2009in the Jun´ın Region of Peru, AKT noticed a lump beneath the head of a passing major worker ant, which turned out to be an attached haeteriine beetle upon inspection. Special attention to the passing major worker ants allowed the collection of multiple specimens of the genera  ClientisterReichensperger,  Ecclisister, and  SternocoelopsisReichenspergeron approximately every 40 thmajor worker ant (see also von Beeren and Tishechkin 2017). Further evidence for phoretic transport of  Ecclisisterbeetles on major army ant comes from the collection of specific army ant castes. Five of the nine specimenscollected at LSBS were found in collection vials into which only major workers had been collected, while no  Ecclisisterspecimens were found in collection vials of minor and intermediate workers. The remaining four specimenswere found in collection vials with representatives of all worker castes. No data are available about host-following mechanisms in  E. bickhardti.  Molecular Analysis. COIbarcodes were successfully obtained for four South American  E. costaericaespecimens and seven Central American  E. costaericaespecimens ( Figure 2A; for detailed collection records as well as GenBank accession numbers, see Table 1). They comprised two and three different COIhaplotypes, respectively. A RAxML tree of COIsequences revealed two distinct genetic clades reflecting the origin of specimens ( Fig. 2B). P-distances within the geographic areas were lower than 1% (range 0.0–0.5%), while they exceeded 2% (range 2.3–2.4%) in pairwise comparisons between specimens from the two geographic areas. P-distances describe the percentage of base-pair differences in two homologous DNA sequences. 1934-11-25 FIMAK F. Nevermann Costa Rica Costa Rica 5 283 2 lectotype 1934-11-25 FIMAK F. Nevermann Costa Rica Reichensperger Reichensperger Type 5 283 4 Limon paralectotype 1986-04-14 IPEAN N. Degallier & Belem & Utinga Brazil -3.15 Altamira - Maraba 1307 -52.05 5 283 5 Para 1965-08-02 2013-04-08 1965-08-02 Costa Rica 10.433333 Puerto Viejo 1298 -83.98333 La Selva Biological Station 5 283 3 Heredia 10. xi. 1927 and 1. i. 1938 Costa Rica 6 284 10.216666 Hamburg Farm 1298 -83.48333 5 283 29 Limon 2008-07-31 Ecuador -0.6376 Tiputini Biodiversity Station 7 -76.1499 6 284 1 Orellana 1919-04-15 Guyana 6.383333 Kartabo 1304 -58.7 6 284 1 Cuyuni-Mazaruni 3. x. 2003 and 21. x. 2003 Panama 9.283334 Smithsonian Research Institution 1300 -79.96667 San Lorenzo Forest 6 284 3 Colon 4. iii. 1955 and 23. iii. 1966 FMNH, UCS C. W. Rettenmeyer Panama 9.183333 Barro Colorado Island 1300 -79.85 San Lorenzo Forest 6 284 7 Panama 26. iii. 1967 and 1. v. 1967 R, WSU Akre & Torgerson Panama 9.183333 Barro Colorado Island 1300 -79.85 San Lorenzo Forest 6 284 4 Panama 2009-03-28 LSAM A. K. Tishechkin Peru 1220 -11.05 Los Olivos 13 -74.25867 Jun'in 6 284 5 1992-08-27 Peru -3.25 Napo 1307 -73.23333 Sucusari 6 284 1 Loreto