(C) 2013 Mladen Kučinić. This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
For reference, use of the paginated PDF or printed version of this article is recommended.
Citation: Kučinić M, Szivák I, Pauls SU, Bálint M, Delić A, Vučković I (2013) Chaetopteryx bucari sp. n., a new species from the Chaetopteryx rugulosa group from Croatia (Insecta, Trichoptera, Limnephilidae) with molecular, taxonomic and ecological notes on the group. ZooKeys 320: 1–28. doi: 10.3897/zookeys.320.4565
We describe a new autumnal caddisfly species Chaetopteryx bucari sp. n. from 8 localities in the Banovina region of Croatia. We also present molecular, taxonomic and ecological notes (emergence, sex ratio and seasonal dynamics) on the new species and discuss the distribution of Chaetopteryx species in general and the Chaetopteryx rugulosa group in particular. Based on Bayesian phylogenetic analysis Chaetopteryx rugulosa schmidi was separated from the clade containing the other subspecies of Chaetopteryx rugulosa. Thus the subspecies Chaetopteryx rugulosa schmidi is here raised to species level, Chaetopteryx schmidi, as it was described originally. We further present distribution data on rare species in the genus Chaetopteryx in Croatia.
Chaetopteryx, aquatic insects, new species, distribution, Croatia
The genus Chaetopteryx belongs to a small number of caddisfly genera with adults that are adapted to low air temperatures and emerge in autumn or winter, mostly from October-January. The larvae of most species live in small headwater streams and springs. This genus is distributed in Europe and parts of Asia (e.g., Asia Minor, Iran) (
Four years ago we started systematically collecting adults of the genus Chaetopteryx, including members of the Chaetopteryx rugulosa group in Croatia. This paper has 2 main objectives, first to present and describe a new species from the Chaetopteryx rugulosa group found in Croatia, and second to present new molecular, taxonomic, distributional, and ecological information on the Chaetopteryx rugulosa group.
Fieldwork. We collected specimens of Chaetopteryx including Chaetopteryx rugulosa group species in the continental (central Croatia, Banovina, Hrvatsko zagorje, Kordun, Slavonia), mountain (Gorski kotar, Lika regions) and Mediterranean (Istria and Dalmatia) regions of Croatia. Collecting methods included the use of entomological nets and handpicking specimens from walls of small buildings or wells, or from the riparian vegetation near springs and headwater streams. In one spring (Pecki spring, Banovina region) (Table 1) we installed 5 pyramid-type emergence traps in 2010 and 2011 to investigate the emergence dynamics of caddisflies (Figure 1). This investigation is part of a multi-year study on emergence dynamics of aquatic insects in springs and other aquatic habitats in Croatia and the Dinaric karst of the Balkan Peninsula (Bosnia and Herzegovina) (
Localities where Chaetopteryx bucari sp. n., was collected, including habitat type, elevation (m a.s.l.), and geographic coordinates.
Location | Character of location | Altitude (m) | Geographic coordinates |
---|---|---|---|
Bijele stijene | wellspring and stream | 144 | 45°25'23"N, 16°13'23"E |
Gore | wellspring | 165 | 45°24'21"N, 16°14'22"E |
Hrvatski Čuntić | stream | 159 | 45°21'28"N, 16°17'04"E |
Marića točak | wellspring | 163 | 45°21'29"N, 16°17'03"E |
Pašino vrelo | spring | 185 | 45°17'16"N, 16°25'13"E |
Pecki | spring | 161 | 45°23'50"N, 16°14'40"E |
Slabinja | wellspring | 104 | 45°13'05"N, 16°37'52"E |
Varoški bunar | wellspring | 130 | 45°13'34"N, 16°33'12"E |
Type locality of Chaetopteryx bucari sp. n., showing pyramid-type emergence traps, Pecki spring, Croatia.
In pyramid-type emergence traps caddisflies were collected in 1% formaldehyde and thereafter stored in 80% alcohol. All other collected specimens were stored directly in 80% or 96% alcohol. All specimens were deposited in the collections of the first and second authors. The holotype is deposited in the Croatian Natural History Museum in Zagreb.
Laboratory work. For the phylogenetic analysis we compiled mtCOI DNA sequence data for 103 specimens from the Chaetopteryx rugulosa group (Table 2). We also sequenced several outgroup taxa of varying putative phylogenetic depths including congeneric species (e.g., Chaetopteryx gessneri McLachlan, 1876, Chaetopteryx fusca Brauer, 1857, Chaetopteryx major McLachalan, 1876, Chaetopteryx villosa (Fabricius, 1798)), other members of the tribe Chaetopterygini (Chaetopterygopsis maclachlani (Stein, 1874)), other members of the subfamily Limnephilinae (Limnephilus centralis Curtis, 1834), and members of a different subfamily of Limnephilidae (e.g. Metanoea rhaetica Schmid, 1955, Drusus alpinus (Meyer-Dür, 1875), Drusus rectus McLachlan, 1868).
Systematic presentation follows
List of species included in the DNA analysis (mtCOI sequences). Localities are given with country code, locality/specimen data, and collection date.
Species name | Locality | Specimen ID | Accession number | Collectors/Source |
---|---|---|---|---|
Chaetopteryx aproka | ROU, Ignis Mts., springs near Desesti-Statiunea Izvoare, 21.10.2010 | CAxJC0101 | HE858253 | Ecsedi, Olah & Szivak |
Chaetopteryx aproka | ROU, Ignis Mts., springs near Desesti-Statiunea Izvoare, 21.10.2010 | CAxJC0102 | HE858254 | Ecsedi, Olah & Szivak |
Chaetopteryx aproka | ROU, Ignis Mts., springs near Desesti-Statiunea Izvoare, 21.10.2010 | CAxJC0103 | HE858255 | Ecsedi, Olah & Szivak |
Chaetopteryx bosniaca | BIH, Livno, Sturba river, 08.11.2009 | CBxED0101 | Kučinić, Delić & Mihoci | |
Chaetopteryx bosniaca | BIH, Livno, Sturba river, 08.11.2009 | CBxED0102 | Kučinić, Delić & Mihoci | |
Chaetopteryx bosniaca | BIH, Livno, Sturba river, 08.11.2009 | CBxED0103 | Kučinić, Delić & Mihoci | |
Chaetopteryx bosniaca | BIH, Livno, Sturba river, 08.11.2009 | CBxED0104 | Kučinić, Delić & Mihoci | |
Chaetopteryx bosniaca | BIH, Livno, Sturba river, 08.11.2009 | CBxED0105 | Kučinić, Delić & Mihoci | |
Chaetopteryx clara | SLO, Ljubljana, Mostec park, Przanec stream, 06.12.2009 | CCxEA0101 | JF891164 | Dery & Szivak |
Chaetopteryx clara | SLO, Ljubljana, Mostec park, Przanec stream, 06.12.2009 | CCxEA0102 | JF891165 | Dery & Szivak |
Chaetopteryx clara | SLO, Ljubljana, Mostec park, Przanec stream, 06.12.2009 | CCxEA0103 | JF891166 | Dery & Szivak |
Chaetopteryx clara | SLO, Ljubljana, Mostec park, Przanec stream, 06.12.2009 | CCxEA0104 | JF891167 | Dery & Szivak |
Chaetopteryx clara | SLO, Ljubljana, Mostec park, Przanec stream, 06.12.2009 | CCxEA0105 | JF891168 | Dery & Szivak |
Chaetopteryx goricensis | SLO, spring of Lokavscek stream near Predmeja, 06.12.2009 | CGREG0101 | JF891159 | Dery & Szivak |
Chaetopteryx goricensis | SLO, spring of Lokavscek stream near Predmeja, 06.12.2009 | CGREG0102 | JF891160 | Dery & Szivak |
Chaetopteryx goricensis | SLO, spring of Lokavscek stream near Predmeja, 06.12.2009 | CGREG0103 | JF891161 | Dery & Szivak |
Chaetopteryx goricensis | SLO, spring of Lokavscek stream near Predmeja, 06.12.2009 | CGREG0104 | JF891162 | Dery & Szivak |
Chaetopteryx goricensis | SLO, spring of Lokavscek stream near Predmeja, 06.12.2009 | CGREG0105 | JF891163 | Dery & Szivak |
Chaetopteryx goricensis | SLO, spring near Čekovnik (Hlevise), 05.12.2009 | CGREG0201 | JF891154 | Dery & Szivak |
Chaetopteryx goricensis | SLO, spring near Čekovnik (Blask), 05.12.2009 | CGREG0301 | JF891155 | Dery & Szivak |
Chaetopteryx goricensis | SLO, spring near Čekovnik (Blask), 05.12.2009 | CGREG0302 | JF891156 | Dery & Szivak |
Chaetopteryx goricensis | SLO, spring near Čekovnik (Blask), 05.12.2009 | CGREG0303 | JF891157 | Dery & Szivak |
Chaetopteryx goricensis | SLO, spring near Čekovnik (Blask), 05.12.2009 | CGREG0304 | JF891158 | Dery & Szivak |
Chaetopteryx irenae | SLO, Susica stream near Misliče, 06.12.2009 | CIxEI0101 | JF891169 | Dery & Szivak |
Chaetopteryx irenae | SLO, Susica stream near Misliče, 06.12.2009 | CIxEI0102 | JF891170 | Dery & Szivak |
Chaetopteryx irenae | SLO, Misliče, Susica stream, 06.12.2009 | CIxEI0103 | JF891171 | Dery & Szivak |
Chaetopteryx irenae | SLO, Misliče, Susica stream, 06.12.2009 | CIxEI0104 | JF891172 | Dery & Szivak |
Chaetopteryx irenae | SLO, Misliče, Susica stream, 06.12.2009 | CIxEI0105 | JF891173 | Dery & Szivak |
Chaetopteryx major | HUN, Mecsek Mts., Vár valley, Pásztor spring 05.11.2010 | CMJKB0101 | JF891233 | Olah, Szivak & Uherkovich |
Chaetopteryx major | HUN, Mecsek Mts., Vár valley, Pásztor spring 05.11.2010 | CMJKB0102 | HE858256 | Olah, Szivak & Uherkovich |
Chaetopteryx major | HUN, Mecsek Mts., Vár valley, Pásztor spring 05.11.2010 | CMJKB0103 | HE858257 | Olah, Szivak & Uherkovich |
Chaetopteryx major | HUN, Mecsek Mts., Vár valley, Pásztor spring 05.11.2010 | CMJKB0104 | HE858258 | Olah, Szivak & Uherkovich |
Chaetopteryx major | AUT, valley Hottmannsgraben, Unteraspang (Aspang Markt) 19.11.2009 | CMJDJ0101 | JF891234 | Dery & Szivak |
Chaetopteryx marinkovicae | CRO, Kompanj, 14.11.2009 | CMREI0101 | JF891174 | Kučinić & Vučković |
Chaetopteryx marinkovicae | CRO, Kompanj, 14.11.2009 | CMREI0102 | JF891175 | Kučinić & Vučković |
Chaetopteryx marinkovicae | CRO, Kompanj, 14.11.2009 | CMREI0103 | JF891176 | Kučinić & Vučković |
Chaetopteryx marinkovicae | CRO, Kompanj, 14.11.2009 | CMREI0104 | JF891177 | Kučinić & Vučković |
Chaetopteryx marinkovicae | CRO, Kompanj, 14.11.2009 | CMREI0105 | JF891178 | Kučinić & Vučković |
Chaetopteryx rugulosa mecsekensis | HUN, Mecsek Mts., Nagy-Mély valley, Kánya spring, 14.11.2009 | CRMKB0101 | JF891179 | Szivak |
Chaetopteryx rugulosa mecsekensis | HUN, Mecsek Mts., Vár valley, Pásztor spring, 06.11.2009 | CRMKB0201 | JF891180 | Szivak & Uherkovich |
Chaetopteryx rugulosa mecsekensis | HUN, Mecsek Mts., Melegmányi valley, Mésztufa spring, 14.11.2009 | CRMKB0301 | JF891203 | Szivak |
Chaetopteryx rugulosa mecsekensis | HUN, Mecsek Mts., Vár valley, Iharos spring, 06.11.2009 | CRMKB0401 | JF891204 | Szivak |
Chaetopteryx rugulosa noricum | AUT, Saualpe, Klieningbach stream near Kliening, 21.11.2009 | CRNDI0101 | JF891187 | Dery & Szivak |
Chaetopteryx rugulosa noricum | AUT, Saualpe, springs of the Klippitzbach stream near Klippitztörl 21.11.2009 | CRNDI0201 | JF891188 | Dery & Szivak |
Chaetopteryx rugulosa noricum | AUT, Saualpe, springs of the Klippitzbach stream near Klippitztörl 21.11.2009 | CRNDI0202 | JF891189 | Dery & Szivak |
Chaetopteryx rugulosa noricum | AUT, Saualpe, springs of the Klippitzbach stream near Klippitztörl 21.11.2009 | CRNDI0203 | JF891219 | Dery & Szivak |
Chaetopteryx rugulosa noricum | AUT, Saualpe, springs of the Klippitzbach stream near Klippitztörl 21.11.2009 | CRNDI0204 | JF891220 | Dery & Szivak |
Chaetopteryx rugulosa noricum | AUT, Saulape, spring of the Löllingbach stream near Stranach, 21.11.2009 | CRNDI0301 | JF891190 | Dery & Szivak |
Chaetopteryx rugulosa noricum | AUT, Saulape, spring of the Löllingbach stream near Stranach, 21.11.2009 | CRNDI0302 | JF891191 | Dery & Szivak |
Chaetopteryx rugulosa noricum | AUT, Saulape, spring of the Löllingbach stream near Stranach, 21.11.2009 | CRNDI0303 | JF891217 | Dery & Szivak |
Chaetopteryx rugulosa noricum | AUT, Saulape, spring of the Löllingbach stream near Stranach, 21.11.2009 | CRNDI0304 | JF891218 | Dery & Szivak |
Chaetopteryx rugulosa rugulosa | HUN, Kőszegi Mts., Hörmann spring near Velem, 18.11.2009 | CRRDJ0101 | Szivak | |
Chaetopteryx rugulosa rugulosa | HUN, Kőszegi Mts., Hörmann spring near Velem, 18.11.2009 | CRRDJ0102 | Szivak | |
Chaetopteryx rugulosa rugulosa | AUT, Mitterneuwald, Hermann spring, 19.11.2009 | CRRDJ0201 | JF891184 | Dery & Szivak |
Chaetopteryx rugulosa rugulosa | AUT, Sommeralm, Mixnitzbach stream, 20.11.2009 | CRRDJ0301 | Dery & Szivak | |
Chaetopteryx rugulosa rugulosa | AUT, Sommeralm, Mixnitzbach stream, 20.11.2009 | CRRDJ0302 | JF891214 | Dery & Szivak |
Chaetopteryx rugulosa rugulosa | AUT, Hochegg bei Grimmenstein, spring and its outlet, 19.11.2009 | CRRDJ0401 | JF891205 | Dery & Szivak |
Chaetopteryx rugulosa rugulosa | AUT, Hochegg bei Grimmenstein, spring and its outlet, 19.11.2009 | CRRDJ0402 | JF891206 | Dery & Szivak |
Chaetopteryx rugulosa rugulosa | AUT, Hochegg bei Grimmenstein, spring and its outlet, 19.11.2009 | CRRDJ0403 | JF891207 | Dery & Szivak |
Chaetopteryx rugulosa rugulosa | AUT, Ausserneuwald, spring, 19.11.2009 | CRRDJ0501 | JF891208 | Dery & Szivak |
Chaetopteryx rugulosa rugulosa | AUT, Ausserneuwald, spring, 19.11.2009 | CRRDJ0502 | JF891209 | Dery & Szivak |
Chaetopteryx rugulosa rugulosa | AUT, Plenzengreith, upper reach of stream Schöcklbach, 20.11.2009 | CRRDJ0601 | JF891230 | Dery & Szivak |
Chaetopteryx rugulosa rugulosa | AUT, Plenzengreith, upper reach of stream Schöcklbach, 20.11.2009 | CRRDJ0602 | JF891231 | Dery & Szivak |
Chaetopteryx rugulosa rugulosa | AUT, Plenzengreith, upper reach of stream Schöcklbach, 20.11.2009 | CRRDJ0603 | JF891232 | Dery & Szivak |
Chaetopteryx rugulosa rugulosa | SLO, Pohorje Mts., Osankarica (Lukanja), 10.11.2008 | CRRDG0101 | JF891186 | Popijač |
Chaetopteryx rugulosa rugulosa | SLO, Pohorje Mts., Osankarica (Lukanja), 10.11.2008 | CRRDG0102 | JF891215 | Popijač |
Chaetopteryx rugulosa rugulosa | SLO, Pohorje Mts., Osankarica (Lukanja), 10.11.2008 | CRRDG0103 | JF891216 | Popijač |
Chaetopteryx rugulosa rugulosa | CRO, Medvednica Mts., Mrzlak spring near Sljeme, 18.11.2006 | CRREE0101 | JF891185 | Popijač |
Chaetopteryx rugulosa rugulosa | CRO, Medvednica Mts., Mrzlak spring near Sljeme, 18.11.2006 | CRREE0102 | JF891213 | Popijač |
Chaetopteryx rugulosa rugulosa | CRO, Medvednica Mts., Kraljičin Zdenac spring, Kraljičin Zdenac, 19.11.2009 | CRREE0201 | JF891210 | Kučinić & Vučković |
Chaetopteryx rugulosa rugulosa | CRO, Medvednica Mts., Bliznec stream, Podsljeme (Pilana), 09.12.2009 | CRREE0301 | JF891211 | Kučinić & Vučković |
Chaetopteryx rugulosa rugulosa | CRO, Žumberak Mts., Slapnica stream, Ribička kuća, 28.10.2009 | CRREF0101 | JF891212 | Kučinić & Vučković |
Chaetopteryx schmidi | ROU, spring brook in Cerna valley near Tatu, 13.11.2010 | CRSJF0101 | HE858259 | Ecsedi & Szivak |
Chaetopteryx schmidi | ROU, spring brook in Cerna valley near Tatu, 13.11.2010 | CRSJF0102 | HE858260 | Ecsedi & Szivak |
Chaetopteryx schmidi | ROU, spring brook in Cerna valley near Tatu, 13.11.2010 | CRSJF0103 | HE858261 | Ecsedi & Szivak |
Chaetopteryx schmidi | SRB, Derdap Mts., stream valley N of Golubinje, 13.10.2006 | CRSGE0101 | JF891182 | Danyi, Kontschan & Muranyi |
Chaetopteryx schmidi | SRB, Derdap Mts., stream valley N of Golubinje, 13.10.2006 | CRSGE0102 | JF891201 | Danyi, Kontschan & Muranyi |
Chaetopteryx schmidi | SRB, Derdap Mts., Grgeci spring, Donji Milankovac, 13.10.2006 | CRSGE0201 | JF891183 | Danyi, Kontschan & Muranyi |
Chaetopteryx schmidi | SRB, Derdap Mts., Grgeci spring, Donji Milankovac, 13.10.2006 | CRSGE0203 | JF891202 | Danyi, Kontschan & Muranyi |
Chaetopteryx bucari sp. n. | CRO, Kriz spring near Petrinja, 08.12.2009 | CxxEC0101 | JF891192 | Kučinić, Delić & Bučar |
Chaetopteryx bucari sp. n. | CRO, Kriz spring near Petrinja, 07.11.2009 | CxxEC0102 | JF891222 | Kučinić, Delić & Bučar |
Chaetopteryx bucari sp. n. | CRO, Kriz spring near Petrinja, 07.11.2009 | CxxEC0103 | JF891223 | Kučinić, Delić & Bučar |
Chaetopteryx bucari sp. n. | CRO, Kriz spring near Petrinja, 04.11.2009 | CxxEC0104 | JF891224 | Bučar |
Chaetopteryx bucari sp. n. | CRO, Kriz spring near Petrinja, 08.12.2009 | CxxEC0105 | JF891225 | Kučinić, Delić, Bučar & Vučković |
Chaetopteryx bucari sp. n. | CRO, Hrvatski Cuntic, Marića točak spring, 22.11.2009 | CxxEC0201 | JF891193 | Kučinić, Delić & Bučar |
Chaetopteryx bucari sp. n. | CRO, Hrvatski Cuntic, Marića točak spring, 21.11.2009 | CxxEC0202 | JF891221 | Kučinić, Delić & Bučar |
Chaetopteryx bucari sp. n. | CRO, Hrvatska Kostajnica, Varoški bunar spring, 06.12.2009 | CxxEC0301 | Kučinić, Delić & Bučar | |
Chaetopteryx bucari sp. n. | CRO, Šuplji Kamen, Slabinja spring, 29.11.2009 | CxxEC0401 | JF891194 | Kučinić, Delić & Bučar |
Chaetopteryx bucari sp. n. | CRO, Banovina region, Pecki spring, 15.12.2009 | CxxEC0501 | JF891195 | Kučinić, Delić & Bučar |
Chaetopteryx bucari sp. n. | CRO, Banovina region, Pecki spring, 21.11.2009 | CxxEC0502 | JF891228 | Kučinić, Delić & Bučar |
Chaetopteryx bucari sp. n. | CRO, Banovina region, Pecki spring, 21.11.2009 | CxxEC0503 | JF891229 | Kučinić, Delić & Bučar |
Chaetopteryx bucari sp. n. | CRO, Banovina region, Gora spring, 10.12.2009 | CxxEC0601 | JF891226 | Bučar |
Chaetopteryx bucari sp. n. | CRO, Mečenčani, Pašino vrelo, 29.11.2009 | CxxEC0701 | JF891227 | Kučinić, Delić & Bučar |
Chaetopterygopsis maclachlani | AUT, Lower Austria, Rohrwiesteich, 20.10.2004 | 08HMCAD-331* | HMTRI331-09* | Malicky |
Chaetopteryx fusca | AUT, Lower Austria, Rohrwiesteich, 20.10.2004 | 08HMCAD-333* | HMTRI333-09* | Malicky |
Chaetopteryx gessneri | ITA, Umbria, Perugia, Fium Nera above Visso, 11.12.2005 | 07HMCAD-0177* | HMCAD177-08* | Malicky |
Chaetopteryx moretti | ITA, Belluno, Val Canzoi, Veneto, 31.10.2003 | HM09Cm7* | HMTRI421-09* | Malicky |
Chaetopteryx villosa | AUT, Lower Austria, Sarleinsbach, 27.06.2005 | 07HMCAD-0134* | HMCAD134-08* | Malicky |
Drusus alpinus | IT, Valprato Soana, Ronchietto, 10.07.2004 | HM09Dalp8* | HMTRI456-09* | Delmaistro |
Drusus discolor | SK, Lower Tatra, Stream above Partizanska L'upča, 09.06.2008 | ESCAD909-17* | KKCAD497-09* | Bonada |
Drusus rectus | ES, Camprodon/Setcases Alta Val de Ter, 27.07.2004 | HM09Drec8* | HMTRI423-09* | Aistleitner |
Metanoea rhaetica | AUT, Carinthia, Valentinbach, Plockenstrasse, 08.07.2007 | 08HMCAD-020* | HMTRI020-08* | Malicky |
Limnephilus centralis | NORWAY | NHRS:FI9 | FN601020 | Malm & Johanson 2011 |
The mitochondrial COI barcodes were generated at the Canadian Centre for DNA Barcoding, University of Guelph, Canada. Standard barcoding protocols for DNA extraction (
Phylogenetic analysis. Sequences were edited manually and aligned using the program Geneious 5.4 (
Microphotography and measuring. Microphotographic images of genitalia and forewing measurements were taken using a Leica Wild MZ8 stereomicroscope and Olympus SP-500 UZ digital camera. The photographs were processed with the Olympus Quick Photo Camera 2.2. software package. Geographic coordinates and altitudes of sampling localities were recorded with a Garmin ‘Oregon 450' GPS device.
Phylogenetic analyses. In the Bayesian phylogenetic tree based on mtCOI sequences the Chaetopteryx rugulosa group species clustered into 4 strongly supported clades (Figure 2). Chaetopteryx marinkovicae was basal within the species group. The remaining species fell into 3 clades: a basal clade with Chaetopteryx rugulosa schmidi, Chaetopteryx bucari sp. n., and 2 derived sister clades comprising Chaetopteryx clara, Chaetopteryx goricensis, Chaetopteryx irenae, and Chaetopteryx rugulosa rugulosa, Chaetopteryx rugulosa noricum, Chaetopteryx rugulosa mecsekensis. Chaetopteryx bucari sp. n. is sister to the highly supported Chaetopteryx rugulosa schmidi. The mean value of the uncorrected pairwise distance (p distance) was 2.02% between them (Table 3). The p distance did not reach 1% within the 2 clades (Chaetopteryx bucari sp. n.: 0.17%; Chaetopteryx rugulosa schmidi: 0.75%). The relationship of the nominal species of the group Chaetopteryx rugulosa rugulosa and Chaetopteryx rugulosa noricum was not resolved, as the 4 subclades formed a polytomy. In the phylogenetic tree Chaetopteryx rugulosa schmidi was clearly separated from the clade containing the subspecies of Chaetopteryx rugulosa (Figure 2). The mean values of p distance between the 3 subspecies of Chaetopteryx rugulosa ranged between 1.61–3.02 %, while the mean values between the Chaetopteryx rugulosa schmidi and the other subspecies of Chaetopteryx rugulosa were distinctly higher (4.66 – 5.85%) (Table 3).
Bayesian tree for members of the Chaetopteryx rugulosa species group based on mitochondrial COI sequence. Black circles on nodes mark Bayesian posterior probabilities pp>0.95.
Estimates of evolutionary divergence over sequence pairs within and between phylogenetically defined species and subspecies based on mtCOI sequence data. Distance matrix is shown with the mean ± SD values of the intraspecific and interspecific pairwise genetic distances for the all Chaetopterygini species included in the analysis. Abbrev.: CRR – Chaetopteryx rugulosa rugulosa, CRN – Chaetopteryx rugulosa noricum, CRM – Chaetopteryx rugulosa mecsekensis, CCX – Chaetopteryx clara, CGR – Chaetopteryx goricensis, CIX – Chaetopteryx irenae, CBU – Chaetopteryx bucari sp.n., CRS – Chaetopteryx schmidi, CMR – Chaetopteryx marinkovicae, CBA – Chaetopteryx bosniaca, CMO – Chaetopteryx morettii, CFU – Chaetopteryx fusca, CVI – C. villosa, CGE – Chaetopteryx gessneri, CAX – Chaetopteryx aproka, CMA – Chaetopterygopsis maclachlani, CMJ – Chaetopteryx major.
CRR | CRN | CRM | CCX | CGR | CIX | CBU | CRS | CMR | CBO | CMO | CFU | CVI | CGE | CAX | CMA | CMJ | |
CRR | 1.05±0.97 | 1.61±0.49 | 3.02±0.17 | 4.87±0.24 | 4.55±0.24 | 4.79±0.27 | 4.63±0.30 | 5.44±0.39 | 9.24±0.46 | 12.64±0.15 | 12.85±0.33 | 12.69±0.20 | 12.69±0.20 | 14.03±0.20 | 12.00±0.39 | 12.32±0.20 | 14.17±0.16 |
CRN | 1.20±0.89 | 2.83±0.31 | 5.06±0.23 | 4.74±0.24 | 5.26±0.20 | 5.17±0.18 | 5.85±0.11 | 9.45±0.27 | 12.85±0.08 | 12.66±0.12 | 12.83±0.12 | 12.83±0.12 | 14.06±0.12 | 12.18±0.14 | 11.98±0.24 | 14.14±0.39 | |
CRM | 0.17±0.11 | 4.79±0.09 | 4.47±0.10 | 4.69±0.12 | 4.38±0.11 | 4.66±0.13 | 8.87±0.07 | 11.89±0.07 | 11.97±0.08 | 11.76±0.14 | 11.76±0.14 | 12.44±0.14 | 11.60±0.12 | 11.47±0.08 | 13.78±0.00 | ||
CCx | 0.00±0.00 | 0.37±0.07 | 3.80±0.08 | 5.86±0.14 | 5.69±0.06 | 9.92±0.00 | 12.61±0.00 | 12.61±0.00 | 12.77±0.00 | 12.77±0.00 | 13.61±0.00 | 12.77±0.00 | 12.61±0.00 | 14.49±0.07 | |||
CGR | 0.03±0.07 | 3.48±0.10 | 5.54±0.15 | 5.37±0.08 | 9.93±0.05 | 12.62±0.05 | 12.62±0.05 | 12.79±0.05 | 12.79±0.05 | 13.63±0.05 | 12.78±0.05 | 12.62±0.05 | 14.17±0.08 | ||||
CIX | 0.10±0.09 | 4.41±0.16 | 5.11±0.18 | 10.15±0.08 | 12.67±0.08 | 13.21±0.09 | 12.54±0.09 | 12.54±0.09 | 13.04±0.09 | 13.38±0.09 | 13.55±0.09 | 15.39±0.11 | |||||
CBa | 0.17±0.13 | 2.02±0.20 | 8.79±0.13 | 12.48±0.13 | 12.65±0.14 | 12.48±0.14 | 12.48±0.14 | 12.65±0.14 | 11.21±0.08 | 12.89±0.08 | 14.00±0.16 | ||||||
CRS | 0.75±0.57 | 8.72±0.06 | 12.24±0.06 | 13.04±0.13 | 12.05±0.21 | 12.05±0.21 | 12.67±0.23 | 11.56±0.22 | 13.93±0.06 | 14.44±0.15 | |||||||
CMR | 0.00±0.00 | 11.60±0.00 | 11.60±0.00 | 11.93±0.00 | 11.93±0.00 | 13.11±0.00 | 11.76±0.00 | 12.94±0.00 | 12.77±0.18 | ||||||||
CBO | 0.00±0.00 | 2.69±0.00 | 4.87±0.00 | 4.87±0.00 | 6.05±0.00 | 12.61±0.00 | 12.61±0.00 | 15.26±0.17 | |||||||||
CMO | 0.00±0.00 | 5.55±0.00 | 5.55±0.00 | 6.55±0.00 | 12.94±0.00 | 11.76±0.00 | 14.82±0.08 | ||||||||||
CFU | 0.00±0.00 | 0.00±0.00 | 5.88±0.00 | 13.05±0.19 | 12.94±0.00 | 15.83±0.08 | |||||||||||
CVI | 0.00±0.00 | 5.88±0.00 | 13.05±0.19 | 12.94±0.00 | 15.83±0.08 | ||||||||||||
CGE | 0.00±0.00 | 13.45±0.00 | 13.44±0.00 | 15.87±0.09 | |||||||||||||
CAX | 0.35±0.00 | 11.71±0.19 | 13.44±0.11 | ||||||||||||||
CMA | 0.00±0.00 | 13.04±0.15 | |||||||||||||||
CMJ | 0.47±0.51 |
http://zoobank.org/E775EC69-0E8A-4AF0-A027-F290BB31E76E
http://species-id.net/wiki/Chaetopteryx_bucari
Figures 3–16Holotype male: CROATIA, Pecki spring, 45°23'50"N, 16°14'40"E, 161 m a.s.l., 15 December 2009, leg. Bučar, Delić, Kučinić, dry specimen, DNA Barcode ID: HGCAD046-10, deposited in the Croatian Natural History Museum in Zagreb.
Paratype: CROATIA, ♂ and ♀ (n=49): 1 female, Pecki spring, 21 November 2009, leg. Bučar, Delić, Kučinić, dry specimen, DNA Barcode ID: HGCAD087-10; 14 males, Pecki spring, 31 October 2011; 9 females, Pecki spring, 31 October 2010; 20 females, Pecki spring, 30 November 2011; 2 males and 2 females, Hrvatski Čuntić stream, 45°21'28"N, 16°17'04"E, 159 m a.s.l., 22 October 2010; 1 male, Marića točak, 45°21'29"N, 16°17'03"E, 163 m a.s.l., 23 November 2012, leg. Bučar, Delić, Kučinić (all specimens in alcohol).
Male of Chaetopteryx bucari is most similar to Chaetopteryx rugulosa mecsekensis and Chaetopteryx rugulosa schmidi but differs in the following features: 1. In lateral view the inferior appendages in Chaetopteryx bucari are always with a pointed apex on the dorsal side, not rounded as in Chaetopteryx rugulosa mecsekensis; 2. Bristles in Chaetopteryx bucari are set more distally from the membranous part of the aedeagus than in Chaetopteryx rugulosa mecsekensis and Chaetopteryx rugulosa schmidi and never reach (touch) the lateral membranous finger, as in Chaetopteryx rugulosa mecsekensis. Female of Chaetopteryx bucari is clearly different from other species in the Chaetopteryx rugulosa group (e.g., form of the visible finger on lateral side, form of the anal tube, form of the supragenital plate of segment X in lateral and ventral views, form of the median lobe of the vulvar scale in ventral view). We did not find strong morphological variability among the females of the new species (except the median lobe of the vulvar scale). Females of Chaetopteryx bucari have in lateral, ventral and dorsal views very visible finger-shaped proturbances (ventral lobes of tergite IX) on the anal tube which is lacking in Chaetopteryx rugulosa mecsekensis and Chaetopteryx rugulosa schmidi. In lateral view the excision of the anal tube in Chaetopteryx rugulosa rugulosa is more pronounced than in Chaetopteryx bucari. The median lobe of the vulvar scale in Chaetopteryx rugulosa mecsekensis, Chaetopteryx rugulosa rugulosa and Chaetopteryx rugulosa schmidi is longer and more visible than in Chaetopteryx bucari.
Wings and legs yellow to yellowish-brown; veins darker in both sexes (Figure 3). Antennae long, grey to fuscous. Scapus yellow to yellowish-brown, thorax and abdomen yellow. Spur formula male 0, 3, 3, female 1, 3, 3. Ocelli present. Forewing with round apex; length 7.7–9.9 mm in males, 7.2–10.1 mm in females.
Chaetopteryx bucari sp. n., adults at type locality, Pecki spring, Croatia.
Male genitalia (Figures 4–11). In dorsal view, spinulose zone of tergite VIII well developed with yellow setae. Segment IX ventrally broad, dorsally narrow in lateral view (Figures 4–5). Superior appendages with small yellow setae, shape of superior appendages variable (Figures 4–7b–d), usually in one of two forms (Figures 4–6). In lateral view, 1st form with posterior edge slightly rounded apically, concave at middle (Figure 5); in 2nd form, dorsal side more protuberant with round or irregular apex (Figures 4, 7b). In some specimens triangular or rectangular intermediate forms are found (Figure 7c-d). Inferior appendages in lateral view rectangular, anterior part broad, posterior part narrow (Figures 4–7a). Apical flap of inferior appendage developed, in lateral view with pointed apex (tip) and ventral side slightly rounded; or with apex forked, long setae present on ventral side (Figures 4–7a). Intermediate appendages (paraproctal complex) elongated in lateral view with long, connecting middle section, apical hook narrowing with upward–curving apex (Figures 4–5), basal triangular part of paraproct relatively large in caudal view (Figures 8–9). Phallic organ (phallus) a single tube consisting of phallic apodeme, phallobase, aedeagus and parameres. Aedeagus relatively long, sclerotized, in posterior part with membranous lobes, lateral lobes membranous finger-like proturbances (endophallus) (Figures 10a–d). Two relatively short parameres set very distant from posterior membranous part of aedeagus (Figures 10a–b, 10d); parameres with sclerotized, straight, stout, brown bristles (Figures 10a–b, 10d, 11a–f). Bristles vary in width and length (Figure 11a–f); lateral bristles shorter; bristles arranged in 1 fan-like row (Figure 11a–f); in specimens with more bristles, some form 2nd row; bristles vary from 5-10.
Chaetopteryx bucari sp. n., male genitalia, lateral view.
Chaetopteryx bucari sp. n., male genitalia, lateral view.
Chaetopteryx bucari sp. n., male genitalia, lateral view.
Chaetopteryx bucari sp. n., male genitalia, lateral view a inferior appendages b–d superior appendages.
Chaetopteryx bucari sp. n., male genitalia, caudal view.
Chaetopteryx bucari sp. n., male genitalia, intermediate appendages (paraproctal complex), caudal view.
Chaetopteryx bucari sp. n., male genitalia, phallic organ (phallus): a dorsal view b ventral view c posterior membranous part of aedeagus d lateral view.
Chaetopteryx bucari sp. n., male genitalia a-f parameres with sclerotized bristles.
Female genitalia (Figures 12–16). Anal tube (fusion of tergites IX and X) in lateral view broad, relatively elongated with one excision and very distinct finger-shaped proturbance (lobes of tergite IX) on ventral side (Figures 12–13). Apex of proturbance rounded or slightly pointed with small yellow setae (Figures 12–15). In 2/3rds of specimens examined ventral and dorsal lips of anal tube equal in length, in 1/3rd ventral lip longer. In dorsal view anal tube thickened with digitate proturbance on lateral side and small excision (recess) in middle (Figure 14). In ventral view anal tube broad with larger excision (recess) in middle than in dorsal side (Figure 15). Supragenital plate of segment X well-developed, triangular in shape in lateral and ventral views (Figures 12, 15). Lateral segment of vulvar scale relatively short in ventral view, with flat or slightly rounded apex (Figure 16a–c). Median lobe of vulvar scale (lower vulvar lip) with very small rounded or pointed apex (Figure 16b–d). In ca. 1/3rd of specimens' median lobe of vulvar scale not visible (Figure 16a).
Chaetopteryx bucari sp. n., female genitalia, lateral view.
Chaetopteryx bucari sp. n., female genitalia, dorso-lateral view.
Chaetopteryx bucari sp. n., female genitalia, dorsal view.
Chaetopteryx bucari sp. n., female genitalia, ventral view.
Chaetopteryx bucari sp. n., female genitaliaa–d vulvar scale and median lobe of vulvar scale, ventral view.
The species is dedicated to Professor Matija Bučar from the Faculty of Education, Department in Petrinja, University of Zagreb.
During our recent faunal surveys in Croatia and the Western Dinaric Balkan Chaetopteryx bucari was found only at 8 localities in the Banovina region (Table 1). The most distant sampling sites are 40 km apart (Slabinja and Gore). We collected Chaetopteryx bucari from 2 springs, 5 wellsprings and 1 location in the stream (Table 1). In total, we collected more than 580 specimens of Chaetopteryx bucari (85% were collected in pyramid-type emergence traps). The most abundant populations were found at Pecki spring and a headwater stream in Hrvatski Ćuntić. Over 150 specimens of Chaetopteryx bucari were observed on the night of October 14, 2010 on the walls of a small building next to the stream in Hrvatski Čuntić. In Pecki spring more than 50 specimens were observed on the night of October 31, 2010. Chaetopteryx bucari was recorded at low altitudes between 104–185 m a.s.l. (Table 1).
Chaetopteryx bucari was collected in pyramid-type emergence traps from the end of September-December. The highest number of specimens was collected in October and November in both years. The sex ratio in both years was biased toward males, 1:1.37 (♀♀: ♂♂) in 2010, and 1:1.40 (♀♀:♂♂) in 2011. Besides Chaetopteryx bucari, Chaetopteryx gonospina Marinković-Gospodnetić, 1966 and 2 additional caddisfly species (Limnephilus rhombicus (Linnaeus, 1758), Potamophylax pallidus Klapálek, 1898) were recorded in the emergence traps.
In addition to Chaetopteryx bucari 2 other species of the Chaetopteryx rugulosa group were collected in Croatia during our recent surveys. Chaetopteryx marinkovicae was collected from its type locality on the stream and spring in Kompanj village (Istria region); Chaetopteryx rugulosa rugulosa was caught on Mt. Žumberak and Mt. Medvednica (northeast and central Croatia). Other species of Chaetopteryx found during this investigation were Chaetopteryx bosniaca Marinković-Gospodnetić, 1959 (Lika region), Chaetopteryx gonospina Marinković-Gospodnetić, 1966 (Banovina region), Chaetopteryx fusca (central Croatia, Dalmatia and Lika regions), and Chaetopteryx major (central Croatia).
Systematic and taxonomic implications. Based on molecular evidence, we could confirm the hypothesis that Chaetopteryx bucari is a distinct species. Although Chaetopteryx bucari does not have a pp>0.95, it represents the sister taxon (pp>0.95) to the highly supported Chaetopteryx rugulosa schmidi. Furthermore, the mean genetic distance (2.02%) between Chaetopteryx bucari and Chaetopteryx rugulosa schmidi barely reached the 2-3% divergence observed as an interspecific genetic divergence in mtCOI sequences among some well-defined caddisfly species (
In both sexes, especially in the adult female, Chaetopteryx bucari is relatively easily distinguishable from other taxa of the Chaetopteryx rugulosa group. The genetic data also show that specimens from 7 populations across the known range of the species from a clearly distinct clade from all other analysed Chaetopteryx. It is interesting that the female of Chaetopteryx bucari is particularly informative in diagnosing the species. In caddisflies this is quite unusual as males are generally more easily distinguished and females are often very difficult to differentiate from one another.
Based on the phylogenetic position of Chaetopteryx rugulosa schmidi in relation to Chaetopteryx rugulosa rugulosa and the other Chaetopteryx rugulosa subspecies, Chaetopteryx rugulosa schmidi is well-defined and quite divergent from other members of the Chaetopteryx rugulosa clade based on molecular data. Thus, the subspecies Chaetopteryx rugulosa schmidi is here re-established as a distinct species, Chaetopteryx schmidi, as it was described originally by
Ecology. The emergence pattern of Chaetopteryx bucari corresponds with the general autumnal emergence patterns of the genus, usually from September-December, though emergence can be prolonged through January for some Chaetopteryx species (
Research on the diversity of large karst springs on the Balkan Peninsula has revealed high levels of caddisfly diversity. In some cases more than 20 species were collected from a single spring (
Distribution of Chaetopteryx rugulosa group in Croatia. At present, the genus Chaetopteryx is represented by 9 taxa in Croatia (
Distribution of Chaetopteryx rugulosa group in Croatia.
Until now, the new species Chaetopteryx bucari was found only in the Banovina region, which is situated between rivers Sava and Kupa to the north and the state border with Bosnia and Herzegovina to the south and east (Figure 17). The Banovina region is characterised by rolling hills up to 600 m a.s.l. There are many small springs and streams in the region, and 3 large rivers, Una, Kupa and Sava, that form the border of the region. It is possible that Chaetopteryx bucari is also distributed in some other parts of continental Croatia or in Bosnia and Herzegovina, because we found this species in the valley of the Una River (Slabinja spring, Varoški bunar spring), which forms the border between these 2 countries.
According to the current findings, Chaetopteryx bucari is not rare in the Croatian fauna. In fact, it is one of the most dominant caddisflies in the Banovina region. Along with Chaetopteryx fusca (
Taxa from the Chaetopteryx rugulosa group have allopatric distributions in Croatia (Figure 17): Chaetopteryx bucari is distributed in the Banovina region, Chaetopteryx rugulosa rugulosa in northern Croatia on Mt. Medvednica and Mt. Žumberak, Chaetopteryx rugulosa mecsekensis in eastern Croatia on Mt. Papuk and Chaetopteryx marinkovicae in the sub-Mediterranean part of Croatia in Istria (
Many members of the genus Chaetopteryx are either small-scale endemics or species with a low number of disjunct populations. This makes the group very interesting for biogeographic studies. There are several reasons that could explain the observed pattern of distribution: small populations, poor mobility of the winter emerging adults, and distribution in springs and in headwater reaches of small streams. Besides naturally isolating individual populations from one another, these aspects can also cause difficulties for investigating the genus, as it is hard to access many of the sites, especially in winter. Future investigations of this genus will be focused on poorly researched areas in Croatia and the western Balkans to gain a better understanding of the distribution and biogeography of Chaetopteryx in the region.
We express our gratitude to Xin Zhou for handling the mitochondrial barcoding of our specimens within the Barcoding of Life initiative. We thank Karl Kjer for sharing the previously unpublished “outgroup” data from BOLD. We thank two anonymous reviewers and Ralph Holzenthal for their valuable comments that helped improve the manuscript. For technical help we acknowledge senior curator Iva Mihoci, PhD, from the Croatian Natural Museum in Zagreb, Nediljko Landeka form the Institute of Public Health in Pula, and Milivoj Franjević from the Foresty Faculty, University of Zagreb. ISz was partially funded through an incoming research grant of the Biodiversity and Climate Research Centre in the frame of the research funding program Landes-Offensive zur Entwicklung Wissenschaftlich-ökonomischer Exzellenz (LOEWE) of Hesse's Ministry of Higher Education, Research, and the Arts, Germany. SUP and MB acknowledge financial support from the research fundprogram “LOEWE – Landes-Offensive zur Entwicklung Wissenschaftlich-ökonomischer Exzellenz” of Hesse's Ministry of Higher Education, Research, and the Arts. This research was supported by the Austrian Science Fund (FWF) as a part of Project P 23687-B17 and Croatian Ministry of Science, Education and Sports as a part of Project No. 119–1193080–1206.